Affinage

ENY2

Transcription and mRNA export factor ENY2 · UniProt Q9NPA8

Round 2 corrected
Length
101 aa
Mass
11.5 kDa
Annotated
2026-04-28
83 papers in source corpus 25 papers cited in narrative 24 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

ENY2 is a conserved adaptor protein that integrates transcription, histone modification, and mRNA export by serving as a shared non-enzymatic subunit of the SAGA/TFTC deubiquitination module (DUBm), the TREX-2 mRNA export complex, and the THO mRNP biogenesis complex (PMID:14718168, PMID:20048002). Within the DUBm, ENY2 and ATXN7L3 scaffold the H2B deubiquitinases USP22, USP27X, and USP51, and their depletion globally elevates H2Bub1 levels; in the TREX-2 complex, ENY2 adopts a helical hairpin fold that wraps around alpha-helical segments of Sac3/GANP, and its binding to Sgf11 (SAGA) versus Sac3/GANP (TREX-2) is mutually exclusive, establishing physically distinct subcomplexes (PMID:19328066, PMID:20007317, PMID:27132940). ENY2 promotes mRNA export by anchoring TREX-2 to the nuclear pore basket via NUP153 and TPR, and loss of this function causes nuclear poly(A)+ RNA accumulation, genome instability through R-loop formation, and shortened replicative lifespan (PMID:18667528, PMID:23591820, PMID:35771153). In Drosophila, ENY2 additionally confers chromatin barrier activity at Su(Hw)- and dCTCF-dependent insulators by preventing the spread of H3K27me3 and Polycomb silencing (PMID:17643381, PMID:25147918).

Mechanistic history

Synthesis pass · year-by-year structured walk · 13 steps
  1. 2004 High

    The discovery that yeast Sus1 co-purifies with both SAGA and the Sac3-Thp1 mRNA export complex established that a single small protein physically links transcription initiation at gene promoters to mRNA export at nuclear pores, revealing an unexpected coupling mechanism.

    Evidence Biochemical co-purification, ChIP at GAL1, and mRNA export assays in S. cerevisiae

    PMID:14718168

    Open questions at the time
    • Structural basis of Sus1 association with both complexes unknown
    • Unclear whether simultaneous or mutually exclusive binding
  2. 2006 High

    Identification of Sus1 as a stable subunit of the Ubp8-Sgf11 deubiquitinase module within SAGA resolved how ENY2 contributes to transcription: it is required for H2B deubiquitination and consequent regulation of H3 methylation, placing it directly in the histone modification cascade.

    Evidence Genetic deletions, high-salt co-purification of DUBm subcomplex, and reciprocal ChIP recruitment in yeast

    PMID:16855026

    Open questions at the time
    • No in vitro reconstitution of DUBm activity with Sus1
    • Relative contribution of Sus1 to DUB catalysis versus substrate access unclear
  3. 2007 High

    Demonstrating that Drosophila ENY2 is recruited by Su(Hw) to chromatin insulators and is specifically required for barrier activity (not enhancer-blocking) revealed a metazoan-specific function in preventing heterochromatin spreading.

    Evidence In vivo insulator recruitment assay, genetic interaction analysis in Drosophila

    PMID:17643381

    Open questions at the time
    • Mechanism by which ENY2 blocks heterochromatin spreading not defined
    • Whether this function exists in mammals unknown
  4. 2008 High

    Three concurrent advances established ENY2's mechanistic breadth: TREX-2 was shown to prevent R-loops and maintain genome stability independently of SAGA; the mammalian DUBm containing ENY2, ATXN7L3, and USP22 was biochemically reconstituted with H2A/H2B deubiquitinase activity; and Sus1 was found on coding regions with elongating RNA Pol II and mRNA export factors.

    Evidence R-loop assays and genetic epistasis in yeast; DUB activity assay and PEV genetics in human/Drosophila; ChIP across coding regions and co-purification with Yra1/Mex67

    PMID:18206972 PMID:18667528 PMID:18923079

    Open questions at the time
    • How Sus1 transitions from promoter-bound SAGA to elongation-associated TREX-2 unknown
    • R-loop suppression mechanism not fully resolved
  5. 2009 High

    Crystal structures of Sus1:Sac3 and Sus1:Sgf11 complexes revealed that ENY2 uses the same helical hairpin surface to grip alpha-helical segments in both partners, establishing mutually exclusive binding and proving that separate pools of ENY2 exist in SAGA versus TREX-2.

    Evidence X-ray crystallography of Sus1:Cdc31:Sac3(CID) and Sus1:Sgf11, validated by site-directed mutagenesis and in vivo NPC localization/mRNA export assays

    PMID:19269973 PMID:19328066 PMID:20007317

    Open questions at the time
    • Full-length TREX-2 or SAGA DUBm structures not available
    • Dynamics of ENY2 exchange between complexes in vivo unresolved
  6. 2010 High

    Discovery that Drosophila ENY2 stably associates with the THO complex—independent of SAGA and TREX-2—and is required for THO recruitment to nascent mRNA and 3′-end processing expanded the functional repertoire to a third mRNP biogenesis complex.

    Evidence Co-purification, ChIP at hsp70, RNAi knockdown with 3′-end processing analysis in Drosophila

    PMID:20048002

    Open questions at the time
    • ENY2-THO structural interface not characterized
    • Whether mammalian ENY2 participates in THO not tested
  7. 2012 High

    Structural and functional characterization of the mammalian TREX-2 complex (GANP-ENY2-PCID2-centrins) showed that the GANP:ENY2 interface differs from the yeast Sac3:Sus1 interface, and ENY2 depletion impairs mRNA export, confirming evolutionary conservation of the core export function.

    Evidence X-ray crystallography of GANP:ENY2, siRNA knockdown, mRNA export and NPC localization assays in human cells

    PMID:22307388

    Open questions at the time
    • Whether ENY2 contributes to TREX-2 substrate selectivity in mammals unknown
  8. 2013 High

    Mapping the NPC docking requirements of human TREX-2 to nucleoporins NUP153 and TPR, with ENY2 dispensable for nuclear protein export, delineated ENY2's role as specific to the mRNA export arm of TREX-2 at the pore basket.

    Evidence siRNA knockdowns of NUP153/TPR with TREX-2 localization and functional assays

    PMID:23591820 PMID:24291146

    Open questions at the time
    • Direct contacts between ENY2 and nucleoporins not structurally resolved
  9. 2014 High

    Extending the insulator function to dCTCF-dependent boundaries and showing that ENY2 loss causes spreading of H3K27me3 and Polycomb occupancy mechanistically linked ENY2's barrier activity to antagonism of Polycomb-mediated silencing.

    Evidence Transgenic insulator assay, RNAi in Drosophila BG3 cells, ChIP for H3K27me3 and Pc

    PMID:25147918

    Open questions at the time
    • Whether ENY2 recruits a chromatin remodeler or acts passively at barriers unclear
    • Mammalian insulator role not tested
  10. 2016 High

    Identification of USP27X and USP51 as additional H2B deubiquitinases that, like USP22, require ENY2 and ATXN7L3 as shared adaptors explained why ENY2 depletion causes global H2Bub1 increase whereas USP22 depletion alone does not, establishing ENY2 as a central node for multiple DUB complexes.

    Evidence Mass spectrometry, systematic siRNA depletion with H2Bub1 immunoblot readout, tumor growth assays

    PMID:27132940

    Open questions at the time
    • How cells regulate partitioning of ENY2/ATXN7L3 among three DUBs unknown
    • Substrate specificity differences among USP22/USP27X/USP51 not defined
  11. 2018 Medium

    Genome-wide ChIP-seq of Sus1 revealed broad occupancy at upstream regulatory regions, dynamic redistribution to stress-responsive genes upon heat shock, and contributions to both RNA synthesis and mRNA stability, indicating ENY2 acts as a general transcriptional cofactor beyond SAGA-regulated genes.

    Evidence ChIP-seq and genomic run-on in S. cerevisiae

    PMID:29598828

    Open questions at the time
    • Mechanism of mRNA stability regulation by ENY2 unresolved
    • Single lab observation awaiting independent replication
  12. 2022 Medium

    Genetic suppression and lifespan assays demonstrated that ENY2's role in replicative longevity operates specifically through TREX-2-mediated mRNA export rather than SAGA-dependent deubiquitination, functionally separating the two pathways at the organismal level.

    Evidence Overexpression rescue by Mex67/Dbp5, lifespan assay, poly(A)+ FISH in yeast sus1Δ

    PMID:35771153

    Open questions at the time
    • Molecular basis of the mRNA export–lifespan connection unclear
    • Not tested in metazoan aging models
  13. 2026 Medium

    Two recent studies broadened ENY2's functional scope: depletion in human cells releases NPM1 from nucleoli, triggering nucleolar stress and p53 stabilization that suppresses tumor growth; in Drosophila, ENY2's presence at the histone locus body as part of SAGA, THO, and TREX-2 links it to histone gene transcription and histone mRNA export.

    Evidence In vivo tumor models, siRNA with nucleolar stress readouts, co-IP with FLASH, ChIP at histone locus in Drosophila polytene chromosomes

    PMID:41642454 PMID:41912850

    Open questions at the time
    • Nucleolar stress mechanism not independently replicated
    • Whether NPM1 release is direct or indirect effect of ENY2 depletion unknown
    • Histone mRNA export role awaits reconstitution

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key unresolved questions include how cells regulate partitioning of ENY2 among its multiple complexes (SAGA DUBm, TREX-2, THO, USP27X/USP51 modules), whether the chromatin insulator barrier function is conserved in mammals, and the structural basis of ENY2's emerging nucleolar and ribosome export roles.
  • No in vivo single-molecule tracking of ENY2 complex dynamics
  • Mammalian insulator role untested
  • Nucleolar function requires independent validation

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 6 GO:0140110 transcription regulator activity 4 GO:0042393 histone binding 3
Localization
GO:0005634 nucleus 5 GO:0005694 chromosome 4 GO:0005635 nuclear envelope 3 GO:0005654 nucleoplasm 2
Pathway
R-HSA-8953854 Metabolism of RNA 5 R-HSA-4839726 Chromatin organization 4 R-HSA-74160 Gene expression (Transcription) 4
Partners
ATXN7L3GANPNUP153PCID2SGF11TPRUSP22USP27X
Complex memberships
SAGA DUBmTHOTREX-2

Evidence

Reading pass · 24 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2004 Sus1 (ENY2 ortholog in yeast) is a component of both the SAGA histone acetyltransferase complex (involved in transcription initiation) and the Sac3-Thp1 mRNA export complex at the nuclear pore basket, physically linking transcription and mRNA export; ChIP showed Sus1 associates with the GAL1 promoter during activation, and sus1 mutants show mRNA export defects. Biochemical co-purification, chromatin immunoprecipitation (ChIP), DNA macroarray, genetic analysis Cell High 14718168
2006 Sus1 (ENY2) is required for SAGA-dependent histone H2B deubiquitination and normal H3 methylation levels; it forms a stable subcomplex with Ubp8 and Sgf11 within SAGA, and in vivo recruitment of Sus1 to the GAL1 promoter depends on Ubp8 and vice versa. Genetic deletion analyses, co-purification under high salt, ChIP, in vivo recruitment assays Molecular biology of the cell High 16855026
2007 Drosophila E(y)2/ENY2 is recruited to Su(Hw) insulators via the zinc-finger domain of Su(Hw) and is required specifically for the barrier (anti-spreading) activity of these insulators but not their enhancer-blocking activity; genetic interaction between su(Hw) and e(y)2 is lethal. In vivo recruitment assays, genetic interaction analysis, partial loss-of-function allele (e(y)2^u1) Molecular cell High 17643381
2008 Thp1-Sac3-Sus1-Cdc31 (TREX-2) forms a functional unit with a role in mRNP biogenesis and genome integrity that is independent of SAGA; R-loop formation is implicated in the transcription/genome instability phenotypes of TREX-2 mutants, similar to THO/TREX mutants, defining a pathway connecting transcription elongation with mRNA export. Genetic epistasis, R-loop analysis (ribozyme constructs, RNase H), AID-induced instability assays Molecular biology of the cell High 18667528
2008 ENY2/Sus1 (TFTC/STAGA complex) is part of a histone H2A and H2B deubiquitinase module together with ATXN7L3 and USP22; this module is required for nuclear receptor-dependent transcriptional activation and counteracts heterochromatin silencing; ENY2 acts as an enhancer of position-effect variegation in Drosophila. Complex purification, deubiquitinase activity assays, Drosophila genetic PEV assay, nuclear receptor transcription assays Molecular cell High 18206972
2008 Sus1 is recruited to coding regions and is associated with the elongating form of RNA Pol II (phosphorylated on Ser5 and Ser2 of CTD); it co-purifies with mRNA export factors Yra1 and Mex67; Sgf73 is necessary for Sus1 association with both SAGA and TREX-2 and for its chromatin occupancy. ChIP, co-purification/co-IP, chromatin occupancy analysis Genes & development High 18923079
2009 Crystal structure of the Sus1:Cdc31:Sac3(CID) complex reveals Sus1 has an articulated helical hairpin fold that wraps around a long alpha-helix in Sac3; two Sus1 chains and one Cdc31 bind Sac3(CID); engineered mutations disrupting individual chain binding show Sus1 and Cdc31 act synergistically to promote NPC association of TREX-2 and mRNA export. X-ray crystallography, site-directed mutagenesis, in vivo NPC association assay, mRNA export assay Molecular cell High 19328066
2009 Crystal structure of the Sus1:Sgf11 complex shows Sus1 wraps around an extended alpha-helix of Sgf11's N-terminal region via a hydrophobic stripe; the Sus1-binding site on Sgf11 is shorter than on Sac3; a single Sus1 molecule cannot simultaneously bind Sgf11 and Sac3, indicating Sus1 forms separate subcomplexes in SAGA and TREX-2. X-ray crystallography, in vitro mutagenesis disruption assays The Journal of biological chemistry High 20007317
2009 Mutational dissection of Sus1 identified alleles (sus1-10, sus1-12) that dissociate from TREX-2 (Sac3) while retaining SAGA (Sgf11) interaction, and allele sus1-11 that disrupts both; all three TREX-2-dissociated mutants showed impaired NPC targeting of Sac3 and mRNA export defects; reduced affinity for Sac3/Sgf11 was confirmed in vitro. Mutagenesis, in vitro binding assays, in vivo NPC localization assay, mRNA export assay, genetic analysis The Journal of biological chemistry High 19269973
2010 Drosophila ENY2 is stably associated with the THO complex (involved in mRNP biogenesis), functioning independently of SAGA and AMEX/TREX-2; ENY2 and THO are loaded onto nascent mRNA after transcription activation of hsp70; ENY2 plays an important role in THO recruitment; ENY2/THO knockdown affects 3' end processing of transcripts. Co-purification/co-IP, ChIP, RNA knockdown (RNAi), 3' end processing analysis Genes & development High 20048002
2010 Sus1 has genetic interactions with cytoplasmic mRNA decay factors (LSM1, PAT1, LSM6, DHH1); sus1Δ is synthetic lethal with lsm1Δ and pat1Δ; Sus1 overexpression leads to its accumulation in cytoplasmic P-body/stress granule-like granules; Sus1 physically interacts with P-body/stress granule components. Genetic interaction analysis, co-localization by fluorescence microscopy, co-IP/pulldown BMC cell biology Medium 20230609
2012 Mammalian TREX-2 complex consists of GANP (scaffold), ENY2, PCID2, and centrins; the crystal structure of the GANP:ENY2 complex shows two ENY2 chains interacting with GANP in orientations different from those on yeast Sac3; depletion of any TREX-2 component including ENY2 inhibits mRNA export; GANP recruits ENY2 to nuclear pore complexes (NPCs), while ENY2 is not required for GANP recruitment. X-ray crystallography, siRNA knockdown, mRNA export assay, co-IP, NPC localization assay Nucleic acids research High 22307388
2013 Human TREX-2 stably associates with the nuclear pore basket and requires basket nucleoporins NUP153 and TPR but not active transcription for NPC association; ENY2 (unlike PCID2) is not involved in nuclear protein export or centrosome localization, indicating ENY2's role is specific to the classical mRNA export function of TREX-2. siRNA knockdown, co-IP, fluorescence microscopy, nuclear protein export assay Journal of cell science / Experimental cell research High 23591820 24291146
2014 Drosophila ENY2 is recruited to dCTCF-dependent insulators via the zinc-finger domain of dCTCF and is required for barrier activity; ENY2 RNAi in BG3 cells leads to spreading of H3K27 trimethylation and Polycomb protein at dCTCF boundaries. In vivo transgenic insulator assay, RNAi knockdown, chromatin immunoprecipitation for H3K27me3/Pc Epigenetics High 25147918
2016 ENY2 and ATXN7L3 serve as non-enzymatic adaptor subunits that orchestrate the activity of multiple H2B deubiquitinases: USP22 (SAGA DUBm), USP27X, and USP51; USP27X and USP51 function independently of SAGA and compete with USP22 for ATXN7L3 and ENY2; depletion of ENY2 or ATXN7L3 increases global H2Bub1 (unlike USP22 depletion which paradoxically decreases it); USP51 and USP27X depletion suppresses tumor growth. siRNA knockdown, mass spectrometry, H2Bub1 immunoblot, tumor growth assay Molecular cell High 27132940
2016 In Drosophila, ENY2 forms mutually exclusive complexes with insulator proteins (Su(Hw), dCTCF) and with Sgf11 (SAGA component); direct binding between ENY2 and insulator proteins or Sgf11 was demonstrated. Co-IP, direct binding assays (pulldown) Doklady. Biochemistry and biophysics Medium 27417714
2017 Sus1/ENY2 absence leads to elongated telomeres in yeast; Sus1 physically interacts with telomere maintenance factors; sus1Δ mutants show higher H2BK123ub1 levels, and this increase is correlated with longer telomeres; Sus1's H2B de-ubiquitination activity negatively regulates telomere length. Telomere length assay (Southern blot), co-IP, genetic epistasis with telomere maintenance mutants, H2Bub1 quantification Current genetics Medium 29116388
2018 Arabidopsis ENY2 is a component of the SAGA deubiquitination module (DUBm), interacting directly with SGF11 and UBP22; unlike yeast/metazoan orthologs, plant ENY2 is not enriched at the nuclear envelope and does not co-purify with TREX-2; plants depleted in ENY2 or SGF11 show increased H2Bub1 levels. Affinity purification-mass spectrometry, targeted proteomics (reciprocal tagging), homology modeling, H2Bub1 immunoblot Journal of molecular biology High 29588169
2018 Sus1/ENY2 binds widely to upstream regulatory regions of yeast genes genome-wide; heat shock relocates Sus1 from SAGA-dominated genes to stress-responsive genes; Sus1 contributes to both RNA synthesis and mRNA stability at a broad range of transcripts as shown by genomic run-on experiments. ChIP-seq, genomic run-on (GRO), genome-wide binding analysis Epigenetics & chromatin Medium 29598828
2020 Drosophila ENY2 interacts with RNA helicase MLE; this interaction was confirmed by independent methods and shown to be evolutionarily conserved; the interaction is important for MLE function in both sexes. Co-IP, pulldown, independent interaction assays Doklady. Biochemistry and biophysics Low 32130612
2022 Sus1/ENY2 maintains normal yeast replicative lifespan specifically through its role in TREX-2-mediated mRNA export (independently of SAGA DUBm); increased dosage of mRNA export factors Mex67 and Dbp5 rescues growth defects, shortened lifespan, and nuclear poly(A)+ RNA accumulation in sus1Δ cells; Sus1 is required for proper NPC association of Mex67 and Dbp5. Genetic epistasis, lifespan assay, mRNA export (FISH for poly(A)+), suppression by overexpression Aging Medium 35771153
2024 Drosophila ENY2 interacts with RNA-binding protein Paip2; Paip2 directly binds ENY2 in vitro and associates with the ENY2-containing TREX-2 complex in vivo; both Paip2 and ENY2 localize to the histone locus body (HLB); Paip2 knockdown reduces binding of TREX-2 subunits to histone mRNP particles, implicating ENY2/TREX-2 in histone mRNA export. Yeast two-hybrid, in vitro direct binding, co-IP, immunofluorescence, RNAi knockdown with mRNP association assay Molekuliarnaia biologiia Medium 39707855
2026 ENY2 overexpression enhances tumor growth and cell cycle progression; ENY2 depletion causes release of NPM1 into the nucleoplasm, impairing ribosomal subunit export and inducing nucleolar stress; released NPM1 interacts with MDM2 to stabilize p53 and suppress tumor growth; in p53-mutant cells, ENY2 knockdown augments RISC-mediated mRNA silencing (p53-independent pathway). In vivo tumor models, siRNA knockdown, co-IP, RNA-seq, polysome profiling, agarose gel electrophoresis (ribosome export), immunofluorescence, ubiquitination assays, molecular docking Cellular oncology Medium 41642454
2026 ENY2 is present at the histone locus body (HLB) in Drosophila polytene chromosomes as a component of SAGA, THO, and TREX-2 complexes; all three complexes interact with FLASH (HLB scaffold component); ENY2/SAGA and ENY2/THO associate with histone gene chromatin, while ENY2/TREX-2 interacts with histone mRNA for export. Immunofluorescence on polytene chromosomes, co-IP with FLASH, ChIP Doklady. Biochemistry and biophysics Medium 41912850

Source papers

Stage 0 corpus · 83 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2005 A human protein-protein interaction network: a resource for annotating the proteome. Cell 1704 16169070
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
2009 Defining the human deubiquitinating enzyme interaction landscape. Cell 1282 19615732
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
2015 A human interactome in three quantitative dimensions organized by stoichiometries and abundances. Cell 1015 26496610
2020 A reference map of the human binary protein interactome. Nature 849 32296183
2003 Complete sequencing and characterization of 21,243 full-length human cDNAs. Nature genetics 754 14702039
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
2010 Quantitative interaction proteomics and genome-wide profiling of epigenetic histone marks and their readers. Cell 639 20850016
2022 OpenCell: Endogenous tagging for the cartography of human cellular organization. Science (New York, N.Y.) 432 35271311
2005 Diversification of transcriptional modulation: large-scale identification and characterization of putative alternative promoters of human genes. Genome research 409 16344560
2015 Panorama of ancient metazoan macromolecular complexes. Nature 407 26344197
1996 Normalization and subtraction: two approaches to facilitate gene discovery. Genome research 401 8889548
2001 Human STAGA complex is a chromatin-acetylating transcription coactivator that interacts with pre-mRNA splicing and DNA damage-binding factors in vivo. Molecular and cellular biology 320 11564863
2008 A TFTC/STAGA module mediates histone H2A and H2B deubiquitination, coactivates nuclear receptors, and counteracts heterochromatin silencing. Molecular cell 307 18206972
2004 Sus1, a functional component of the SAGA histone acetylase complex and the nuclear pore-associated mRNA export machinery. Cell 299 14718168
1998 Function of TAF(II)-containing complex without TBP in transcription by RNA polymerase II. Nature 221 9603525
2014 Proximity biotinylation and affinity purification are complementary approaches for the interactome mapping of chromatin-associated protein complexes. Journal of proteomics 215 25281560
2001 UV-damaged DNA-binding protein in the TFTC complex links DNA damage recognition to nucleosome acetylation. The EMBO journal 169 11406595
1999 Identification of TATA-binding protein-free TAFII-containing complex subunits suggests a role in nucleosome acetylation and signal transduction. The Journal of biological chemistry 164 10373431
2019 H4K20me0 recognition by BRCA1-BARD1 directs homologous recombination to sister chromatids. Nature cell biology 162 30804502
2020 Synthetic Lethal and Resistance Interactions with BET Bromodomain Inhibitors in Triple-Negative Breast Cancer. Molecular cell 159 32416067
2019 A protein-interaction network of interferon-stimulated genes extends the innate immune system landscape. Nature immunology 159 30833792
2018 MYC Protein Interactome Profiling Reveals Functionally Distinct Regions that Cooperate to Drive Tumorigenesis. Molecular cell 152 30415952
2011 Sucrose synthase activity in the sus1/sus2/sus3/sus4 Arabidopsis mutant is sufficient to support normal cellulose and starch production. Proceedings of the National Academy of Sciences of the United States of America 135 22184213
2011 Interactions of pathological hallmark proteins: tubulin polymerization promoting protein/p25, beta-amyloid, and alpha-synuclein. The Journal of biological chemistry 131 21832049
2020 PROTAC-mediated degradation reveals a non-catalytic function of AURORA-A kinase. Nature chemical biology 129 32989298
2008 The THP1-SAC3-SUS1-CDC31 complex works in transcription elongation-mRNA export preventing RNA-mediated genome instability. Molecular biology of the cell 123 18667528
2012 Functional and structural characterization of the mammalian TREX-2 complex that links transcription with nuclear messenger RNA export. Nucleic acids research 119 22307388
2016 ATXN7L3 and ENY2 Coordinate Activity of Multiple H2B Deubiquitinases Important for Cellular Proliferation and Tumor Growth. Molecular cell 118 27132940
2009 Sus1, Cdc31, and the Sac3 CID region form a conserved interaction platform that promotes nuclear pore association and mRNA export. Molecular cell 118 19328066
2013 The human TREX-2 complex is stably associated with the nuclear pore basket. Journal of cell science 115 23591820
2006 The mRNA export factor Sus1 is involved in Spt/Ada/Gcn5 acetyltransferase-mediated H2B deubiquitinylation through its interaction with Ubp8 and Sgf11. Molecular biology of the cell 109 16855026
2008 Sus1 is recruited to coding regions and functions during transcription elongation in association with SAGA and TREX2. Genes & development 85 18923079
2007 Evolutionarily conserved E(y)2/Sus1 protein is essential for the barrier activity of Su(Hw)-dependent insulators in Drosophila. Molecular cell 74 17643381
2014 Novel three-component Rieske non-heme iron oxygenase system catalyzing the N-dealkylation of chloroacetanilide herbicides in sphingomonads DC-6 and DC-2. Applied and environmental microbiology 73 24928877
2020 Transactivation of Sus1 and Sus2 by Opaque2 is an essential supplement to sucrose synthase-mediated endosperm filling in maize. Plant biotechnology journal 62 32004404
2010 Multifunctional factor ENY2 is associated with the THO complex and promotes its recruitment onto nascent mRNA. Genes & development 61 20048002
2007 Sus1, Sac3, and Thp1 mediate post-transcriptional tethering of active genes to the nuclear rim as well as to non-nascent mRNP. RNA (New York, N.Y.) 51 18003937
1994 Structural features of the maize sus1 gene and protein. Plant physiology 47 7846165
2011 Key features of the two-intron Saccharomyces cerevisiae gene SUS1 contribute to its alternative splicing. Nucleic acids research 41 21749978
2009 The cap binding complex influences H2B ubiquitination by facilitating splicing of the SUS1 pre-mRNA. RNA (New York, N.Y.) 37 19561118
2009 Structural basis for the interaction between yeast Spt-Ada-Gcn5 acetyltransferase (SAGA) complex components Sgf11 and Sus1. The Journal of biological chemistry 33 20007317
2018 The Adaptor Protein ENY2 Is a Component of the Deubiquitination Module of the Arabidopsis SAGA Transcriptional Co-activator Complex but not of the TREX-2 Complex. Journal of molecular biology 32 29588169
2011 SUS1 introns are required for efficient mRNA nuclear export in yeast. Nucleic acids research 27 21749979
2009 A tale of coupling, Sus1 function in transcription and mRNA export. RNA biology 24 19246994
2010 ENY2: couple, triple...more? Cell cycle (Georgetown, Tex.) 21 20090412
2009 Transcription at the proximity of the nuclear pore: a role for the THP1-SAC3-SUS1-CDC31 (THSC) complex. RNA biology 19 19229139
2009 Mutational uncoupling of the role of Sus1 in nuclear pore complex targeting of an mRNA export complex and histone H2B deubiquitination. The Journal of biological chemistry 19 19269973
2014 Highly conserved ENY2/Sus1 protein binds to Drosophila CTCF and is required for barrier activity. Epigenetics 16 25147918
2018 The SAGA/TREX-2 subunit Sus1 binds widely to transcribed genes and affects mRNA turnover globally. Epigenetics & chromatin 15 29598828
2013 Human TREX2 components PCID2 and centrin 2, but not ENY2, have distinct functions in protein export and co-localize to the centrosome. Experimental cell research 13 24291146
2010 E(y)2/Sus1 is required for blocking PRE silencing by the Wari insulator in Drosophila melanogaster. Chromosoma 13 20082086
2010 A novel link between Sus1 and the cytoplasmic mRNA decay machinery suggests a broad role in mRNA metabolism. BMC cell biology 13 20230609
2019 Comparative genome analysis reveals the evolution of chloroacetanilide herbicide mineralization in Sphingomonas wittichii DC-6. Archives of microbiology 12 30997539
2022 Integrated multi-omics analysis identifies ENY2 as a predictor of recurrence and a regulator of telomere maintenance in hepatocellular carcinoma. Frontiers in oncology 10 35992857
2017 The evolutionarily conserved factor Sus1/ENY2 plays a role in telomere length maintenance. Current genetics 10 29116388
2014 Unveiling novel interactions of histone chaperone Asf1 linked to TREX-2 factors Sus1 and Thp1. Nucleus (Austin, Tex.) 10 24824343
1990 A molecular dynamics simulation of the (dG)6 . (dC)6 minihelix including counterions and water. Biopolymers 10 2369613
2022 Sus1 maintains a normal lifespan through regulation of TREX-2 complex-mediated mRNA export. Aging 8 35771153
2020 An insight into structural plasticity and conformational transitions of transcriptional co-activator Sus1. PloS one 8 32134955
2017 Protein arginine methylation of Npl3 promotes splicing of the SUS1 intron harboring non-consensus 5' splice site and branch site. Biochimica et biophysica acta. Gene regulatory mechanisms 6 28392442
2015 An intronic RNA structure modulates expression of the mRNA biogenesis factor Sus1. RNA (New York, N.Y.) 6 26546116
1991 DC6, a novel type of Dictyostelium discoideum gene regulated by secreted factors but not by cAMP. Differentiation; research in biological diversity 6 1717330
2023 An integrative analysis of enhancer of yellow 2 homolog (ENY2) as a molecular biomarker in pan-cancer. Functional & integrative genomics 4 36862319
2018 An exon three-way junction structure modulates splicing and degradation of the SUS1 yeast pre-mRNA. Biochimica et biophysica acta. Gene regulatory mechanisms 4 29966763
2012 Suppression of the nuclear factor Eny2 increases insulin secretion in poorly functioning INS-1E insulinoma cells. Experimental diabetes research 4 22649445
2009 Getting to the gate: crystallization of a Sac3(CID):Sus1:Cdc31 complex. Molecular cell 4 19328059
2020 Multifunctional ENY2 Protein Interacts with RNA Helicase MLE. Doklady. Biochemistry and biophysics 3 32130612
2007 A hexanucleotide sequence (dC1-dC6 tract) restricts the dC-specific cleavage of single-stranded DNA by endonuclease IV of bacteriophage T4. Nucleic acids research 3 17940096
2018 Role of the mRNA export factor Sus1 in oxidative stress tolerance in Candida albicans. Biochemical and biophysical research communications 2 29326041
2018 Zinc Finger Protein CG9890 - New Component of ENY2-Containing Complexes of Drosophila. Acta naturae 2 30713769
2024 [Drosophila melanogaster Paip2 Binds ENY2 and Interacts with the TREX-2 Complex in Histone mRNP Particles]. Molekuliarnaia biologiia 1 39707855
2018 Identification of the ORC Complex Subunits That Can Interact with the ENY2 Protein of Drosophila melanogaster. Doklady. Biochemistry and biophysics 1 30008099
2016 CTCF and Sgfl1 proteins form alternative complexes with ENY2 proteins. Doklady. Biochemistry and biophysics 1 27417714
2009 [Conservative E(y)2/Sus1 protein interacts with the Su(Hw)-dependent insulators in Drosophila]. Genetika 1 19382683
2009 [Conservative E(y)2/Sus1 protein is the member of SAGA complex and new nuclear pore-associated complex in Drosophila]. Genetika 1 19947544
2026 ENY2 transcription and export complex 2 subunit deficiency induces nucleolar stress to inhibit tumor progression through NPM1/MDM2/p53-dependent and -independent responses. Cellular oncology (Dordrecht, Netherlands) 0 41642454
2026 ENY2 Protein Is Present at the Histone Locus Body HLB As Part of Complexes with Different Functions in Transcription and mRNA Export. Doklady. Biochemistry and biophysics 0 41912850
2025 Characterization of stem cell landscape and identification of stemness-related gene ENY2 in colorectal cancer by intergrated transcriptomic analysis. Global medical genetics 0 40735504