Affinage

IWS1

Protein IWS1 homolog · UniProt Q96ST2

Length
819 aa
Mass
92.0 kDa
Annotated
2026-06-10
14 papers in source corpus 10 papers cited in narrative 10 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 6/6 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

IWS1 is an intrinsically disordered scaffold of the RNA Polymerase II transcription elongation machinery that couples elongation to chromatin modification, mRNA processing, and nuclear export (PMID:17234882, PMID:19141475, PMID:40835814). Cryo-EM of the activated elongation complex shows that the disordered C-terminal region of IWS1 uses short linear motifs to contact Pol II subunits RPB1, RPB2, and RPB5 together with the elongation factors DSIF, SPT6, and ELOF1, positioning downstream DNA within the Pol II cleft; recruitment depends on the RPB1 jaw and downstream-DNA contacts while stimulation of elongation velocity depends on the RPB2 lobe and ELOF1, and IWS1 also protects the complex from RECQL5 inhibition (PMID:40835814, PMID:42134803). Rapid IWS1 depletion globally lowers RNA synthesis and Pol II elongation velocity, establishing this elongation-stimulating activity as its core function (PMID:40835814). IWS1 bridges the CTD-binding proteins SPT6 and HYPB/Setd2 and is required for co-transcriptional H3K36me3 deposition across transcribed regions, and it recruits the export factor REF1/Aly to license bulk poly(A)+ RNA export downstream of SPT6 engagement of the Ser2-phosphorylated Pol II CTD (PMID:17234882, PMID:19141475). AKT-mediated phosphorylation of IWS1 amplifies an H3K36me3-dependent, LEDGF/SRSF1-directed splicing program controlling inclusion of U2AF2 exon 2, which in turn governs CDCA5/Sororin expression, ERK signaling, G2/M progression, and—via the RS-domain U2AF65 isoform that recruits Prp19—the export of CAR-element intronless mRNAs including interferon transcripts (PMID:34330897, PMID:34635782). In mouse preimplantation embryos IWS1 acts with SPT6 and nuclear AKT to support splicing, export, and Nanog expression required for development beyond the 8/16-cell stage (PMID:30846735).

Mechanistic history

Synthesis pass · year-by-year structured walk · 9 steps
  1. 2007 High

    Established IWS1 as the molecular link coupling transcription elongation to mRNA nuclear export, answering how export machinery is loaded co-transcriptionally.

    Evidence Co-IP, siRNA knockdown, ChIP, and nuclear poly(A)+ retention assays in HeLa cells

    PMID:17234882

    Open questions at the time
    • Direct structural basis of the IWS1–REF1/Aly contact not resolved
    • Whether export recruitment is separable from elongation function untested at this stage
  2. 2008 High

    Showed IWS1 is a physical bridge between SPT6 and Setd2 that directs co-transcriptional H3K36 trimethylation, defining its role in writing an elongation-coupled chromatin mark.

    Evidence siRNA knockdown, ChIP for H3K36me3/H3K27me3/acetylation, Co-IP, and in vitro Spt6-CTD binding with recombinant proteins

    PMID:19141475

    Open questions at the time
    • Did not distinguish whether H3K36me3 loss is direct or secondary to reduced transcription
    • No structural map of the SPT6–IWS1–Setd2 megacomplex
  3. 2019 Medium

    Extended the IWS1/SPT6 axis to development and linked nuclear AKT to H3K36me3 control, addressing how a signaling kinase modulates this chromatin output in vivo.

    Evidence siRNA microinjection in mouse embryos, Co-IP of IWS1 with nuclear AKT, H3K36me3 immunofluorescence, PI3K/AKT modulation

    PMID:30846735

    Open questions at the time
    • Phosphorylation site not mapped in this system
    • Causal chain from AKT to H3K36me3 inferred pharmacologically, not by direct enzymatic assay
  4. 2021 High

    Dissected how AKT-phosphorylated IWS1 drives an H3K36me3-dependent splicing program controlling U2AF2 exon 2 inclusion and downstream cell-cycle progression, connecting the scaffold to proliferation and cancer.

    Evidence RNA-seq after phosphorylation block, RT-PCR, ChIP, rescue experiments, xenografts, and EGFR-mutant lung adenocarcinoma specimens

    PMID:34330897

    Open questions at the time
    • IWS1 phosphosite kinetics relative to elongation not defined
    • Whether splicing effects persist independent of bulk elongation changes untested
  5. 2021 Medium

    Showed the phospho-IWS1-dependent RS-domain U2AF65 isoform recruits Prp19 to export CAR-element intronless mRNAs including interferon transcripts, linking the pathway to antiviral defense.

    Evidence RNA immunoprecipitation, Co-IP, siRNA knockdown, viral infection and caspase assays

    PMID:34635782

    Open questions at the time
    • Single-lab reciprocal Co-IP without structural validation
    • Direct contribution of IWS1 versus the downstream U2AF65 isoform not fully separated
  6. 2022 Medium

    Genetic epistasis in yeast placed Spn1/Iws1 at the intersection of multiple histone chaperone and acetylation pathways, framing its role as overcoming repressive chromatin during elongation.

    Evidence Bypass suppressor screen and genetic epistasis in S. cerevisiae

    PMID:35977387

    Open questions at the time
    • Genetic, not biochemical, relationships
    • Conservation of specific suppressor interactions to human IWS1 untested
  7. 2025 High

    Cryo-EM and degron kinetics redefined the core IWS1 function as direct stimulation of Pol II elongation via downstream-DNA positioning, and showed the H3K36me3 decrease is an indirect consequence of reduced transcription.

    Evidence Cryo-EM of the activated elongation complex, auxin-inducible degron multi-omics, in vitro transcription, and C-terminal SLiM mutagenesis

    PMID:40835814

    Open questions at the time
    • Reconciliation with earlier models placing H3K36me3 as a direct IWS1 output
    • Whether export/splicing roles are likewise downstream of elongation not fully resolved
  8. 2025 High

    Mapped the specific SLiMs by which IWS1 contacts Pol II subunits and elongation factors and defined which contacts drive recruitment versus stimulation, providing a structural mechanism for its scaffolding.

    Evidence Cryo-EM with individual SLiM mutants, in vitro transcription, recruitment assays, and RECQL5 competition assays

    PMID:42134803

    Open questions at the time
    • Functional consequence of RECQL5 protection in cells not quantified
    • How LEDGF nucleosome binding integrates with IWS1 elongation function unresolved
  9. 2025 Low

    Docking-guided screening identified the SPT6-binding region of IWS1 and candidate disruptors that alter IWS1 localization and tumor-cell behavior, opening a therapeutic targeting hypothesis.

    Evidence Molecular docking, Co-IP validation, migration/invasion/spheroid assays in dedifferentiated liposarcoma cells

    PMID:40594510

    Open questions at the time
    • No structural or biochemical reconstitution of the interaction domain
    • Inhibitor specificity and on-target mechanism not established

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved which IWS1 outputs (H3K36me3, splicing, export, antiviral signaling, cancer phenotypes) are direct functions versus indirect consequences of its core elongation-stimulating activity.
  • Causal hierarchy between elongation velocity and chromatin/splicing/export outputs not disentangled
  • AKT phosphosite role in the structural elongation mechanism unmapped
  • In vivo relevance of the RECQL5-protection function unknown

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
Localization
GO:0005634 nucleus 2
Pathway
R-HSA-74160 Gene expression (Transcription) 4 R-HSA-8953854 Metabolism of RNA 3 R-HSA-4839726 Chromatin organization 2
Complex memberships
RNA Pol II elongation complex

Evidence

Reading pass · 10 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2007 Human IWS1 (hIws1) was identified as an Spt6-interacting protein that associates with the mRNA nuclear export factor REF1/Aly; depletion of hIws1 caused mRNA processing defects, reduced REF1/Aly occupancy at the c-myc gene, and nuclear retention of bulk poly(A)+ RNAs, establishing IWS1 as a cotranscriptional recruiter of mRNA export machinery downstream of Spt6 binding to Ser2-phosphorylated RNAPII CTD. Co-immunoprecipitation, siRNA knockdown, chromatin immunoprecipitation (ChIP), nuclear poly(A)+ RNA retention assay in HeLa cells Genes & development High 17234882
2008 IWS1 bridges two CTD-binding proteins, Spt6 and HYPB/Setd2, in a megacomplex on the RNAPII elongation machinery; knockdown of IWS1 abolished H3K36me3 across the transcribed regions of c-Myc, HIV-1, and PABPC1 genes and also increased H3K27me3 at the 5' end of PABPC1 and histone acetylation across coding regions, demonstrating that IWS1 recruits Setd2 to direct co-transcriptional H3K36 trimethylation. siRNA knockdown, ChIP for H3K36me3/H3K27me3/acetylation, Co-IP, in vitro Spt6-CTD binding assay with recombinant proteins Genes & development High 19141475
2021 AKT-mediated phosphorylation of IWS1 promotes H3K36me3 deposition in target gene bodies, which controls LEDGF/SRSF1-dependent inclusion of exon 2 in U2AF2 pre-mRNA; the resulting exon-2-containing U2AF65 is required for proper CDCA5 pre-mRNA processing, Sororin expression, ERK phosphorylation, and G2/M progression. Loss of IWS1 phosphorylation produces an RS-domain-deficient U2AF65 that cannot support these downstream events, impairing cell proliferation. RNA-seq after IWS1 phosphorylation block, RT-PCR, ChIP for H3K36me3, functional rescue experiments, xenograft tumor growth assays, analysis of EGFR-mutant lung adenocarcinoma specimens Nature communications High 34330897
2021 The RS-domain-containing U2AF65 isoform (produced under phospho-IWS1-dependent splicing) recruits Prp19 to CAR-element-containing intronless mRNAs and promotes their nuclear export; U2AF65 loading to CAR-elements was RS-domain-independent but RNA Pol II-dependent. IWS1-phosphorylation-deficient cells express reduced IFNα1/IFNβ1 protein and show enhanced sensitivity to cytolytic virus infection. RNA immunoprecipitation, Co-IP, siRNA knockdown, viral infection assays, caspase activation assays Communications biology Medium 34635782
2019 In mouse preimplantation embryos, IWS1 interacts with nuclear AKT, and inhibition of the PI3K/AKT pathway reduced global H3K36me3 whereas activation increased it, suggesting AKT modulates H3K36me3 through interaction with IWS1; knockdown of Iws1 or Supt6 individually blocked development at the 8/16-cell stage with defects in pre-mRNA splicing, mRNA export, and Nanog expression. siRNA microinjection in mouse embryos, Co-IP (IWS1 with nuclear AKT), immunofluorescence for H3K36me3, PI3K/AKT pathway inhibitor/activator treatment Scientific reports Medium 30846735
2025 Cryo-EM structure of the activated Pol II elongation complex shows IWS1 acts as a scaffold that positions downstream DNA within the cleft of Pol II. The intrinsically disordered C-terminal region of IWS1 contacts the Pol II cleft and, together with ELOF1, stimulates Pol II elongation velocity. Rapid depletion of IWS1 in human cells caused a global decrease in RNA synthesis and Pol II elongation velocity; the associated decrease in H3K36me3 was found to be an indirect, secondary consequence of reduced transcription rather than a direct IWS1 function. Cryo-EM structure determination, multi-omics kinetic analysis after auxin-inducible IWS1 degron, in vitro transcription assays, mutagenesis of IWS1 C-terminal disordered region Nature communications High 40835814
2025 Cryo-EM mapping revealed that the intrinsically disordered C-terminal region of IWS1 contains short linear motifs (SLiMs) that contact Pol II subunits RPB1, RPB2, and RPB5, elongation factors DSIF, SPT6, and ELOF1; IWS1 recruitment to the elongation complex requires the RPB1 jaw interaction and downstream DNA binding, while transcription stimulation requires RPB2 lobe and ELOF1 contacts. IWS1 was found to protect the elongation complex from RECQL5 inhibition. Additionally, the histone reader LEDGF (an IWS1 interactor) was shown to bind a transcribed downstream nucleosome. Cryo-EM structure determination, in vitro transcription assays with IWS1 SLiM mutants, functional recruitment assays, RECQL5 competition assays Nucleic acids research High 42134803
2025 Molecular docking using the AlphaFold-predicted IWS1 structure identified a core Spt6-binding region (AA 545–694) of IWS1; candidate small-molecule inhibitors Ketotifen and Desloratadine were predicted to mimic Spt6 Phe217 and disrupt the IWS1/Spt6 complex, which was confirmed by co-immunoprecipitation. Disruption of this interaction increased nuclear localization of IWS1 and reduced migration, invasion, and spheroid formation in dedifferentiated liposarcoma cells. Molecular docking/virtual screening, Co-immunoprecipitation, cell migration/invasion assays, spheroid formation assay, immunofluorescence for subcellular localization Scientific reports Low 40594510
2022 Genetic suppressor screen in S. cerevisiae showed that viability in the absence of Spn1/Iws1 can be achieved by mutations in FACT, Set2, Rpd3S, Rtt109, Chd1, or Sgf73, placing Spn1/Iws1 function at the intersection of histone acetylation and multiple histone chaperone pathways; this epistasis indicates Spn1 acts to overcome repressive chromatin through multiple mechanisms during transcription elongation. Suppressor genetics (bypass suppressor screen), yeast viability assays, genetic epistasis Genetics Medium 35977387
2025 IWS1's intrinsically disordered C-terminal region (SLiMs) is responsible for stimulation of Pol II elongation activity, as demonstrated by in vitro assays; this region acts together with ELOF1 to enhance Pol II processivity. In vitro transcription elongation assay, cryo-EM, SLiM mutagenesis bioRxivpreprint Medium 40909601

Source papers

Stage 0 corpus · 14 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2007 The Spt6 SH2 domain binds Ser2-P RNAPII to direct Iws1-dependent mRNA splicing and export. Genes & development 214 17234882
2008 The Iws1:Spt6:CTD complex controls cotranscriptional mRNA biosynthesis and HYPB/Setd2-mediated histone H3K36 methylation. Genes & development 200 19141475
2019 Iws1 and Spt6 Regulate Trimethylation of Histone H3 on Lysine 36 through Akt Signaling and are Essential for Mouse Embryonic Genome Activation. Scientific reports 24 30846735
2022 Suppressor mutations that make the essential transcription factor Spn1/Iws1 dispensable in Saccharomyces cerevisiae. Genetics 10 35977387
2018 Genetic ablation of interacting with Spt6 (Iws1) causes early embryonic lethality. PloS one 10 30208029
2021 AKT3-mediated IWS1 phosphorylation promotes the proliferation of EGFR-mutant lung adenocarcinomas through cell cycle-regulated U2AF2 RNA splicing. Nature communications 9 34330897
2023 Phosphorylation of IWS1 by AKT maintains liposarcoma tumor heterogeneity through preservation of cancer stem cell phenotypes and mesenchymal-epithelial plasticity. Oncogenesis 7 37237004
2023 Toxoplasma IWS1 Determines Fitness in Interferon-γ-Activated Host Cells and Mice by Indirectly Regulating ROP18 mRNA Expression. mBio 5 36715543
2025 IWS1 positions downstream DNA to globally stimulate Pol II elongation. Nature communications 3 40835814
2010 Crystallization and preliminary crystallographic analysis of eukaryotic transcription and mRNA export factor Iws1 from Encephalitozoon cuniculi. Acta crystallographica. Section F, Structural biology and crystallization communications 3 20124725
2025 Structure and function of IWS1 in transcription elongation. bioRxiv : the preprint server for biology 1 40909601
2021 Phosphor-IWS1-dependent U2AF2 splicing regulates trafficking of CAR-E-positive intronless gene mRNAs and sensitivity to viral infection. Communications biology 1 34635782
2026 Structure and function of IWS1 in transcription elongation. Nucleic acids research 0 42134803
2025 Molecular docking and biological evaluation of a novel IWS1 inhibitor for the treatment of human retroperitoneal liposarcoma. Scientific reports 0 40594510

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