Affinage

STAG1

Protein TMEPAI · UniProt Q969W9

Length
287 aa
Mass
31.6 kDa
Annotated
2026-06-10
94 papers in source corpus 24 papers cited in narrative 26 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

STAG1 (SA1) is a core HEAT-repeat subunit of the cohesin complex that defines a functionally distinct variant, cohesin-SA1, mechanistically separable from the paralogous cohesin-SA2 (PMID:19822671, PMID:29867216). STAG1 binds tightly to the SMC1/SMC3/SCC1(RAD21) trimer to form a tetramer that cooperatively engages DNA, and an ATP-modulated DNA-binding site created at this interface supports the repeated DNA grabbing-and-release cycles of loop extrusion (PMID:40763028); the STAG1–RAD21 interface has been resolved crystallographically and depends on defined SA-binding motifs on RAD21 (PMID:23874961, PMID:32467316). A unique N-terminal AT-hook motif absent from SA2 lets STAG1 bind AT-rich and telomeric DNA directly and drives efficient loop extrusion and stable G1 chromatin association without changing ATPase activity (PMID:23729739). Through these properties cohesin-SA1 preferentially accumulates at CTCF sites and gene promoters to stabilize TAD boundaries and long-lived chromatin loops, a role SA2-cohesin cannot replace (PMID:22415368, PMID:29867216, PMID:32065581); ESCO1-mediated acetylation and CTCF protect cohesin-SA1 from WAPL eviction, conferring hours-long chromatin residence in G1 (PMID:32065581). Functionally, STAG1 is specifically required for sister telomere cohesion—not centromere cohesion—via its AT-hook and cooperation with the shelterin proteins TRF1 and TIN2 to pair and compact telomeric DNA (PMID:19822671, PMID:23729739, PMID:27298259, PMID:37081787), and SA1-null cells show telomere replication defects, aneuploidy, and altered transcription of Cornelia de Lange-associated genes (PMID:22415365, PMID:22415368). Loss of STAG1 abrogates residual sister chromatid cohesion selectively in STAG2-mutant cells, establishing a synthetic-lethal vulnerability exploitable in cancer (PMID:28691904, PMID:28430577, PMID:32467316). STAG1 haploinsufficiency, via nonsense-mediated decay of a frameshift allele, is a pathogenic mechanism underlying the intellectual disability disorder MRD47 (PMID:41210242).

Mechanistic history

Synthesis pass · year-by-year structured walk · 12 steps
  1. 1997 Medium

    Established STAG1 as a discrete nuclear protein and gene, providing the molecular entity later placed within cohesin.

    Evidence cDNA cloning with subcellular fractionation and immunoprecipitation/Western blot

    PMID:9305759

    Open questions at the time
    • No functional or complex assignment
    • Discrepancy between ubiquitous mRNA and hematopoietic-restricted protein left unexplained
  2. 2009 High

    Resolved whether the two cohesin SA paralogs are functionally redundant by showing SA1 governs telomere cohesion while SA2 governs centromere cohesion, defining variant-specific roles.

    Evidence siRNA depletion of SA1 vs SA2 in human cells with cytological cohesion analysis

    PMID:19822671

    Open questions at the time
    • Molecular basis of telomere specificity not yet identified
    • No structural detail on SA1-chromatin engagement
  3. 2012 High

    Defined the in vivo consequences of SA1 loss and its genome-wide chromatin role, linking SA1 to telomere replication, CTCF/promoter occupancy, and Cornelia de Lange-associated transcription.

    Evidence SA1 knockout mouse, MEF cytogenetics/FISH, cohesin ChIP-seq and RNA-seq; separate NLS/NES mapping in yeast and HeLa

    PMID:22415365 PMID:22415368 PMID:22715410

    Open questions at the time
    • Direct DNA-binding determinant not yet identified
    • Mechanism connecting cohesin distribution to specific gene regulation incomplete
  4. 2013 High

    Identified the molecular determinants of SA1 function: a unique AT-hook for direct telomeric DNA binding, defined RAD21-binding motifs, and a C-terminal region dictating damage-site recruitment versus SA2.

    Evidence Domain mutagenesis and ChIP for the AT-hook; in vitro binding, AUC and rescue for RAD21 motifs; chimeric SA1/SA2 constructs and laser micro-irradiation for DSB recruitment

    PMID:23729739 PMID:23874961 PMID:24324008

    Open questions at the time
    • AT-hook DNA-binding dynamics not yet resolved at single-molecule level
    • How AT-hook contribution differs from cohesin ring entrapment unclear
  5. 2016 High

    Provided biophysical and chromatin-level mechanism: single-molecule imaging showed SA1 searches DNA by 1D diffusion and is tethered at telomeres by TRF1, and SA1 modulates higher-order chromatin compaction relevant to tumor suppression.

    Evidence DNA tightrope/single-molecule imaging and AFM with AT-hook mutants; MNase accessibility, nanocytology and proliferation assays in HT29 cells

    PMID:27298259 PMID:27549371

    Open questions at the time
    • In vitro telomere pairing not yet linked to cellular cohesion quantitatively
    • Tumor suppressor mechanism in colorectal cells limited to single line
  6. 2017 High

    Demonstrated a therapeutically relevant synthetic-lethal dependency: STAG1 loss selectively kills STAG2-mutant cells by abrogating residual cohesion, reversible by STAG2 restoration.

    Evidence RNAi/CRISPR STAG1 inactivation in isogenic STAG2-WT vs mutant lines, metaphase spreads, viability and rescue assays

    PMID:28430577 PMID:28691904

    Open questions at the time
    • Druggable interface not yet defined
    • Breadth across cancer genotypes incompletely mapped
  7. 2018 High

    Separated the genome-organization functions of the two variants, assigning cohesin-SA1 to CTCF-anchored TAD boundaries and cohesin-SA2 to CTCF-independent enhancer-promoter contacts.

    Evidence ChIP-seq and Hi-C with individual SA1 or SA2 depletion in human cells

    PMID:29867216

    Open questions at the time
    • Mechanism of SA1 preference for CTCF sites not yet molecular
    • Polycomb-related contributions addressed only later
  8. 2019 High

    Extended variant-specific roles to development: SA1/SA2 cooperate in HSPC genome occupancy and segregation, while SA1 specifically preserves TAD borders and limits Polycomb domain interactions in stem cells.

    Evidence ChIP-seq/ATAC-seq and conditional single/double knockout mouse models in HSPCs; Hi-C/ChIP-seq in mESCs

    PMID:31216471 PMID:31495782

    Open questions at the time
    • Direct biochemical basis of SA1 versus SA2 boundary preference still unresolved
    • Polycomb antagonism mechanism not fully defined
  9. 2020 High

    Mechanistically explained SA1's preferential boundary role through chromatin kinetics (hours-long acetylation/CTCF/ESCO1-protected residence), defined RNA/R-loop binding by both paralogs, and crystallized the STAG1–RAD21 interface underpinning the synthetic-lethal dependency.

    Evidence FRAP, degron, ChIP-seq, Hi-C for residence/acetylation; EMSA/AFM/tightrope and DRIP-seq for RNA binding; X-ray crystallography plus degron/CRISPR for STAG1–RAD21

    PMID:32065581 PMID:32352519 PMID:32467316

    Open questions at the time
    • Functional consequence of SA1 RNA/R-loop binding in vivo unclear
    • How acetylation status is set specifically on SA1-cohesin not resolved
  10. 2023 Medium

    Defined SA1's telomere compaction biochemistry with TRF1/TIN2 and its developmental role at the nucleolus, showing the N-terminus regulates rRNA/repeat expression and pluripotency in mESCs.

    Evidence Nanochannel DNA stretching and AFM for TRF1-TIN2-SA1 compaction; STAG1 depletion with N-terminal rescue, RNA-seq and nucleolar imaging in mESCs

    PMID:37081787 PMID:37802073

    Open questions at the time
    • Telomere compaction reconstituted in vitro, not in cells
    • Link between nucleolar/repeat regulation and cohesin activity unestablished
  11. 2024 Medium

    Advanced toward pharmacology and structural mechanism: a small molecule (KPT-6566) directly disrupts STAG1/STAG2 binding to SCC1 and DNA, and simulations predict STAG1 DNA-clamping patches contributing to loop extrusion.

    Evidence FP-based screening and biophysical binding plus chromosome/γH2AX and synergy assays for KPT-6566; molecular dynamics simulations for DNA-binding patches

    PMID:39541712

    Open questions at the time
    • KPT-6566 selectivity between paralogs not established
    • MD-predicted clamping patches lack experimental validation
  12. 2025 High

    Provided quantitative biochemical and physiological mechanism: STAG1 builds an ATP-modulated DNA-binding site within the SMC1/SMC3/SCC1 tetramer for extrusion, the AT-hook enhances extrusion efficiency and G1 chromatin retention, and STAG1 haploinsufficiency via NMD causes MRD47.

    Evidence Mass photometry of cohesin subcomplexes with mutants; single-molecule loop extrusion comparing SA1 vs SA2 with AT-hook mutants (preprint); mutant expression with RT-qPCR/Western for the MRD47 frameshift variant

    PMID:40763028 PMID:41210242

    Open questions at the time
    • AT-hook extrusion data from preprint single-molecule assay
    • How CTCF asymmetrically halts SA1 extrusion still being mapped (preprint)

Open questions

Synthesis pass · forward-looking unresolved questions
  • How cohesin-SA1's biochemical features (AT-hook DNA binding, ATP-modulated DNA site, RNA/R-loop binding, acetylation-dependent residence) are integrated and selectively regulated to specify telomere cohesion, TAD boundary maintenance, and developmental gene control remains unresolved.
  • No unified structure of chromatin-engaged cohesin-SA1 during extrusion
  • Functional role of SA1 RNA/R-loop binding in vivo undefined
  • Mechanism linking nucleolar/repeat regulation to cohesin activity unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003677 DNA binding 4 GO:0005198 structural molecule activity 2 GO:0060090 molecular adaptor activity 2 GO:0003723 RNA binding 1
Localization
GO:0000228 nuclear chromosome 3 GO:0005634 nucleus 2 GO:0005730 nucleolus 1
Pathway
R-HSA-1266738 Developmental Biology 3 R-HSA-1640170 Cell Cycle 3 R-HSA-4839726 Chromatin organization 3 R-HSA-74160 Gene expression (Transcription) 2 R-HSA-73894 DNA Repair 1
Partners
CTCFESCO1NIPBLRAD21SMC1ASMC3TIN2TRF1
Complex memberships
cohesin complex (cohesin-SA1)

Evidence

Reading pass · 26 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2009 Cohesin-SA1 is specifically required for telomere cohesion in human cells, whereas cohesin-SA2 is required for centromere cohesion. Cells depleted of SA1 fail to establish or maintain cohesion between sister telomeres after DNA replication in S phase, leading to inability to repair chromatid breaks and sister telomere loss. siRNA depletion of SA1 or SA2 in human cells with cytological analysis of telomere and centromere cohesion The Journal of cell biology High 19822671
2012 SA1-null mouse embryonic fibroblasts show defective telomere replication (not centromeric cohesion) leading to chromosome segregation defects and aneuploidy. Complete SA1 ablation causes embryonic lethality; heterozygous mice show shorter lifespan and earlier tumourigenesis. SA1 knockout mouse model; analysis of MEFs by FISH, cytogenetics, and BrdU incorporation; telomere FISH The EMBO journal High 22415365
2012 SA1 is largely responsible for cohesin accumulation at gene promoters and CTCF-binding sites in mouse cells. Loss of SA1 alters transcription of genes related to Cornelia de Lange syndrome, including myc and protocadherin genes, a function that cannot be assumed by cohesin-SA2. ChIP-seq of cohesin in wild-type vs SA1-null MEFs; RNA-seq; comparison of genome-wide cohesin distribution The EMBO journal High 22415368
1997 SA1 (stromal antigen 1) is a nuclear protein of ~120 kDa encoded by a novel gene family. It is ubiquitously expressed at the mRNA level but the protein is predominantly detected in hematopoietic organs, suggesting post-transcriptional regulation. Molecular cloning from mouse/human cDNA libraries; subcellular fractionation; immunoprecipitation + Western blot with polyclonal antiserum Gene Medium 9305759
2013 SA1 binds directly to telomeric DNA through a unique AT-hook motif in its N-terminal domain (not present in SA2). Overexpression of SA1 alone is sufficient to induce telomere cohesion independent of the cohesin ring, dependent on this AT-hook. SA1 is highly enriched at telomeres by ChIP and enrichment decreases at mitosis when cohesion is resolved. ChIP of SA1 at telomeres; overexpression of SA1 truncation/domain mutants; FISH-based cohesion assay; identification of AT-hook by sequence analysis and mutagenesis Journal of cell science High 23729739
2013 Cohesin-SA2, not cohesin-SA1, is the main complex recruited to DNA double-strand break sites in S/G2 phase, co-recruited with the cohesin-loading factor NIPBL. Replacing the diverged C-terminal region of SA1 with the corresponding SA2 region confers damage-site recruitment on SA1. SA2 but not SA1 depletion decreases sister chromatid homologous recombination repair and affects repair pathway choice. Both SA1 and SA2 function in the intra-S checkpoint. Laser micro-irradiation with live-cell imaging; siRNA depletion; SA1/SA2 chimeric protein constructs; HR repair assay Molecular and cellular biology High 24324008
2013 SA1/SA2 interact with Rad21 through two SA-binding motifs on Rad21: an N-terminal motif (residues 60–81) and a middle-part 10-amino-acid α-helical motif (residues 383–392). Mutation of conserved residues L385, F389, T390 in the middle motif significantly hinders Rad21 from binding SA1/2. SA2 interacts with Rad21 through a broad region (301–750 aa). Mutant Rad21 fails to rescue precocious chromosome separation caused by Rad21 depletion. In vitro binding assays; co-immunoprecipitation; analytical ultracentrifugation; site-directed mutagenesis; chromosome segregation rescue assay PloS one High 23874961
2016 SA1 uses two-state binding on DNA: 1D free diffusion for searching and subdiffusive sliding for recognition at telomeric regions. The AT-hook motif modulates both non-specific DNA binding and subdiffusive dynamics at telomeric sequences. TRF1 tethers SA1 within telomeric regions. SA1 and TRF1 together form longer DNA-DNA pairing tracts than TRF1 alone. Single-molecule fluorescence imaging; DNA tightrope assay; atomic force microscopy; SA1 AT-hook mutant analysis Nucleic acids research High 27298259
2017 STAG1 loss abrogates sister chromatid cohesion specifically in STAG2-mutated cells, leading to mitotic catastrophe, defective cell division, and apoptosis — demonstrating a synthetic lethal interaction between STAG1 and STAG2. Restoration of STAG2 expression in a STAG2-mutated cell line alleviates STAG1 dependency. RNAi/CRISPR-mediated STAG1 inactivation in isogenic STAG2-wildtype vs. mutant cell lines; metaphase spread analysis; cell viability assays; STAG2 re-expression rescue eLife High 28430577 28691904
2018 Cohesin-SA1 preferentially stabilizes topologically associating domain (TAD) boundaries together with CTCF, while cohesin-SA2 promotes cell-type-specific contacts between enhancers and promoters independently of CTCF. Both variants are found at CTCF sites, but SA2-cohesin also localizes to enhancers lacking CTCF, which SA1-cohesin cannot replace when SA2 is absent. ChIP-seq and Hi-C in human cells with SA1 or SA2 downregulation; comparison of genomic distributions; RNA-seq Nature structural & molecular biology High 29867216
2019 Stag2 and Stag1 bind a shared set of genomic loci in hematopoietic stem and progenitor cells, but a component of Stag2 binding sites is unoccupied by Stag1 even in Stag2-deficient cells. Stag2 loss alone decreases chromatin accessibility and transcription of lineage-specification genes; concurrent loss of both Stag2 and Stag1 abrogates hematopoiesis. Both share a role in chromosomal segregation. ChIP-seq for Stag1 and Stag2 in HSPCs; ATAC-seq; conditional Stag2 knockout and Stag1/Stag2 double knockout mouse models Cell stem cell High 31495782
2019 In mouse embryonic stem cells, cohesin-SA1 preserves TAD borders and disrupts Polycomb domain interaction networks, while cohesin-SA2 facilitates Polycomb domain compaction through PRC1 recruitment and promotes long-range interaction networks between Polycomb-bound promoters. Hi-C, ChIP-seq, and RNA-seq in mESCs with SA1 or SA2 depletion; comparison of 3D genome organization Cell reports High 31216471
2020 During G1 phase, acetylated cohesin-STAG1 binds chromatin for hours (long residence time), whereas cohesin-STAG2 binds for minutes. CTCF and the acetyltransferase ESCO1 protect a subset of cohesin-STAG1 from WAPL-mediated removal, enabling formation of long and long-lived chromatin loops. ESCO1 contributes to boundary formation in chromatin looping similarly to CTCF. FRAP, ChIP-seq, Hi-C, auxin-inducible degron system for ESCO1/WAPL; acetylation analysis; live-cell imaging eLife High 32065581
2020 SA1 (STAG1) and SA2 (STAG2) directly bind RNA, including ssRNA, dsRNA, RNA:DNA hybrids, and R-loops, using bulk biochemical assays and single-molecule techniques. SA1 and SA2 localize to regions on dsDNA that contain RNA; SA1/SA2 binding sites significantly overlap with R-loop sites genome-wide by ChIP-seq/DRIP-seq comparison. Electrophoretic mobility shift assay (EMSA), atomic force microscopy, DNA tightrope assay with RNA substrates; ChIP-seq and DRIP-seq data comparison Nucleic acids research High 32352519
2020 STAG1 is required for sister chromatid cohesion specifically in STAG2-deficient cells. Chemical-genetic degradation of STAG1 protein (inducible degron) results in loss of sister chromatid cohesion and rapid cell death selectively in STAG2-deficient cells. X-ray crystallography identifies STAG1 regions that interact with the RAD21 subunit; STAG1 mutations abrogating this interaction selectively compromise viability of STAG2-deficient cells. Genome-wide CRISPR screens; auxin-inducible degron for STAG1 degradation; X-ray crystallography of STAG1–RAD21 interaction domain; metaphase spread assay; biochemical interaction assays Life science alliance High 32467316
2012 SA1 and SA2 contain nuclear localization signals (NLS) and export signals (NES). Only the N-terminal NLS of SA1 is functional for nuclear import (validated in both yeast and HeLa cells), whereas SA2 has at least two functional NLS and two functional NES allowing nucleocytoplasmic shuttling. This represents a difference in nuclear transport organization between the two paralogs. Expression of fluorescently tagged SA1/SA2 in S. cerevisiae and HeLa cells; deletion/mutation analysis of putative NLS and NES sequences; fluorescence microscopy PloS one Medium 22715410
2023 TRF1, TIN2, and SA1 work synergistically to compact telomeric DNA. SA1 alone can drive DNA compaction. In the ternary TRF1-TIN2-SA1 system, compaction is an additive effect of TRF1-TIN2 and SA1-based compaction pathways. Compaction by TRF1 and TIN2 proceeds via TRF1 binding followed by TIN2 recognition of TRF1. Nanochannel DNA stretching to quantify compaction; atomic force microscopy of intermolecular aggregation; systematic comparison of protein combinations Biophysical journal Medium 37081787
2023 STAG1 is the dominant paralog in mouse embryonic stem cells and is required for pluripotency. The N-terminus of STAG1 specifically regulates repeat (rRNA) expression, nucleolar integrity, and repression of the two-cell (2C) state to maintain mESC identity. Skewing the balance of naturally occurring Stag1 isoforms impacts cell identity. STAG1 depletion in mESCs; rescue with N-terminal domain constructs; RNA-seq for repeat expression; immunofluorescence of nucleolar markers; STAG1 isoform characterization Stem cell reports Medium 37802073
2025 STAG1 binds tightly to the trimeric SMC1/SMC3/SCC1 complex; this tetramer then weakly but cooperatively binds DNA. Full-length NIPBL acts as a DNA anchor during loop extrusion. An ATP-modulated DNA binding site created by STAG1's interaction with SMC1/SMC3/SCC1 is important for repeated grabbing and release of DNA critical to extrusion. Cohesin mutants lacking DNA-binding domains in ATPase heads show negligible differences in overall DNA affinity. Mass photometry to quantify biomolecular interactions; systematic binding measurements of cohesin subcomplexes; ATPase-dead and head-engagement mutants Proceedings of the National Academy of Sciences of the United States of America High 40763028
2025 Cohesin-STAG1 extrudes DNA loops more efficiently than cohesin-STAG2 despite comparable ATPase activity and topological DNA entrapment. The AT-hook motif unique to the STAG1 N-terminus promotes loop extrusion without altering ATPase activity or DNA binding. In human somatic cells, the AT-hook is required for stable cohesin-chromatin association during G1 phase but is dispensable for sister chromatid cohesion. Mutation of the AT-hook markedly impairs TAD and chromatin loop formation. Single-molecule loop extrusion assay comparing cohesin-STAG1 vs cohesin-STAG2; AT-hook mutant constructs; ChIP-seq; Hi-C; ATPase assay; cohesion assay in human cells bioRxivpreprint High
2024 KPT-6566 directly binds to both STAG1 and STAG2, disrupting their interactions with SCC1 and double-stranded DNA. KPT-6566 causes premature chromosome separation and chromosome damage in HeLa cells, impairs DNA damage repair, and accumulates DSBs. KPT-6566 sensitizes cells to PARP inhibitor Olaparib and NHEJ inhibitor UMI-77. Fluorescence polarization-based high-throughput screening; biochemical and biophysical binding assays; metaphase chromosome spread; γH2AX foci assay; cell viability synergy assay DNA repair Medium 39541712
2025 The CTCF N-terminal region (NTR) contains two motifs, YDF and KTYQR, that hinder cohesin-mediated loop extrusion. YDF hinders loop extrusion step cycles and converts cohesin into a unidirectional extruder by strengthening STAG1's affinity to DNA. KTYQR fully impedes loop extrusion activity. Single-molecule DNA loop extrusion assay with CTCF NTR fragments and motif mutants; measurement of STAG1-DNA affinity bioRxivpreprint Medium
2024 Molecular dynamics simulations predict DNA binding patches on STAG1 that, together with patches on SMC1, SMC3, and NIPBL, form a DNA clamping patch group. This group is predicted to facilitate DNA bending and capture of DNA within the cohesin ring, critical steps in loop extrusion. Molecular dynamics simulations with amino-acid-residue resolution structural models of cohesin subunits including STAG1; quantification of DNA-binding affinities via dissociation rate constants bioRxivpreprint Low
2020 In zebrafish, stag1a (but not stag1b) contributes to primitive embryonic haematopoiesis. Loss of stag1a expands the lateral plate mesoderm zone of scl-positive haematopoietic/vascular progenitors, reduces kidney progenitors, and increases spi1-positive myeloid-biased cells, consistent with skewing toward primitive myelopoiesis. CRISPR-Cas9 zebrafish loss-of-function mutants; in situ hybridization for haematopoietic markers; phenotypic analysis at 24 hpf Frontiers in cell and developmental biology Medium 33365313
2016 SA1 modulates high-order chromatin structure in colorectal cancer cells. SA1 knockdown in HT29 cells increases chromatin accessibility (by MNase assay and nanocytology) and increases cell proliferation, supporting a tumor suppressor role through chromatin organization. siRNA knockdown of SA1 in HT29 cells; micrococcal nuclease (MNase) chromatin accessibility assay; nanocytology (biophotonic); WST-1 proliferation assay; colony formation assay Cancer prevention research Medium 27549371
2025 A frameshift STAG1 variant (c.500dup; p.Gly168TrpfsTer13) leads to dramatic reduction in mutant mRNA levels due to nonsense-mediated mRNA decay, resulting in impaired production of functional STAG1 protein. This provides functional evidence that STAG1 haploinsufficiency is the pathogenic mechanism underlying MRD47. Mutant expression vector transfection into HEK293T cells; RT-qPCR of mutant mRNA; Western blot of STAG1 protein; comparison with wild-type construct Frontiers in pediatrics Medium 41210242

Source papers

Stage 0 corpus · 94 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2012 Cohesin-SA1 deficiency drives aneuploidy and tumourigenesis in mice due to impaired replication of telomeres. The EMBO journal 150 22415365
2009 Differential regulation of telomere and centromere cohesion by the Scc3 homologues SA1 and SA2, respectively, in human cells. The Journal of cell biology 140 19822671
2020 N6-Methyladenosine Modification of Fatty Acid Amide Hydrolase Messenger RNA in Circular RNA STAG1-Regulated Astrocyte Dysfunction and Depressive-like Behaviors. Biological psychiatry 138 32387133
2020 ESCO1 and CTCF enable formation of long chromatin loops by protecting cohesinSTAG1 from WAPL. eLife 134 32065581
2019 Cohesin Members Stag1 and Stag2 Display Distinct Roles in Chromatin Accessibility and Topological Control of HSC Self-Renewal and Differentiation. Cell stem cell 133 31495782
2018 Distinct roles of cohesin-SA1 and cohesin-SA2 in 3D chromosome organization. Nature structural & molecular biology 131 29867216
2012 A unique role of cohesin-SA1 in gene regulation and development. The EMBO journal 113 22415368
2020 Thermotolerance effect of plant growth-promoting Bacillus cereus SA1 on soybean during heat stress. BMC microbiology 103 32571217
2017 Synthetic lethality between the cohesin subunits STAG1 and STAG2 in diverse cancer contexts. eLife 94 28691904
2019 Specific Contributions of Cohesin-SA1 and Cohesin-SA2 to TADs and Polycomb Domains in Embryonic Stem Cells. Cell reports 77 31216471
2013 Distinct functions of human cohesin-SA1 and cohesin-SA2 in double-strand break repair. Molecular and cellular biology 75 24324008
1988 Purification, molecular cloning, and sequencing of salivary cystatin SA-1. The Journal of biological chemistry 75 2837486
2001 Characterization of a novel gene, STAG1/PMEPA1, upregulated in renal cell carcinoma and other solid tumors. Molecular carcinogenesis 71 11568975
2003 EGF- and cell-cycle-regulated STAG1/PMEPA1/ERG1.2 belongs to a conserved gene family and is overexpressed and amplified in breast and ovarian cancer. Molecular carcinogenesis 64 14639658
1999 Increased platinum accumulation in SA-1 tumour cells after in vivo electrochemotherapy with cisplatin. British journal of cancer 61 10188880
2017 Synthetic lethal interaction between the tumour suppressor STAG2 and its paralog STAG1. Oncotarget 60 28430577
2010 A randomized study of pegylated liposomal doxorubicin versus continuous-infusion doxorubicin in elderly patients with acute lymphoblastic leukemia: the GRAALL-SA1 study. Haematologica 58 20971822
2008 Pediocin SA-1, an antimicrobial peptide from Pediococcus acidilactici NRRL B5627: production conditions, purification and characterization. Bioresource technology 58 18093831
2020 Specialized functions of cohesins STAG1 and STAG2 in 3D genome architecture. Current opinion in genetics & development 52 32294612
2020 Cohesin SA1 and SA2 are RNA binding proteins that localize to RNA containing regions on DNA. Nucleic acids research 48 32352519
2016 Preliminary treatment of bovine mastitis caused by Staphylococcus aureus, with trx-SA1, recombinant endolysin of S. aureus bacteriophage IME-SA1. Veterinary microbiology 48 27374909
2017 STAG1 mutations cause a novel cohesinopathy characterised by unspecific syndromic intellectual disability. Journal of medical genetics 47 28119487
2023 The synthetic TRPML1 agonist ML-SA1 rescues Alzheimer-related alterations of the endosomal-autophagic-lysosomal system. Journal of cell science 45 36825945
2020 Extending thermotolerance to tomato seedlings by inoculation with SA1 isolate of Bacillus cereus and comparison with exogenous humic acid application. PloS one 40 32353077
2013 Characterization of the interaction between the cohesin subunits Rad21 and SA1/2. PloS one 40 23874961
2004 Identification of STAG1 as a key mediator of a p53-dependent apoptotic pathway. Oncogene 40 15361841
2020 ML-SA1, a selective TRPML agonist, inhibits DENV2 and ZIKV by promoting lysosomal acidification and protease activity. Antiviral research 38 32858116
1997 SA-1, a nuclear protein encoded by one member of a novel gene family: molecular cloning and detection in hemopoietic organs. Gene 38 9305759
2013 Enhanced CO2 sequestration by a novel microalga: Scenedesmus obliquus SA1 isolated from bio-diversity hotspot region of Assam, India. Bioresource technology 37 23811524
2013 SA1 binds directly to DNA through its unique AT-hook to promote sister chromatid cohesion at telomeres. Journal of cell science 34 23729739
1991 Induction of systemic lupus erythematosus in naive mice with T-cell lines specific for human anti-DNA antibody SA-1 (16/6 Id+) and for mouse tuberculosis antibody TB/68 (16/6 Id+). Clinical immunology and immunopathology 34 1830833
2014 Differential mechanisms of action of the mucolipin synthetic agonist, ML-SA1, on insect TRPML and mammalian TRPML1. Cell calcium 31 25266962
2020 STAG1 vulnerabilities for exploiting cohesin synthetic lethality in STAG2-deficient cancers. Life science alliance 29 32467316
2006 The structures of glycolipids isolated from the highly thermophilic bacterium Thermus thermophilus Samu-SA1. Glycobiology 29 16636007
2018 The genome of 'Candidatus Phytoplasma solani' strain SA-1 is highly dynamic and prone to adopting foreign sequences. Systematic and applied microbiology 28 30455068
2014 CO2 biofixation and carbonic anhydrase activity in Scenedesmus obliquus SA1 cultivated in large scale open system. Bioresource technology 28 24865325
2004 Setting optimal parameters for in vitro electrotransfection of B16F1, SA1, LPB, SCK, L929 and CHO cells using predefined exponentially decaying electric pulses. Bioelectrochemistry (Amsterdam, Netherlands) 27 14990328
2004 NT3 expressed in skin causes enhancement of SA1 sensory neurons that leads to postnatal enhancement of Merkel cells. The Journal of comparative neurology 26 14991566
2016 Functional interplay between SA1 and TRF1 in telomeric DNA binding and DNA-DNA pairing. Nucleic acids research 25 27298259
2012 Autoimmunity and cystatin SA1 deficiency behind chronic mucocutaneous candidiasis in autoimmune polyendocrine syndrome type 1. Journal of autoimmunity 20 23122533
2018 SA1/SA2 cohesion proteins and SIRT1-NAD+ deacetylase modulate telomere homeostasis in cumulus cells and are eligible biomarkers of ovarian aging. Human reproduction (Oxford, England) 19 29481647
1978 A continuous culture study of the regulation of extracellular protease production in Vibrio SA1. Antonie van Leeuwenhoek 19 582093
2016 Complete genome sequence of Novosphingobium resinovorum SA1, a versatile xenobiotic-degrading bacterium capable of utilizing sulfanilic acid. Journal of biotechnology 17 27851894
2024 Jianpi qingre tongluo prescription alleviates the senescence-associated secretory phenotype with osteoarthritis by regulating STAG1/TP53/P21 signaling pathway. Journal of ethnopharmacology 15 39423944
2021 ML-SA1 and SN-2 inhibit endocytosed viruses through regulating TRPML channel expression and activity. Antiviral research 15 34687820
2013 Comparative phenotypic analysis and genome sequence of Clostridium beijerinckii SA-1, an offspring of NCIMB 8052. Microbiology (Reading, England) 15 24068240
2022 Characterization and Genomic Analysis of a Novel Jumbo Bacteriophage vB_StaM_SA1 Infecting Staphylococcus aureus With Two Lysins. Frontiers in microbiology 14 35572667
2021 Degradation of salicylic acid to catechol in Solanaceae by SA 1-hydroxylase. Plant physiology 14 33793924
2021 Novel STAG1 Frameshift Mutation in a Patient Affected by a Syndromic Form of Neurodevelopmental Disorder. Genes 14 34440290
2024 Characterization of bacteriophage vB_AbaS_SA1 and its synergistic effects with antibiotics against clinical multidrug-resistant Acinetobacter baumannii isolates. Pathogens and disease 13 39435653
2020 Cohesin Components Stag1 and Stag2 Differentially Influence Haematopoietic Mesoderm Development in Zebrafish Embryos. Frontiers in cell and developmental biology 13 33365313
2018 Serpin A1 and the modulation of type I collagen turnover: Effect of the C-terminal peptide 409-418 (SA1-III) in human dermal fibroblasts. Cell biology international 12 29908000
2007 High level expression of STAG1/PMEPA1 in an androgen-independent prostate cancer PC3 subclone. Cellular & molecular biology letters 12 17318295
2017 Starvation- and xenobiotic-related transcriptomic responses of the sulfanilic acid-degrading bacterium, Novosphingobium resinovorum SA1. Applied microbiology and biotechnology 11 29051988
2016 Higher Order Chromatin Modulator Cohesin SA1 Is an Early Biomarker for Colon Carcinogenesis: Race-Specific Implications. Cancer prevention research (Philadelphia, Pa.) 11 27549371
1991 Presence of a 16/6 related human anti-DNA common idiotype (SA1) in the anticardiolipin antibodies of patients with primary antiphospholipid syndrome. The Journal of rheumatology 11 1765979
2016 Cohesin subunits, STAG1 and STAG2, and cohesin regulatory factor, PDS5b, in oral squamous cells carcinomas. Journal of oral pathology & medicine : official publication of the International Association of Oral Pathologists and the American Academy of Oral Pathology 9 27341316
1998 Association of 16/6 and SA1 anti-DNA idiotypes with anticardiolipin antibodies and clinical manifestations in a large cohort of SLE patients. European Concerted Action on the Immunogenetics of SLE. Clinical and experimental rheumatology 8 9844765
2024 A Novel De Novo STAG1 Variant in Monozygotic Twins with Neurodevelopmental Disorder: New Insights in Clinical Heterogeneity. Genes 7 39336775
2012 Nuclear import and export signals of human cohesins SA1/STAG1 and SA2/STAG2 expressed in Saccharomyces cerevisiae. PloS one 7 22715410
1993 SA-1 antigen expression in small intensely fluorescent cells is associated with proliferation. Developmental biology 6 8099047
2024 A novel STAG1 variant associated with congenital clubfoot and microphthalmia: A case report. SAGE open medical case reports 5 39224759
2023 The N-terminus of Stag1 is required to repress the 2C program by maintaining rRNA expression and nucleolar integrity. Stem cell reports 5 37802073
2007 Overlaps between the various biodegradation pathways in Sphingomonas subarctica SA1. Acta biologica Hungarica 5 18297793
2004 Biochemical and genetic characterization of a D(-)-3-hydroxybutyrate dehydrogenase from Acidovorax sp. strain SA1. Journal of bioscience and bioengineering 5 16233594
2003 Altered kinetic properties of tyrosine-183 to cysteine mutation in glutamine synthetase of anabaena variabilis strain SA1 is responsible for excretion of ammonium ion produced by nitrogenase. Current microbiology 5 12732949
2002 Cloning and sequencing of an intracellular D(-)-3-hydroxybutyrate oligomer hydrolase from Acidovorax sp. strain SA1 and purification of the enzyme. Current microbiology 5 12070691
2024 (S)-ML-SA1 Activates Autophagy via TRPML1-TFEB Pathway. Chembiochem : a European journal of chemical biology 4 38923811
2022 Interactions between Jumbo Phage SA1 and Staphylococcus: A Global Transcriptomic Analysis. Microorganisms 4 36014008
2018 High-Quality Draft Single-Cell Genome Sequence Belonging to the Archaeal Candidate Division SA1, Isolated from Nereus Deep in the Red Sea. Genome announcements 4 29748404
2018 Draft genome sequence of Colletotrichum sansevieriae Sa-1-2, the anthracnose pathogen of Sansevieria trifasciata. Data in brief 4 29900221
2007 [Isolation and characterization of an atrazine-degrading bacterium strain SA1]. Wei sheng wu xue bao = Acta microbiologica Sinica 4 17672324
2006 Prostate--specific G protein couple receptor genes and STAG1/PMEPA1 in peripheral blood from patients with prostatic cancer. International journal of immunopathology and pharmacology 4 17166409
1995 Effect of surface antigen-1 (SA-1) immune lymphocyte subsets and naive cell subsets in protecting scid mice from initial and persistent infection with Cryptosporidium parvum. Veterinary immunology and immunopathology 4 8533299
1994 Effects of vinblastine on cell membrane fluidity and the growth of SA-1 tumor in mice. Cancer letters 4 8187053
1978 Purification and some properties of two extracellular proteolytic enzymes produced by Vibrio SA1. Antonie van Leeuwenhoek 4 36029
2025 Neurological Disease Syndrome Caused by a STAG1 Gene Variant: A Case Report and Literature Review. International journal of developmental neuroscience : the official journal of the International Society for Developmental Neuroscience 3 40625213
2025 NIPBL and STAG1 enable loop extrusion by providing differential DNA-cohesin affinity. Proceedings of the National Academy of Sciences of the United States of America 3 40763028
2020 Chromosomal localization of PemIK toxin-antitoxin system results in the loss of toxicity - Characterization of pemIKSa1-Sp from Staphylococcus pseudintermedius. Microbiological research 3 32622987
2013 Complete Genome Sequence of Simiduia agarivorans SA1(T), a Marine Bacterium Able To Degrade a Variety of Polysaccharides. Genome announcements 3 23405302
2024 Discovery of KPT-6566 as STAG1/2 Inhibitor sensitizing PARP and NHEJ Inhibitors to suppress tumor cells growth in vitro. DNA repair 2 39541712
2023 Assembly path dependence of telomeric DNA compaction by TRF1, TIN2, and SA1. Biophysical journal 2 37081787
2025 STAG1 Disease, Central Precocious Puberty, and Bone Fragility-A Case Report. Diagnostics (Basel, Switzerland) 1 40361893
2025 Therapeutic potential of ECP-SA1: A novel lytic phage against multidrug-resistant Escherichia coli. Microbial pathogenesis 1 40769226
2025 A Novel Variant of STAG1 Gene and Literature Review. Annals of human genetics 1 40899458
2025 A novel frameshift STAG1 variant exhibiting haploinsufficiency due to the nonsense-mediated mRNA decay: a case report and literature review. Frontiers in pediatrics 1 41210242
2023 Periosteal Fasciitis With Unusual Radiologic Features Harboring a Novel STAG1::USP6 Fusion Gene. International journal of surgical pathology 1 36694417
2020 Crystal structure of the YoeBSa1-YefMSa1 complex from Staphylococcus aureus. Biochemical and biophysical research communications 1 32446378
1985 Expression of an antigen defined by a monoclonal antibody (SA-1) on normal and malignant cells. Scandinavian journal of haematology 1 4081640
2026 Tumor-suppressive activities of SA1/STAG2 and effects of PARP impairment during brain development. Disease models & mechanisms 0 41502392
2026 Analysis of STAG1 gene variants identified in a patient with intellectual disability and speech delay. Yi chuan = Hereditas 0 41669812
2026 STAG1: Bridging the Gap Between Cohesin Complex and Epigenetic Machinery. Genes 0 42074601
2025 The synthetic TRPML1 agonist ML-SA1 mitigates intracellular lipid accumulation induced by antipsychotics in vitro by stimulating release of extracellular microvesicles. Biochimica et biophysica acta. Molecular and cell biology of lipids 0 40222411
1989 [Immunochemical analysis of the products of translation of mRNA hybridized with the left HindIII-A-Sa1-fragment of vaccinia virus DNA]. Molekuliarnaia genetika, mikrobiologiia i virusologiia 0 2628751

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