Affinage

SAFB

Scaffold attachment factor B1 · UniProt Q15424

Length
915 aa
Mass
102.6 kDa
Annotated
2026-06-10
100 papers in source corpus 17 papers cited in narrative 17 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

SAFB (SAFB1; also called HET/SAF-B) is a multifunctional nuclear scaffold protein that directly binds scaffold/matrix attachment region (S/MAR) DNA and integrates higher-order chromatin organization with transcription and RNA processing (PMID:8600450, PMID:9671816). Through its SAP DNA-binding domain it tethers itself to chromatin and physically associates with RNA polymerase II and SR splicing factors, localizing to SC35 nuclear speckles and coupling S/MAR chromatin to pre-mRNA splicing (PMID:9671816, PMID:27731383); SUMO-1 modification at ribosomal protein gene promoters enhances Pol II recruitment and splicing (PMID:25800734). iCLIP maps SAFB1 to a purine-rich GAAGA motif genome-wide and shows it controls alternative splicing of targets such as NCAM1, with consequences for neuronal dendritic spine density (PMID:26694817). SAFB acts predominantly as a transcriptional corepressor: it binds the estrogen receptor DNA-binding domain (an interaction strengthened by tamoxifen) and cooperates with the corepressor N-CoR, which is required for its repressive activity, to silence estrogen-repressed apoptotic genes (PMID:10707955, PMID:16195251, PMID:19901029). It also directly represses defined promoters including hsp27, the hXOR E-box (in complex with Ku86 and BRG1), and the TAK1 promoter, the latter suppressing downstream NF-κB signaling and tumor-aggressive phenotypes (PMID:9328833, PMID:18772145, PMID:28912140). Conversely, SAFB1 promotes myogenic differentiation by associating with MyoD and enabling loss of repressive H3K27me3/Ezh2 and recruitment of myogenin and Brg1 at muscle gene promoters (PMID:23609547). SAFB1 is recruited to damaged chromatin in a PARP1/poly(ADP-ribose)-dependent manner to facilitate γH2AX spreading and checkpoint activation (PMID:24055346), and it stabilizes pericentromeric heterochromatin architecture through RNA-dependent phase separation mediated by its R/G-rich region binding satellite repeat RNAs (PMID:31677973). During apoptosis SAFB1 relocalizes from nucleolus to a peri-nucleolar ring via its coiled-coil domains and is cleaved by caspase-3 (PMID:17643427).

Mechanistic history

Synthesis pass · year-by-year structured walk · 16 steps
  1. 1996 High

    Established the molecular identity of SAFB as a chromatin protein that specifically recognizes S/MAR DNA, defining its foundational role in nuclear architecture.

    Evidence Protein purification, cDNA cloning, and subcellular fractionation from HeLa cells

    PMID:8600450

    Open questions at the time
    • No functional consequence of S/MAR binding established
    • Domain responsible for DNA binding not yet mapped
  2. 1997 High

    Showed SAFB directly binds a specific promoter (hsp27) and represses its activity, introducing SAFB as a sequence-targeted transcriptional repressor.

    Evidence EMSA, GST-pulldown, nuclear matrix fractionation, and luciferase reporter assays in breast cancer cells

    PMID:9328833

    Open questions at the time
    • Mechanism of repression not defined
    • No corepressor partner identified
  3. 1998 High

    Connected SAFB's chromatin role to RNA processing by demonstrating physical and functional coupling with RNA Pol II and SR proteins.

    Evidence Co-IP, SC35 colocalization, and E1A/S-MAR splicing and CAT reporter assays

    PMID:9671816

    Open questions at the time
    • Direct RNA target sequences not identified
    • Domain mediating SR-protein/Pol II contact unmapped
  4. 2000 High

    Defined SAFB as an estrogen receptor corepressor by mapping its binding to the ER DNA-binding/hinge region and showing tamoxifen-enhanced antagonism.

    Evidence In vitro binding, co-IP, and ER transactivation reporter assays with domain mapping

    PMID:10707955

    Open questions at the time
    • Corepressor machinery recruited by SAFB not yet identified
    • Endogenous ER target genes not defined
  5. 2000 Medium

    Expanded SAFB's RNA-processing interactome and linked it to negative gene regulation through isoform-specific binding to AUF1/hnRNP D.

    Evidence Yeast two-hybrid, nuclear colocalization, heterokaryon shuttling, and reporter assays

    PMID:10933876

    Open questions at the time
    • No reciprocal Co-IP from mammalian cells
    • Target transcripts of the complex not identified
  6. 2000 Medium

    Linked SAFB to growth control by showing overexpression drives growth arrest and multinucleation in breast cancer cells.

    Evidence GFP-fusion overexpression, proliferation assays, and tumor sample Western blots

    PMID:10999774

    Open questions at the time
    • Molecular basis of growth arrest unresolved
    • Single-lab overexpression phenotype
  7. 2005 High

    Identified N-CoR as the corepressor required for SAFB-mediated ER repression, providing a mechanistic basis for its repressive activity at endogenous promoters.

    Evidence Reciprocal Co-IP, in vitro domain-mapped binding, ChIP at pS2, and siRNA functional rescue

    PMID:16195251

    Open questions at the time
    • Whether additional corepressors act at other targets unknown
    • Dynamics of co-release upon estrogen not structurally resolved
  8. 2008 Medium

    Demonstrated context-specific repression and signal-responsive regulation of SAFB via the hXOR promoter and an OSM/DNA-PK phosphorylation input.

    Evidence GST-pulldown, Co-IP, ChIP, siRNA, and promoter reporter assays

    PMID:18772145

    Open questions at the time
    • Phosphorylation sites on SAFB not mapped
    • Functional role of Ku86/BRG1 association not dissected
  9. 2009 Medium

    Defined the genome-wide promoter occupancy and corepressor logic of SAFB1/SAFB2, confirming a predominant role in repressing estrogen-regulated apoptotic genes.

    Evidence ChIP-on-chip, expression microarray, and siRNA knockdown in MCF-7 cells

    PMID:19901029

    Open questions at the time
    • Direct vs indirect target distinction limited
    • Functional redundancy of SAFB1 vs SAFB2 not resolved
  10. 2013 High

    Placed SAFB1 in the DNA damage response as a PARP1/PAR-recruited factor enabling γH2AX spreading and checkpoint activation.

    Evidence FRAP at damage sites, PARP inhibition, γH2AX spreading, and checkpoint assays with siRNA

    PMID:24055346

    Open questions at the time
    • Direct PAR-binding domain not defined
    • Mechanism coupling SAFB to γH2AX spreading unresolved
  11. 2013 High

    Revealed a positive transcriptional role for SAFB1 in myogenic differentiation by relieving Polycomb repression and enabling myogenin/Brg1 recruitment.

    Evidence Co-IP with MyoD, ChIP for histone marks and chromatin regulators, and gain/loss-of-function in C2C12 cells

    PMID:23609547

    Open questions at the time
    • How SAFB1 directs Ezh2/H3K27me3 removal mechanistically unknown
    • Generality beyond muscle lineage untested
  12. 2015 Medium

    Showed SUMO-1 modification at ribosomal protein gene promoters as a switch that stimulates Pol II recruitment and pre-mRNA splicing by SAFB.

    Evidence ChIP for SUMO-1 and Pol II, siRNA depletion, and splicing assays

    PMID:25800734

    Open questions at the time
    • SUMOylation inferred from ChIP co-occupancy rather than direct site mapping
    • SUMO E3 ligase not identified
  13. 2015 High

    Provided a nucleotide-resolution RNA-binding map defining the GAAGA motif and demonstrating direct control of alternative splicing with a neuronal phenotype.

    Evidence iCLIP, NCAM1 minigene mutagenesis, exon array, and hippocampal neuron knockdown

    PMID:26694817

    Open questions at the time
    • Structural basis of motif recognition unknown
    • Breadth of physiologically relevant splicing targets incomplete
  14. 2016 Medium

    Established SAFB1 as a chromatin tether for FUS via its SAP domain and linked it to AR-regulated transcription and ALS-relevant cytoplasmic sequestration.

    Evidence Yeast two-hybrid, Co-IP, chromatin fractionation with domain mapping, and splicing/AR transactivation assays

    PMID:27731383

    Open questions at the time
    • DNA-dependence of FUS interaction not fully resolved
    • Disease relevance of sequestration in patient tissue untested
  15. 2017 Medium

    Defined the SAFB–TAK1–NF-κB axis whereby SAFB directly represses TAK1 to suppress NF-κB signaling and colorectal cancer aggressiveness.

    Evidence ChIP, reporter assays, and TAK1 rescue epistasis in vitro and in xenografts

    PMID:28912140

    Open questions at the time
    • Whether N-CoR or other corepressors mediate TAK1 repression untested
    • Single-lab in vivo model
  16. 2019 High

    Demonstrated that SAFB stabilizes pericentromeric heterochromatin architecture through RNA-dependent phase separation, defining a structural genome-organizing function.

    Evidence RNA-IP of satellite RNAs, phase separation assays, Hi-C, and R/G-rich domain loss-of-function

    PMID:31677973

    Open questions at the time
    • Quantitative phase-separation parameters in vivo not defined
    • How this links to its transcriptional/splicing roles unclear

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved how SAFB's distinct activities — S/MAR scaffolding, RNA-guided phase separation, sequence-specific transcriptional repression, splicing regulation, and DNA-damage signaling — are coordinated and switched within a single protein.
  • No integrated model linking phase separation, chromatin scaffolding, and transcriptional output
  • Post-translational switches (SUMO, phosphorylation, caspase cleavage) not unified
  • Functional division of labor between SAFB1 and SAFB2 not defined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 5 GO:0003677 DNA binding 4 GO:0003723 RNA binding 2 GO:0060090 molecular adaptor activity 2 GO:0098772 molecular function regulator activity 2
Localization
GO:0000228 nuclear chromosome 2 GO:0005634 nucleus 2 GO:0005654 nucleoplasm 2 GO:0005730 nucleolus 1
Pathway
R-HSA-74160 Gene expression (Transcription) 4 R-HSA-8953854 Metabolism of RNA 3 R-HSA-162582 Signal Transduction 2 R-HSA-4839726 Chromatin organization 2 R-HSA-73894 DNA Repair 1
Partners
AUF1/HNRNP DBRG1ESR1FUSKU86MYODNCOR1RNA POL II

Evidence

Reading pass · 17 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1996 SAF-B (SAFB1) was purified from HeLa cells as a novel nuclear protein that specifically binds scaffold/matrix attachment region (S/MAR) DNA. The cDNA encodes an 849-amino acid protein with two putative bipartite nuclear localization signals and aberrant SDS-PAGE migration due to high charged residue content. SAF-B was found as an abundant chromatin component but not a nuclear matrix component. Protein purification to near-homogeneity, immunoscreening cDNA library, antibody generation, subcellular fractionation Nucleic acids research High 8600450
1997 HET (SAFB1) binds to the hsp27 promoter region containing an imperfect ERE and TATA box. A GST-fusion of partial HET clone demonstrated direct binding to the hsp27 promoter fragment in gel-retardation assays. HET localizes to the nuclear matrix in breast cancer cell lines, and overexpression caused a dose-dependent decrease of hsp27 promoter activity in transient transfection reporter assays. Gel-shift (EMSA), GST-pulldown, nuclear matrix fractionation, transient transfection with luciferase reporter Journal of cellular biochemistry High 9328833
1998 SAF-B interacts with RNA polymerase II and a subset of SR proteins (serine/arginine-rich RNA processing factors). SAF-B localizes to the nucleus in a speckled pattern co-incident with SR protein SC35. Overexpression of SAF-B induced an increase of the 10S E1A splice product and repressed activity of an S/MAR-flanked CAT reporter gene, indicating SAF-B couples S/MAR chromatin organization with transcription and pre-mRNA splicing. Co-immunoprecipitation, immunofluorescence colocalization, E1A splicing reporter assay, CAT reporter assay with S/MAR elements Nucleic acids research High 9671816
2000 HET/SAF-B is an estrogen receptor (ER)-interacting protein. In vitro binding assays showed HET binds to the ER DNA-binding domain and hinge region. Co-immunoprecipitation revealed HET/SAF-B and ER associate in cell lines with or without estradiol, but binding is increased by tamoxifen. HET/SAF-B enhances tamoxifen antagonism of estrogen-induced ER transactivation and, at high concentrations, represses both estrogen- and tamoxifen-induced ER activity in a manner dependent on ER-DBD interaction. In vitro binding assay, co-immunoprecipitation, transactivation reporter assay Molecular endocrinology (Baltimore, Md.) High 10707955
2000 SAF-B interacts specifically with p45 and p42 isoforms of AUF1/hnRNP D (but not p40/p37) via yeast two-hybrid and co-localizes with these isoforms as nuclear speckles. The p45/p42-specific C-terminal sequence mediates nuclear retention and interaction with SAF-B. The p45/p42–SAF-B complex acts as a negative regulator of gene expression. Yeast two-hybrid screening, nuclear colocalization (immunofluorescence), heterokaryon shuttling assay, gene expression reporter assay Archives of biochemistry and biophysics Medium 10933876
2000 Overexpression of HET/SAF-B caused growth arrest, formation of multinucleated cells, and was inversely correlated with cell proliferation in breast cancer cell lines, demonstrating a direct role for SAFB1 in cell division and growth control. Transient transfection with GFP-fusion, cell proliferation assays, Western blot in breast tumor samples Clinical cancer research : an official journal of the American Association for Cancer Research Medium 10999774
2005 SAFB1 interacts directly with the nuclear receptor corepressor N-CoR. This interaction is mediated in vitro and in vivo through the C-terminal region of SAFB1 (aa 600–915) and N-terminal region of N-CoR (aa 1–373). SAFB1 and N-CoR are co-recruited to the pS2 promoter in the absence of estrogen and are co-released upon estrogen addition. SAFB1-mediated repression of ER activity is significantly reduced by N-CoR siRNA knockdown, indicating the interaction is necessary for SAFB1's repressive activity. Co-immunoprecipitation, co-fractionation, colocalization, in vitro binding with defined domains, siRNA knockdown, chromatin immunoprecipitation (ChIP), reporter assay Molecular endocrinology (Baltimore, Md.) High 16195251
2007 During apoptosis, SAFB1 undergoes dynamic relocalization: it moves to the nucleolus within 15 min of apoptosis induction (before cytochrome c release), then forms a peri-nucleolar ring-like structure (after cytochrome c release, before PARP cleavage). The peri-nucleolar ring depends on RNA integrity and is mediated by the S/K and R/E coiled-coil domains (not the SAP DNA-binding or RRM RNA-binding domains). SAFB1 is subsequently cleaved by caspase-3, with the determinant for cleavage located in the DNA-binding domain. Live-cell fluorescence imaging, deletion mutant analysis, RNase treatment, caspase-3 cleavage assay, time-course fractionation Experimental cell research Medium 17643427
2008 SAFB1 directly binds the E-box element of the human xanthine oxidoreductase (hXOR) promoter. SAFB1, Ku86, and BRG1 associate with each other in pulldown, co-immunoprecipitation, and ChIP assays. SAFB1 silencing increases hXOR expression. Oncostatin M (OSM) induces phosphorylation of SAFB1 and promotes hXOR mRNA expression in a manner inhibited by silencing DNA-PK catalytic subunit or SAFB1. GST-pulldown, co-immunoprecipitation, chromatin immunoprecipitation, siRNA knockdown, promoter reporter assay The Journal of biological chemistry Medium 18772145
2009 SAFB1 and SAFB2 occupy promoters of endogenous target genes including immune regulators and apoptotic genes (BBC3, NEDD9, OPG) in MCF-7 cells. ChIP-on-chip identified 541 SAFB1/SAFB2-binding sites. Most target genes are induced upon SAFB1/SAFB2 depletion. 12% of estrogen-regulated genes depend on SAFB1, primarily estrogen-repressed apoptotic genes, confirming SAFB1's primary role as a corepressor. Chromatin immunoprecipitation-on-chip (ChIP-on-chip), gene expression microarray, siRNA knockdown The Journal of biological chemistry Medium 19901029
2013 SAFB1 (chromatin scaffold protein) is a component of the DNA damage response. SAFB1 undergoes highly dynamic exchange at damaged chromatin in a PARP1- and poly(ADP-ribose)-dependent manner (measured by FRAP). SAFB1 cooperates with histone acetylation to allow efficient γH2AX spreading and genotoxic stress signaling. Loss of SAFB1 impairs cell-cycle checkpoint activation and increases susceptibility to replicative stress. FRAP (live-cell imaging at damaged chromatin), siRNA knockdown, γH2AX spreading assay, cell-cycle checkpoint assay, PARP inhibitor treatment Molecular cell High 24055346
2013 Safb1 functions as a positive regulator of myogenic differentiation. Safb1 co-immunoprecipitated with MyoD and co-localized on myogenic gene promoters. Knockdown of Safb1 caused persistence of the repressive H3K27me3 mark and Polycomb methyltransferase Ezh2 at myogenic promoters, and blocked recruitment of myogenin and SWI/SNF ATPase Brg1. Overexpression of Safb1 caused premature expression of muscle structural proteins and enlarged myotubes. Co-immunoprecipitation with MyoD, ChIP for histone marks and chromatin regulators, siRNA knockdown, overexpression in C2C12 myoblasts Nucleic acids research High 23609547
2015 SAFB is SUMOylated by SUMO-1 at ribosomal protein gene promoters. SUMOylated SAFB stimulates both RNA polymerase II binding to promoters and pre-mRNA splicing. Depletion of SAFB decreased RNA Pol II binding to promoters and impaired nuclear mRNA processing without affecting mRNA stability. ChIP for SUMO-1 and RNA Pol II, SAFB depletion (siRNA), splicing assays, promoter binding assays Nucleic acids research Medium 25800734
2015 iCLIP mapping demonstrated that SAFB1 binds RNA genome-wide with enrichment in exons, ncRNAs, 3' and 5' UTRs, recognizing a purine-rich GAAGA (core AGA/GAA/AAG) motif. SAFB1 regulates alternative splicing of NCAM1 via these motifs (mutating the AGA/GAA/AAG sites abolished SAFB1-dependent splicing changes). SAFB1 knockdown altered dendritic spine density in hippocampal neurons. iCLIP (individual-nucleotide resolution CLIP), RT-PCR, NCAM1 minigene splicing assay with motif mutagenesis, exon array, neuronal knockdown and spine density imaging BMC biology High 26694817
2016 FUS interacts with nuclear matrix-associated protein SAFB1 in a DNA-dependent manner, as shown by yeast two-hybrid screening and co-immunoprecipitation. The N-terminal SAP (DNA-binding) domain of SAFB1 is required for its localization to the chromatin-bound fraction and for splicing regulation. Depletion of SAFB1 reduced FUS localization to the chromatin-bound fraction and splicing activity, suggesting SAFB1 tethers FUS to chromatin. FUS and SAFB1 also interact with androgen receptor (AR) to regulate ligand-dependent transcription. ALS-linked FUS mutants sequestered endogenous SAFB1 into cytoplasmic aggregates. Yeast two-hybrid, co-immunoprecipitation, chromatin fractionation, siRNA knockdown, splicing assay, AR transactivation assay Scientific reports Medium 27731383
2017 SAFB directly binds the TAK1 promoter (shown by ChIP) and represses TAK1 transcription, thereby suppressing downstream NF-κB signaling. Ectopic SAFB expression inhibited NF-κB activity and aggressive CRC phenotypes in vitro and in vivo; overexpression of TAK1 rescued these effects, establishing the SAFB–TAK1–NF-κB axis. Chromatin immunoprecipitation (ChIP), luciferase reporter assay, Western blot, RT-PCR, siRNA/overexpression in vitro and in vivo (xenograft) Clinical cancer research : an official journal of the American Association for Cancer Research Medium 28912140
2019 SAFB interacts with major satellite repeat RNAs via its R/G-rich region, and these RNAs promote phase separation driven by SAFB. Depletion of SAFB leads to decondensation of pericentromeric heterochromatin, increased interchromosomal interactions adjacent to pericentromeric regions, and decreased genomic compartmentalization, demonstrating SAFB stabilizes heterochromatin architecture through an RNA-dependent phase separation mechanism. RNA immunoprecipitation, phase separation assay, Hi-C (3D genome organization), SAFB depletion (siRNA/knockout), domain mapping of R/G-rich region Molecular cell High 31677973

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2008 Amyloid fibrils of the HET-s(218-289) prion form a beta solenoid with a triangular hydrophobic core. Science (New York, N.Y.) 794 18339938
1997 The protein product of the het-s heterokaryon incompatibility gene of the fungus Podospora anserina behaves as a prion analog. Proceedings of the National Academy of Sciences of the United States of America 371 9275200
2005 Correlation of structural elements and infectivity of the HET-s prion. Nature 358 15944710
1990 Addition of telomere-associated HeT DNA sequences "heals" broken chromosome ends in Drosophila. Cell 219 2111731
2002 Amyloid aggregates of the HET-s prion protein are infectious. Proceedings of the National Academy of Sciences of the United States of America 208 12032295
2003 Domain organization and structure-function relationship of the HET-s prion protein of Podospora anserina. The EMBO journal 165 12727874
1998 SAF-B protein couples transcription and pre-mRNA splicing to SAR/MAR elements. Nucleic acids research 155 9671816
1998 Phospholipid signalling in the nucleus. Een DAG uit het leven van de inositide signalering in de nucleus. Biochimica et biophysica acta 155 9838115
1992 HeT-A, a transposable element specifically involved in "healing" broken chromosome ends in Drosophila melanogaster. Molecular and cellular biology 135 1324409
1994 Analysis of a Het- mutation in Anabaena sp. strain PCC 7120 implicates a secondary metabolite in the regulation of heterocyst spacing. Journal of bacteriology 133 8157596
1992 Frequent transpositions of Drosophila melanogaster HeT-A transposable elements to receding chromosome ends. The EMBO journal 127 1330538
1996 Purification and molecular cloning of the scaffold attachment factor B (SAF-B), a novel human nuclear protein that specifically binds to S/MAR-DNA. Nucleic acids research 126 8600450
2019 The Nuclear Matrix Protein SAFB Cooperates with Major Satellite RNAs to Stabilize Heterochromatin Architecture Partially through Phase Separation. Molecular cell 122 31677973
2012 The mechanism of toxicity in HET-S/HET-s prion incompatibility. PLoS biology 112 23300377
2011 The [Het-s] prion of Podospora anserina and its role in heterokaryon incompatibility. Seminars in cell & developmental biology 99 21334447
2001 The HET-s prion protein of the filamentous fungus Podospora anserina aggregates in vitro into amyloid-like fibrils. The Journal of biological chemistry 87 11733532
1997 Novel nuclear matrix protein HET binds to and influences activity of the HSP27 promoter in human breast cancer cells. Journal of cellular biochemistry 87 9328833
1999 Vestibular responses to linear acceleration are absent in otoconia-deficient C57BL/6JEi-het mice. Hearing research 85 10491954
2008 On the binding of Thioflavin-T to HET-s amyloid fibrils assembled at pH 2. Journal of structural biology 84 18406172
2000 Tamoxifen-bound estrogen receptor (ER) strongly interacts with the nuclear matrix protein HET/SAF-B, a novel inhibitor of ER-mediated transactivation. Molecular endocrinology (Baltimore, Md.) 84 10707955
2020 Electrophotocatalytic Undirected C-H Trifluoromethylations of (Het)Arenes. Chemistry (Weinheim an der Bergstrasse, Germany) 80 31875327
1988 A spontaneously opened ring chromosome of Drosophila melanogaster has acquired He-T DNA sequences at both new telomeres. Proceedings of the National Academy of Sciences of the United States of America 80 3141921
2003 Sexual transmission of the [Het-S] prion leads to meiotic drive in Podospora anserina. Proceedings of the National Academy of Sciences of the United States of America 78 12719532
2008 Effect of curcumin on acidic pH-induced expression of IL-6 and IL-8 in human esophageal epithelial cells (HET-1A): role of PKC, MAPKs, and NF-kappaB. American journal of physiology. Gastrointestinal and liver physiology 70 19074641
2000 Attachment of HeT-A sequences to chromosomal termini in Drosophila melanogaster may occur by different mechanisms. Molecular and cellular biology 67 11003659
2012 Genomic clustering and homology between HET-S and the NWD2 STAND protein in various fungal genomes. PloS one 66 22493719
2016 FUS interacts with nuclear matrix-associated protein SAFB1 as well as Matrin3 to regulate splicing and ligand-mediated transcription. Scientific reports 65 27731383
2000 A nuclear matrix-associated factor, SAF-B, interacts with specific isoforms of AUF1/hnRNP D. Archives of biochemistry and biophysics 65 10933876
1989 Studies of He-T DNA sequences in the pericentric regions of Drosophila chromosomes. Chromosoma 65 2565198
2002 HET-E and HET-D belong to a new subfamily of WD40 proteins involved in vegetative incompatibility specificity in the fungus Podospora anserina. Genetics 63 12019224
2013 The chromatin scaffold protein SAFB1 renders chromatin permissive for DNA damage signaling. Molecular cell 61 24055346
2007 The fungus-specific HET domain mediates programmed cell death in Podospora anserina. Eukaryotic cell 59 17873080
2000 Vegetative incompatibility in the het-6 region of Neurospora crassa is mediated by two linked genes. Genetics 59 10880472
1994 Structure of the Drosophila HeT-A transposon: a retrotransposon-like element forming telomeres. Chromosoma 59 7924625
2007 Prion and non-prion amyloids of the HET-s prion forming domain. Journal of molecular biology 58 17532341
1996 The product of the het-C heterokaryon incompatibility gene of Neurospora crassa has characteristics of a glycine-rich cell wall protein. Genetics 57 8844148
2011 Structural dependence of HET-s amyloid fibril infectivity assessed by cryoelectron microscopy. Proceedings of the National Academy of Sciences of the United States of America 56 21300906
2006 HET-SOFAST NMR for fast detection of structural compactness and heterogeneity along polypeptide chains. Magnetic resonance in chemistry : MRC 56 16823898
1990 HeT DNA: a family of mosaic repeated sequences specific for heterochromatin in Drosophila melanogaster. Proceedings of the National Academy of Sciences of the United States of America 55 2122452
2010 Bile acid toxicity structure-activity relationships: correlations between cell viability and lipophilicity in a panel of new and known bile acids using an oesophageal cell line (HET-1A). Bioorganic & medicinal chemistry 53 20713311
2014 Evolutionary link between metazoan RHIM motif and prion-forming domain of fungal heterokaryon incompatibility factor HET-s/HET-s. Scientific reports 51 25500536
2010 6-(Het)aryl-7-deazapurine ribonucleosides as novel potent cytostatic agents. Journal of medicinal chemistry 49 19929004
1998 Microscopic and Ultrastructural Examination of Vegetative Incompatibility in Partial Diploids Heterozygous at het Loci in Neurospora crassa. Fungal genetics and biology : FG & B 49 9514694
2016 The HET-S/s Prion Motif in the Control of Programmed Cell Death. Cold Spring Harbor perspectives in biology 48 27352624
2005 Scaffold attachment factor SAFB1 suppresses estrogen receptor alpha-mediated transcription in part via interaction with nuclear receptor corepressor. Molecular endocrinology (Baltimore, Md.) 48 16195251
2003 Scaffold attachment factors SAFB1 and SAFB2: Innocent bystanders or critical players in breast tumorigenesis? Journal of cellular biochemistry 48 14587024
2001 In vivo aggregation of the HET-s prion protein of the fungus Podospora anserina. Molecular microbiology 48 11886562
1998 Deletion mapping of the head tilt (het) gene in mice: a vestibular mutation causing specific absence of otoliths. Genetics 48 9755211
2006 Nonallelic interactions between het-c and a polymorphic locus, pin-c, are essential for nonself recognition and programmed cell death in Neurospora crassa. Genetics 46 16554411
1994 Comparison of two active HeT-A retroposons of Drosophila melanogaster. Chromosoma 46 8055715
2010 Analyzing the birth and propagation of two distinct prions, [PSI+] and [Het-s](y), in yeast. Molecular biology of the cell 42 20219972
1997 Allelic specificity at the het-c heterokaryon incompatibility locus of Neurospora crassa is determined by a highly variable domain. Genetics 42 9258675
2016 The increasing diversity of functions attributed to the SAFB family of RNA-/DNA-binding proteins. The Biochemical journal 39 27888239
2011 Localization of HET-S to the cell periphery, not to [Het-s] aggregates, is associated with [Het-s]-HET-S toxicity. Molecular and cellular biology 37 22037764
2003 Glycolipid intermembrane transfer is accelerated by HET-C2, a filamentous fungus gene product involved in the cell-cell incompatibility response. Biochemistry 37 12525182
2010 A comparison of primary oesophageal squamous epithelial cells with HET-1A in organotypic culture. Biology of the cell 36 20843300
2010 SAFB1's multiple functions in biological control-lots still to be done! Journal of cellular biochemistry 35 20014070
2009 A combined solid-state NMR and MD characterization of the stability and dynamics of the HET-s(218-289) prion in its amyloid conformation. Chembiochem : a European journal of chemical biology 35 19504509
2009 SAFB1 mediates repression of immune regulators and apoptotic genes in breast cancer cells. The Journal of biological chemistry 35 19901029
2003 Conformational transition occurring upon amyloid aggregation of the HET-s prion protein of Podospora anserina analyzed by hydrogen/deuterium exchange and mass spectrometry. Biochemistry 35 12873146
2000 HET/SAF-B overexpression causes growth arrest and multinuclearity and is associated with aneuploidy in human breast cancer. Clinical cancer research : an official journal of the American Association for Cancer Research 35 10999774
2005 HeT-A and TART, two Drosophila retrotransposons with a bona fide role in chromosome structure for more than 60 million years. Cytogenetic and genome research 33 16093667
2004 Multiple roles of CLAN (caspase-associated recruitment domain, leucine-rich repeat, and NAIP CIIA HET-E, and TP1-containing protein) in the mammalian innate immune response. Journal of immunology (Baltimore, Md. : 1950) 33 15528373
2003 HeT-A elements in Drosophila virilis: retrotransposon telomeres are conserved across the Drosophila genus. Proceedings of the National Academy of Sciences of the United States of America 33 14614149
2007 The extracellular calcium-sensing receptor (CaSR) on human esophagus and evidence of expression of the CaSR on the esophageal epithelial cell line (HET-1A). American journal of physiology. Gastrointestinal and liver physiology 32 17962359
2017 Downregulation of SAFB Sustains the NF-κB Pathway by Targeting TAK1 during the Progression of Colorectal Cancer. Clinical cancer research : an official journal of the American Association for Cancer Research 31 28912140
2009 Low SAFB levels are associated with worse outcome in breast cancer patients. Breast cancer research and treatment 31 19137425
2004 Het-PDB Navi.: a database for protein-small molecule interactions. Journal of biochemistry 31 14999012
1999 Mutational analysis of the [Het-s] prion analog of Podospora anserina. A short N-terminal peptide allows prion propagation. Genetics 31 10581272
2020 Effect of different molecular weight PLGA on flurbiprofen nanoparticles: formulation, characterization, cytotoxicity, and in vivo anti-inflammatory effect by using HET-CAM assay. Drug development and industrial pharmacy 29 32281428
2009 Identification of the het-r vegetative incompatibility gene of Podospora anserina as a member of the fast evolving HNWD gene family. Current genetics 29 19137300
2004 The sequences appended to the amyloid core region of the HET-s prion protein determine higher-order aggregate organization in vivo. Journal of cell science 29 15159455
2002 Nonself recognition is mediated by HET-C heterocomplex formation during vegetative incompatibility. The EMBO journal 29 12234924
2017 Gluc-HET, a complementary chick embryo model for the characterization of antidiabetic compounds. PloS one 27 28777818
2015 The chromatin scaffold protein SAFB1 localizes SUMO-1 to the promoters of ribosomal protein genes to facilitate transcription initiation and splicing. Nucleic acids research 27 25800734
2014 Natural variation of heterokaryon incompatibility gene het-c in Podospora anserina reveals diversifying selection. Molecular biology and evolution 27 24448643
2013 The Scaffold attachment factor b1 (Safb1) regulates myogenic differentiation by facilitating the transition of myogenic gene chromatin from a repressed to an activated state. Nucleic acids research 27 23609547
2008 Flexible heteroarotinoid (Flex-Het) SHetA2 inhibits angiogenesis in vitro and in vivo. Investigational new drugs 27 18802666
2015 iCLIP identifies novel roles for SAFB1 in regulating RNA processing and neuronal function. BMC biology 26 26694817
2008 Identification of proteins binding to E-Box/Ku86 sites and function of the tumor suppressor SAFB1 in transcriptional regulation of the human xanthine oxidoreductase gene. The Journal of biological chemistry 26 18772145
1997 Reactivity in vegetative incompatibility of the HET-E protein of the fungus Podospora anserina is dependent on GTP-binding activity and a WD40 repeated domain. Molecular & general genetics : MGG 26 9435787
2005 Heterochromatic distribution of HeT-A- and TART-like sequences in several Drosophila species. Cytogenetic and genome research 25 16093664
2017 Partially-deuterated samples of HET-s(218-289) fibrils: assignment and deuterium isotope effect. Journal of biomolecular NMR 24 28074361
2012 Degradation of fungal prion HET-s(218-289) induces formation of a generic amyloid fold. Biophysical journal 24 22677387
1996 Escape from het-6 incompatibility in Neurospora crassa partial diploids involves preferential deletion within the ectopic segment. Genetics 24 8889517
1994 Drosophila telomere transposon HeT-A produces a transcript with tightly bound protein. Proceedings of the National Academy of Sciences of the United States of America 24 7517558
2013 Distribution and evolution of het gene homologs in the basidiomycota. Fungal genetics and biology : FG & B 23 24380733
2012 SAFB1- and SAFB2-mediated transcriptional repression: relevance to cancer. Biochemical Society transactions 23 22817742
2010 Structural determination and tryptophan fluorescence of heterokaryon incompatibility C2 protein (HET-C2), a fungal glycolipid transfer protein (GLTP), provide novel insights into glycolipid specificity and membrane interaction by the GLTP fold. The Journal of biological chemistry 23 20164530
2002 Some new methodological aspects of the hen's egg test for micronucleus induction (HET-MN). Mutation research 23 11815245
2000 Molecular and functional analyses of incompatibility genes at het-6 in a population of Neurospora crassa. Fungal genetics and biology : FG & B 23 11035941
1993 The genomic organization of HeT-A retroposons in Drosophila melanogaster. Chromosoma 23 8391971
2015 Theme and variations: evolutionary diversification of the HET-s functional amyloid motif. Scientific reports 22 26219477
2014 Structural basis for inhibition of mycobacterial and human adenosine kinase by 7-substituted 7-(Het)aryl-7-deazaadenine ribonucleosides. Journal of medicinal chemistry 22 25259627
2004 Expression of the telomeric retrotransposon HeT-A in Drosophila melanogaster is correlated with cell proliferation. Development genes and evolution 22 15069641
2017 Chaperonin GroEL accelerates protofibril formation and decorates fibrils of the Het-s prion protein. Proceedings of the National Academy of Sciences of the United States of America 21 28784759
2015 Ochratoxin A induces DNA damage and G2 phase arrest in human esophageal epithelium Het-1A cells in vitro. The Journal of toxicological sciences 21 26354382
2020 Synergistic Carcinogenesis of HPV18 and MNNG in Het-1A Cells through p62-KEAP1-NRF2 and PI3K/AKT/mTOR Pathway. Oxidative medicine and cellular longevity 20 33123313
2016 Telomeric Retrotransposon HeT-A Contains a Bidirectional Promoter that Initiates Divergent Transcription of piRNA Precursors in Drosophila Germline. Journal of molecular biology 19 27939293
2007 SAFB re-distribution marks steps of the apoptotic process. Experimental cell research 19 17643427

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