Affinage

SATB2

DNA-binding protein SATB2 · UniProt Q9UPW6

Length
733 aa
Mass
82.6 kDa
Annotated
2026-04-28
100 papers in source corpus 29 papers cited in narrative 31 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

SATB2 is a nuclear matrix-attachment region (MAR) binding protein containing two CUT domains and a homeodomain that functions as a context-dependent transcriptional regulator and higher-order chromatin organizer across multiple developmental and tissue contexts. In osteoblast differentiation, SATB2 directly interacts with RUNX2 and ATF4 to activate bone-specific genes and independently upregulates Osterix, while its haploinsufficiency causes cleft palate (PMID:16751105, PMID:12915443, PMID:21385070). In the nervous system, SATB2 directs neocortical projection neuron identity by directly activating Fezf2 and Sox5 and repressing Ctip2, and in adult hippocampal neurons it binds coding and miRNA gene promoters to stabilize long-term potentiation and memory (PMID:26324926, PMID:27897969). SATB2 recruits chromatin-modifying co-activators (CBP/p300) or co-repressors (HDAC1, LSD1/CoREST, SWI/SNF) to target loci in a context-dependent manner, organizes chromatin loop architecture whose topology shifts developmentally, and is regulated post-translationally by ZFP451-mediated SUMOylation and caspase-7 cleavage, with additional roles in maintaining colonic stem cell identity through modulation of CDX2/HNF4A enhancer binding (PMID:33124191, PMID:30858153, PMID:34244292, PMID:35326417, PMID:34582804, PMID:36598364).

Mechanistic history

Synthesis pass · year-by-year structured walk · 17 steps
  1. 2003 High

    Identification of SATB2 as a CUT-domain/homeodomain MAR-binding protein at a cleft-palate translocation breakpoint established a new chromatin-associated factor with a clear developmental haploinsufficiency phenotype.

    Evidence FISH mapping of translocation breakpoints and gene structure analysis in patients with cleft palate, with expression validated by whole-mount in situ hybridization in mouse embryos

    PMID:12915443

    Open questions at the time
    • Transcriptional targets in palate unidentified
    • Mechanism of MAR binding by CUT domains unresolved at structural level
  2. 2006 High

    Knockout and genetic epistasis studies revealed SATB2 as a molecular node in osteoblast differentiation, showing it physically interacts with RUNX2 and ATF4 and directly represses Hoxa2 to drive skeletal development.

    Evidence Co-immunoprecipitation, reporter assays, Satb2 knockout mice, and double heterozygous (Satb2/Runx2, Satb2/Atf4) genetic epistasis

    PMID:16751105

    Open questions at the time
    • How SATB2 distinguishes activating versus repressive targets in bone unknown
    • Genome-wide binding sites in osteoblasts not mapped
  3. 2010 Medium

    Discovery that SATB2 physically binds ΔNp63α and enhances its transrepression at p53-family targets expanded SATB2's role beyond skeletal biology into epithelial transcriptional regulation and chemoresistance.

    Evidence Reciprocal co-immunoprecipitation, reporter assays for transrepression, RNAi knockdown with apoptosis readout in HNSCC cells

    PMID:20829881

    Open questions at the time
    • Structural basis of SATB2-p63 interaction unresolved
    • In vivo relevance in epithelial tissues not tested
  4. 2011 Medium

    Demonstration that SATB2 upregulates Osterix independently of Runx2 and that AEC-specific p63 mutations alter the SATB2-p63 interaction defined SATB2's independent and partner-specific transcriptional activities.

    Evidence Osterix promoter-luciferase in Runx2-null calvarial cells with in vivo bone defect model; co-IP and ChIP at perp promoter with domain-specific p63 mutation panel

    PMID:21385070 PMID:21965674

    Open questions at the time
    • Runx2-independent mechanism of Osterix activation not fully dissected
    • Whether SATB2-p63 interaction is relevant for craniofacial malformations not tested genetically
  5. 2011 Medium

    In vivo knockdown showed SATB2 controls dendritic arborization and soma spacing of cortical pyramidal neurons, establishing its neuronal morphogenesis function.

    Evidence In vivo RNAi knockdown in mouse cerebral cortex with live imaging and immunohistochemistry

    PMID:21885532

    Open questions at the time
    • Transcriptional targets underlying dendritic and spacing phenotype unknown
    • Cell-autonomous versus non-cell-autonomous mechanisms not fully separated
  6. 2012 Medium

    Genetic epistasis with CB1 cannabinoid receptor revealed that SATB2 directly represses the Ctip2 promoter, positioning it as a binary switch between callosal and subcerebral cortical neuron fates.

    Evidence CB1 knockout mouse models, Ctip2 promoter reporter assays, immunostaining, and genetic epistasis

    PMID:23175820

    Open questions at the time
    • Whether SATB2 binds Ctip2 promoter directly via ChIP not shown in this study
    • How CB1 signaling molecularly modifies SATB2 repression unclear
  7. 2014 High

    Conditional knockout established that SATB2 is required for both callosal and subcerebral projection neuron differentiation with layer-specific roles, resolving the apparent paradox of its repression of Ctip2 yet necessity for corticospinal tract formation.

    Evidence Conditional Satb2 knockout in developing neocortex with electrophysiology, axon tracing, and gene expression analysis

    PMID:25037921

    Open questions at the time
    • Molecular basis of layer-specific SATB2 activity not identified
    • Whether SATB2 partner complexes differ between deep and upper layers not tested
  8. 2015 High

    ChIP and reporter assays demonstrated SATB2 directly activates Fezf2 and Sox5, revealing the specific gene targets through which it promotes subcerebral identity while repressing the same in callosal neurons.

    Evidence ChIP showing direct SATB2 binding at Fezf2 and Sox5 loci, reporter assays, in utero electroporation, conditional knockout

    PMID:26324926

    Open questions at the time
    • How SATB2 switches from activation to repression at the same loci in different neuron types unresolved
    • Chromatin loop structures at these loci not examined
  9. 2017 Medium

    FRAP analysis of disease-associated missense variants in CUT1, CUT2, and inter-domain regions showed that each functional domain differentially controls SATB2's chromatin/matrix residence time, linking domain structure to nuclear dynamics.

    Evidence Fluorescently tagged SATB2 expressed in patient fibroblasts with FRAP measurements across a domain-specific mutation panel

    PMID:28151491

    Open questions at the time
    • How altered nuclear mobility translates to transcriptional output not measured
    • Structural basis for domain-specific mobility differences not resolved
  10. 2019 High

    Conditional knockout in adult forebrain proved SATB2 is required for LTP stabilization and long-term memory, and ChIP showed it directly binds promoters of synaptic plasticity-related coding genes and miRNAs in hippocampal neurons, extending its function beyond development to adult neural circuit maintenance.

    Evidence Conditional Satb2 knockout in adult forebrain, ChIP at promoters, miRNA profiling, LTP electrophysiology, behavioral fear conditioning and object discrimination

    PMID:27897969

    Open questions at the time
    • Specific miRNA targets mediating LTP stabilization not individually validated
    • Whether SATB2 organizes chromatin loops in adult neurons not tested
  11. 2019 Medium

    Mass-spectrometry interactomics revealed a developmental shift in SATB2 protein complexes from transcriptional repressor-enriched in neonatal cortex to chromatin-loop-stabilizer-enriched in adult cortex, providing a molecular basis for its context-dependent functions.

    Evidence Immunoprecipitation-mass spectrometry of SATB2 complexes from neonatal and adult mouse cortex

    PMID:30726206

    Open questions at the time
    • Individual interactors not validated by reciprocal co-IP
    • Whether complex composition is cause or consequence of chromatin state not tested
  12. 2019 High

    ChIP demonstrated SATB2 recruits HDAC1 to the Snail promoter to repress EMT, linking its chromatin-remodeling recruitment activity to epithelial-mesenchymal regulation in colorectal carcinoma.

    Evidence ChIP for SATB2 and HDAC1 at Snail promoter, RNA pulldown for SATB2-AS1/p300 scaffold, reporter assays, in vivo xenograft validation

    PMID:30858153

    Open questions at the time
    • Whether HDAC1 recruitment is direct or mediated by additional adaptor proteins unclear
    • Generalizability beyond CRC not established
  13. 2020 High

    Co-immunoprecipitation and ChIP-seq identified SATB2 recruitment of CBP to MAR sequences at the FOXM1 locus in glioma stem cells, establishing a specific mechanism by which SATB2 uses MAR-binding to deliver histone acetyltransferases to target genes.

    Evidence Co-IP of SATB2-CBP, ChIP at FOXM1 MAR, shRNA knockdown, in vivo xenograft, CBP inhibitor C646 treatment

    PMID:33124191

    Open questions at the time
    • How SATB2 selects specific MAR sequences genome-wide not addressed
    • Whether CBP recruitment is a general mechanism at all SATB2-activated loci unknown
  14. 2021 High

    Identification of ZFP451 as the E3 SUMO ligase that SUMOylates SATB2, and demonstration that SUMO2-SATB2 interacts with LSD1/CoREST to silence pluripotency genes and rewire chromatin loops, provided the first post-translational modification switch controlling SATB2's repressive versus activating modes.

    Evidence SUMO acceptor site mutagenesis, ZFP451 knockout, SUMO2-SATB2 fusion rescue, co-IP with LSD1/CoREST, Hi-C and ChIP-seq in ESCs

    PMID:34244292

    Open questions at the time
    • Whether SUMOylation controls SATB2 in other tissue contexts (bone, brain) not tested
    • DeSUMOylation enzyme not identified
  15. 2021 High

    ATAC-seq and ChIP-seq in conditional intestinal knockout revealed SATB2 maintains colonic stem cell identity by modulating enhancer accessibility for CDX2 and HNF4A, demonstrating a non-neuronal, non-skeletal identity-maintenance function.

    Evidence Conditional Satb2 knockout in adult mouse colon, ATAC-seq, ChIP-seq, organoid culture, scRNA-seq, human colonic organoid SATB2 knockout

    PMID:34582804

    Open questions at the time
    • Direct physical interaction with CDX2/HNF4A not demonstrated
    • Whether SATB2 organizes chromatin loops at intestinal enhancers not tested by Hi-C
  16. 2022 Medium

    Caspase-7-mediated cleavage of SATB2 was identified as a trigger for myogenic differentiation, and Hi-C showed SATB2 binding at chromatin loop anchors influences sub-TAD organization, establishing proteolytic removal as a second PTM mechanism regulating SATB2 function.

    Evidence SATB2 deletion in myoblasts, caspase 7 cleavage assay, Hi-C chromatin interaction analysis, SATB2 ChIP-seq, differentiation assays

    PMID:35326417

    Open questions at the time
    • Specific caspase-7 cleavage site not mapped at residue level
    • Whether cleavage generates a functional fragment versus simple degradation not resolved
  17. 2023 High

    Discovery that SATB2 coordinates with NRF2 and recruits SWI/SNF subunits BRD7/BRG1 to sustain chromatin accessibility at antioxidative genes expanded the repertoire of chromatin-remodeling complexes SATB2 deploys in a context-dependent manner.

    Evidence ChIP-seq, ATAC-seq, co-IP of SATB2-NRF2 and SATB2-BRD7/BRG1, patient-derived xenografts and organoids in renal cell carcinoma

    PMID:36598364

    Open questions at the time
    • Whether SATB2-SWI/SNF interaction occurs in non-cancer contexts unknown
    • Structural basis of SATB2-BRG1 interaction not resolved

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved how SATB2 selects between its multiple co-regulator complexes (CBP/p300, HDAC1, LSD1/CoREST, SWI/SNF) at specific loci, whether SUMOylation and caspase cleavage constitute a general regulatory code across tissues, and what structural features of MAR sequences dictate SATB2 binding specificity genome-wide.
  • No structural model of SATB2 bound to DNA at atomic resolution
  • Integration of SUMOylation, caspase cleavage, and miRNA-mediated translational control into a unified regulatory logic not achieved
  • Comprehensive interactome comparisons across bone, brain, gut, and cancer contexts lacking

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003677 DNA binding 9 GO:0140110 transcription regulator activity 8 GO:0042393 histone binding 2
Localization
GO:0005634 nucleus 4 GO:0000228 nuclear chromosome 2
Pathway
R-HSA-1266738 Developmental Biology 6 R-HSA-74160 Gene expression (Transcription) 6 R-HSA-112316 Neuronal System 5 R-HSA-4839726 Chromatin organization 5
Complex memberships
LSD1/CoREST complexSWI/SNF (BRD7/BRG1)

Evidence

Reading pass · 31 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2006 SATB2 directly interacts with and enhances the activity of both RUNX2 and ATF4 transcription factors to regulate osteoblast differentiation; genetic confirmation via bone formation defects in Satb2/Runx2 and Satb2/Atf4 double heterozygous mice. SATB2 also directly represses Hoxa2 expression. Co-immunoprecipitation, reporter assays, double heterozygous mouse genetic epistasis, Satb2 knockout mouse phenotyping Cell High 16751105
2006 SATB2 is expressed as a nuclear matrix protein in branchial arches and osteoblast lineage cells; its loss causes craniofacial abnormalities and defects in osteoblast differentiation and function, identifying it as a molecular node in the transcriptional network for skeletal development. Satb2 knockout mouse generation, in situ hybridization, histology, gene expression analysis Cell High 16751105
2003 SATB2 encodes a DNA-binding protein with two CUT domains and a homeodomain, belonging to the nuclear matrix-attachment region (MAR) binding protein family; it is expressed in the developing palate and haploinsufficiency causes cleft palate. FISH mapping of translocation breakpoints, gene structure analysis, whole-mount in situ hybridization in mouse embryos Human molecular genetics High 12915443
2010 SATB2 directly binds and is targeted by miR-31 in cancer-associated fibroblasts; miR-31 directly targets the SATB2 3'UTR to regulate tumor cell migration and invasion. Microarray, luciferase reporter assay, RNA interference, overexpression studies in CAFs Cell cycle (Georgetown, Tex.) Medium 20980827
2011 SATB2 upregulates Osterix (Osx/Sp7) expression independent of Runx2 and synergistically enhances Runx2's regulatory effect on the Osx promoter; SATB2 overexpression in adult stem cells increases osteogenic gene expression and promotes bone regeneration in vivo. Osterix promoter-luciferase reporter assay in Runx2-deficient calvarial cells, lentiviral transduction, in vivo transplantation/bone defect model Tissue engineering. Part A Medium 21385070
2010 SATB2 binds ΔNp63α and enhances ΔNp63α-mediated transrepression by augmenting ΔNp63α engagement to p53-family responsive elements, promoting chemoresistance in head and neck squamous cell carcinoma. Co-immunoprecipitation (ΔNp63α-SATB2 interaction), reporter assay for transrepression, RNA interference knockdown with apoptosis readout EMBO reports Medium 20829881
2011 Satb2 regulates dendritic arborization and soma spacing of layer II/III pyramidal neurons in mouse cerebral cortex in a cell-autonomous and non-cell-autonomous manner, controlling adhesive/repulsive properties of neurons. In vivo RNA interference knockdown, live imaging, immunohistochemistry Cerebral cortex (New York, N.Y. : 1991) Medium 21885532
2012 CB1 cannabinoid receptor signaling prevents Satb2-mediated repression of the Ctip2 promoter, thereby increasing Ctip2-positive neuron generation and driving corticospinal motor neuron differentiation; CB1-/- mice show altered Ctip2/Satb2 axis and corticospinal tract defects. CB1 knockout mouse models, promoter reporter assays, immunostaining, genetic epistasis The Journal of neuroscience Medium 23175820
2012 Satb1 and Satb2 are dispensable for X chromosome inactivation in mice; female Satb1−/−Satb2−/− fibroblasts contain proper Barr bodies and no upregulation of X-linked genes. Double knockout mouse fibroblasts, FISH for Xist RNA, immunohistochemistry for H3K27me3, gene expression analysis Developmental cell Medium 23079603
2013 BMP (via Smad5/Hand2) and Hedgehog (via Smo/Shha) signaling pathways converge on neural crest cells to induce satb2 expression in pharyngeal arches; BMP signaling establishes competence for neural crest response to Hh signaling. Zebrafish mutant analysis, transplantation experiments, pharmacological inhibition (cyclopamine, dorsomorphin), in situ hybridization PloS one Medium 23555697
2014 Satb2 is required for proper differentiation of both callosal projection neurons (CPNs) and subcerebral projection neurons (SCPNs), including formation of the corticospinal tract; Satb2 conditional knockout reveals distinct early roles in deep-layer neurons for subcerebral axon development. Conditional Satb2 knockout in developing neocortex, electrophysiology, axon tracing, gene expression analysis Cerebral cortex (New York, N.Y. : 1991) High 25037921
2014 SATB2 promotes osteosarcoma migration and invasion by regulating cytoskeletal organization; SATB2 knockdown upregulates EPLIN (actin-binding protein), increases RhoA, decreases Rac1, and increases phosphorylation of FAK and paxillin. Rescuing EPLIN reverses the decreased invasion from SATB2 knockdown. shRNA knockdown, microarray, migration/invasion assays, Western blot for cytoskeletal signaling proteins, rescue experiments Oncogene Medium 25220418
2015 SATB2 directly activates transcription of Fezf2 and Sox5 (genes essential for subcerebral neuron development), and mutual regulation between Satb2 and Fezf2 enables Satb2 to promote subcerebral neuron identity in layer 5 and repress subcerebral characters in callosal neurons in a context-dependent manner. ChIP, reporter assays, in utero electroporation, conditional knockout mice, gene expression analysis Proceedings of the National Academy of Sciences of the United States of America High 26324926
2016 Satb2 in adult hippocampal neurons binds promoters of coding and non-coding genes; synaptic activity and BDNF upregulate Satb2, which controls hippocampal miRNA levels and is required for stabilization of long-term potentiation and long-term fear and object discrimination memory. Conditional Satb2 knockout in adult mouse forebrain, electrophysiology (LTP), behavioral memory tests, ChIP, miRNA profiling eLife High 27897969
2017 SATB2 nuclear mobility is mutation-dependent; missense variants in CUT1 (p.Arg389Cys) increase SATB2 nuclear mobility while CUT2 (p.Gly515Ser) and between CUT2-HOX domain variants reduce mobility, demonstrating that functional domains control chromatin/matrix association. Fluorescently tagged SATB2 expressed in patient fibroblasts, FRAP (fluorescence recovery after photobleaching), subcellular localization imaging Genetics in medicine Medium 28151491
2019 SATB2-AS1 lncRNA serves as a scaffold to recruit p300, whose acetylation of H3K27 and H3K9 at the SATB2 promoter upregulates SATB2 expression; SATB2 then recruits HDAC1 to the Snail promoter, repressing Snail transcription and inhibiting EMT in colorectal carcinoma. ChIP, RNA pulldown/scaffold assay, reporter assays, overexpression/knockdown in vitro and in vivo xenograft models Cancer research High 30858153
2017 Estrogen upregulates SATB2 through ERβ binding to estrogen response elements (ERE) at position -488 of the SATB2 gene, and this ERβ-SATB2 pathway mediates estrogen's effects on BMSC pluripotency, senescence, and osteogenic differentiation. ChIP for ERβ at SATB2 promoter ERE, ERα/ERβ inhibitor studies, SATB2 overexpression in OVX-BMSCs, in vivo transplantation Journal of cellular physiology Medium 29030963
2020 SATB2 recruits histone acetyltransferase CBP to the MAR sequence of the FOXM1 gene locus, driving FOXM1 expression and cell proliferation in glioma stem cells; pharmacological inhibition with CBP inhibitor C646 suppresses GSC proliferation and GBM growth. Co-immunoprecipitation (SATB2-CBP), ChIP (SATB2 binding to FOXM1 MAR), shRNA knockdown, in vivo xenograft, CBP inhibitor C646 treatment EMBO molecular medicine High 33124191
2021 SATB2 preserves LGR5+ colonic stem cell identity by modulating enhancer binding of intestinal transcription factors CDX2 and HNF4A; Satb2 loss in adult mice leads to stable conversion of colonic stem cells into small intestine ileal-like stem cells. Conditional Satb2 knockout in adult mouse colon, ATAC-seq, ChIP-seq, organoid culture, scRNA-seq, human colonic organoid SATB2 knockout Cell stem cell High 34582804
2021 SATB2 is SUMOylated on two acceptor lysines by the E3 SUMO ligase ZFP451 (SUMO2 modification); SUMO2-SATB2 drives ESC differentiation by silencing pluripotency genes and activating differentiation genes through interaction with the LSD1/CoREST complex and rewiring of higher-order chromatin interactions. Conditional Satb2 knockout, SUMO acceptor site mutagenesis (Satb2-KR), ZFP451 knockout, SUMO2-SATB2 fusion rescue, co-IP (SATB2-ZFP451, SATB2-LSD1/CoREST), Hi-C chromatin interaction analysis, ChIP-seq Genes & development High 34244292
2022 Caspase 7-mediated cleavage of SATB2 is a key event initiating myogenic differentiation; SATB2 binding within chromatin loops and near anchor points influences loop and sub-TAD domain formation, and its removal derepresses differentiation-inducing factors. SATB2 deletion in myoblasts, caspase 7 cleavage assay, genome-wide Hi-C chromatin interaction analysis, SATB2 ChIP-seq, differentiation assays Cells Medium 35326417
2011 SATB2 interacts with p63 (specifically ΔNp63α); AEC-associated p63 mutations (SAM domain) but not EEC-associated (DNA binding domain) mutations alter SATB2-p63 interaction. SATB2 attenuates p63-mediated transactivation of perp and specifically decreases p63 perp promoter binding. Co-immunoprecipitation, ChIP at perp promoter, reporter assay, co-expression analysis in branchial arch development The Journal of biological chemistry Medium 21965674
2019 SATB2 directly binds to the SLC26A3 (Cl-/HCO3- exchanger) promoter to regulate its expression; Satb2 intestinal epithelial-specific knockout disrupts Cl-/HCO3- exchange, alters gut microbiota homeostasis, and promotes colitis and colitis-associated colorectal cancer. ChIP demonstrating SATB2 binding at SLC26A3 promoter, luciferase reporter assay, conditional intestinal KO mice, 16S rDNA sequencing, fluorometric ion exchange assay Journal of Crohn's & colitis Medium 34019628
2023 YAP/TEAD4 activates SATB2 expression; SATB2 then coordinates with NRF2 to drive enhancer-promoter interactions amplifying antioxidative transcriptional activity, and recruits SWI/SNF complex subunits BRD7 and BRG1 to sustain DNA accessibility in renal cell carcinoma. ChIP-seq, ATAC-seq, co-IP (SATB2-NRF2, SATB2-BRD7/BRG1), transcriptome analysis, patient-derived xenografts and organoids, SATB2 inhibition studies Cancer research High 36598364
2019 EZH2-mediated H3K27me3 enrichment on the SATB2 gene represses SATB2 expression in pre-osteoblasts; maternal obesity increases EZH2 activity and H3K27me3 at the SATB2 locus, impairing fetal osteoblast differentiation. EZH2 knockdown or pre-osteoblast-specific EZH2 deletion increases SATB2 expression and trabecular bone mass. ChIP-seq for H3K27me3 and EZH2 genome-wide, Ezh2 conditional knockout (Ezh2flox/flox Osx-Cre), EZH2 knockdown/overexpression, primary embryonic osteogenic cell isolation FASEB journal Medium 31908011
2019 Satb2 deletion from adult mouse forebrain causes loss of synaptic LTP stabilization and impairs long-term memory; Satb2 binds promoters of both coding genes and miRNA genes in hippocampal neurons, controlling a large cohort of synaptic plasticity-related miRNAs. Conditional Satb2 knockout (adult forebrain), ChIP for Satb2 at gene promoters, miRNA microarray, LTP electrophysiology, behavioral fear conditioning and object discrimination tests eLife High 27897969
2019 SATB2 protein interactomes in neonatal vs adult neocortex show a developmental shift: neonatal SATB2 complexes are enriched for transcriptional repressors, while adult SATB2 complexes are enriched for proteins that stabilize de novo chromatin loops, indicating a shift from transcriptional repression to chromosomal superstructure organization. Immunoprecipitation-mass spectrometry (SATB2 interactome) from neonatal and adult mouse cortex, gene set analysis of GWAS data PLoS genetics Medium 30726206
2021 miR-541 (a eutherian-specific miRNA) and miR-92a/b inhibit SATB2 translation through its 3'UTR during early cortical development; their inactivation triggers premature SATB2 protein production in mouse and human cortical progenitors, demonstrating RNA interference as a mechanism timing cortical cell identity. miRNA reporter assays using SATB2 3'UTR, miRNA inhibition in mouse cortex and human cortical cells, immunofluorescence for SATB2 protein vs mRNA single-cell analysis Stem cell reports Medium 34019815
2021 A frameshift SATB2 mutant protein fails to translocate to the nucleus, remaining cytoplasmic, and consequently fails to activate Wnt/β-catenin signaling; wild-type SATB2 translocates to the nucleus and upregulates active β-catenin, promoting odontogenic differentiation. SATB2 mutation upregulates DKK1 and histone demethylase JHDM1D to inhibit Wnt/β-catenin signaling. Fluorescent immunocytochemistry for subcellular localization of WT vs mutant SATB2, Western blot for β-catenin, Wnt inhibitor XAV939 functional assay, WES mutation identification, differentiation assays in hDPSCs Stem cell research & therapy Medium 34863303
2023 SATB2 is a transcriptional suppressor of p16INK4a and competes with CUX1 for binding to the functional SNP rs1537371 at the CDKN2A/B locus; CUX1 transcriptionally represses SATB2 expression, creating a regulatory loop that fine-tunes p16INK4a expression and endothelial senescence. Reporter assay, overexpression competition assay, ChIP for CUX1 and SATB2 at rs1537371, siRNA knockdown, IL-1β senescence induction assay Aging cell Medium 36633253
2017 SATB2 suppresses lung cancer cell invasion and metastasis through regulation of histone methylation via G9a (a histone methyltransferase), affecting EMT-related protein expression. SATB2 knockdown/overexpression, invasion assay, Western blot for G9a and EMT markers, histone methylation analysis Clinical and experimental medicine Low 28667416

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2006 SATB2 is a multifunctional determinant of craniofacial patterning and osteoblast differentiation. Cell 433 16751105
2019 LncRNA MALAT1 shuttled by bone marrow-derived mesenchymal stem cells-secreted exosomes alleviates osteoporosis through mediating microRNA-34c/SATB2 axis. Aging 231 31659145
2003 Identification of SATB2 as the cleft palate gene on 2q32-q33. Human molecular genetics 226 12915443
2010 The role of miR-31 and its target gene SATB2 in cancer-associated fibroblasts. Cell cycle (Georgetown, Tex.) 137 20980827
2014 Satb2 Regulates the Differentiation of Both Callosal and Subcerebral Projection Neurons in the Developing Cerebral Cortex. Cerebral cortex (New York, N.Y. : 1991) 129 25037921
2007 Heterozygous nonsense mutation SATB2 associated with cleft palate, osteoporosis, and cognitive defects. Human mutation 109 17377962
2014 Cadherin-17 and SATB2 are sensitive and specific immunomarkers for medullary carcinoma of the large intestine. Archives of pathology & laboratory medicine 99 24437456
2009 The mRNA expression of SATB1 and SATB2 in human breast cancer. Cancer cell international 94 19642980
2016 SATB2-associated syndrome: Mechanisms, phenotype, and practical recommendations. American journal of medical genetics. Part A 90 27774744
2011 Roles of SATB2 in osteogenic differentiation and bone regeneration. Tissue engineering. Part A 89 21385070
2013 Elevated microRNA-31 expression regulates colorectal cancer progression by repressing its target gene SATB2. PloS one 83 24386467
2019 SATB2-AS1 Suppresses Colorectal Carcinoma Aggressiveness by Inhibiting SATB2-Dependent Snail Transcription and Epithelial-Mesenchymal Transition. Cancer research 79 30858153
2013 Further delineation of the SATB2 phenotype. European journal of human genetics : EJHG 77 24301056
2016 Satb2 determines miRNA expression and long-term memory in the adult central nervous system. eLife 69 27897969
2015 Mutual regulation between Satb2 and Fezf2 promotes subcerebral projection neuron identity in the developing cerebral cortex. Proceedings of the National Academy of Sciences of the United States of America 68 26324926
2012 The CB(1) cannabinoid receptor drives corticospinal motor neuron differentiation through the Ctip2/Satb2 transcriptional regulation axis. The Journal of neuroscience : the official journal of the Society for Neuroscience 68 23175820
2017 SATB2 as an Immunohistochemical Marker for Colorectal Adenocarcinoma: A Concise Review of Benefits and Pitfalls. Archives of pathology & laboratory medicine 66 28968158
2013 The role of SATB2 in skeletogenesis and human disease. Cytokine & growth factor reviews 60 24411565
2017 Estrogen regulates stemness and senescence of bone marrow stromal cells to prevent osteoporosis via ERβ-SATB2 pathway. Journal of cellular physiology 59 29030963
2016 MiR-31 inhibits migration and invasion by targeting SATB2 in triple negative breast cancer. Gene 56 27593563
2013 Expression of transcription factor Satb2 in adult mouse brain. Anatomical record (Hoboken, N.J. : 2007) 56 23386513
2021 SATB2 preserves colon stem cell identity and mediates ileum-colon conversion via enhancer remodeling. Cell stem cell 54 34582804
2013 MicroRNA expression signature for Satb2-induced osteogenic differentiation in bone marrow stromal cells. Molecular and cellular biochemistry 53 24218084
2010 SATB2 augments ΔNp63α in head and neck squamous cell carcinoma. EMBO reports 51 20829881
2019 Mutation update for the SATB2 gene. Human mutation 50 31021519
2017 Clinical and molecular consequences of disease-associated de novo mutations in SATB2. Genetics in medicine : official journal of the American College of Medical Genetics 50 28151491
2016 Special AT-rich sequence-binding protein 2 (SATB2) expression is sensitive but may not be specific for osteosarcoma as compared with other high-grade primary bone sarcomas. Histopathology 50 26644288
2020 SATB2 drives glioblastoma growth by recruiting CBP to promote FOXM1 expression in glioma stem cells. EMBO molecular medicine 48 33124191
2015 miR-211 suppresses hepatocellular carcinoma by downregulating SATB2. Oncotarget 48 25888635
2014 SATB2 enhances migration and invasion in osteosarcoma by regulating genes involved in cytoskeletal organization. Oncogene 46 25220418
2011 Satb2 is required for dendritic arborization and soma spacing in mouse cerebral cortex. Cerebral cortex (New York, N.Y. : 1991) 46 21885532
2015 Dental Follicle Cells Participate in Tooth Eruption via the RUNX2-MiR-31-SATB2 Loop. Journal of dental research 45 25818585
2015 Further supporting evidence for the SATB2-associated syndrome found through whole exome sequencing. American journal of medical genetics. Part A 45 25885067
2017 SATB2/β-catenin/TCF-LEF pathway induces cellular transformation by generating cancer stem cells in colorectal cancer. Scientific reports 43 28887549
2020 SATB2 is a novel biomarker and therapeutic target for cancer. Journal of cellular and molecular medicine 42 32885593
2016 SATB2 is expressed in Merkel cell carcinoma. Archives of dermatological research 41 27262585
2016 MicroRNA-34b/c inhibits aldosterone-induced vascular smooth muscle cell calcification via a SATB2/Runx2 pathway. Cell and tissue research 41 27503378
2019 Loss of Satb2 in the Cortex and Hippocampus Leads to Abnormal Behaviors in Mice. Frontiers in molecular neuroscience 35 30809123
2017 Genotype and phenotype in 12 additional individuals with SATB2-associated syndrome. Clinical genetics 35 28139846
2017 Cellular transformation of human mammary epithelial cells by SATB2. Stem cell research 34 28167342
2016 Role of SATB2 in human pancreatic cancer: Implications in transformation and a promising biomarker. Oncotarget 34 27472393
2014 Intragenic duplication--a novel causative mechanism for SATB2-associated syndrome. American journal of medical genetics. Part A 34 25251319
2021 Silencing of miR-483-5p alleviates postmenopausal osteoporosis by targeting SATB2 and PI3K/AKT pathway. Aging 33 33621956
2018 Genes regulated by SATB2 during neurodevelopment contribute to schizophrenia and educational attainment. PLoS genetics 32 30040823
2016 Downregulation of SATB2 is critical for induction of epithelial-to-mesenchymal transition and invasion of NSCLC cells. Lung cancer (Amsterdam, Netherlands) 28 27393518
2018 Role of conventional immunomarkers, HNF4-α and SATB2, in the differential diagnosis of pulmonary and colorectal adenocarcinomas. Histopathology 27 29243296
2018 Role of miR-31 and SATB2 in arsenic-induced malignant BEAS-2B cell transformation. Molecular carcinogenesis 27 29603397
2018 The Role of SATB2 as a Diagnostic Marker of Sinonasal Intestinal-type Adenocarcinoma. Applied immunohistochemistry & molecular morphology : AIMM 26 27258560
2018 SATB2 Is Superior to CDX2 in Distinguishing Signet Ring Cell Carcinoma of the Upper Gastrointestinal Tract and Lower Gastrointestinal Tract. The American journal of surgical pathology 26 30212392
2017 SATB2 suppresses non-small cell lung cancer invasiveness by G9a. Clinical and experimental medicine 26 28667416
2017 Metformin reduces SATB2-mediated osteosarcoma stem cell-like phenotype and tumor growth via inhibition of N-cadherin/NF-kB signaling. European review for medical and pharmacological sciences 26 29131265
2017 The miR-875-5p inhibits SATB2 to promote the invasion of lung cancer cells. Gene 26 29196257
2016 miR-449a inhibits colorectal cancer progression by targeting SATB2. Oncotarget 26 29254139
2016 miR-33a-5p modulates TNF-α-inhibited osteogenic differentiation by targeting SATB2 expression in hBMSCs. FEBS letters 25 26785690
2019 Value of SATB2, ISL1, and TTF1 to differentiate rectal from other gastrointestinal and lung well-differentiated neuroendocrine tumors. Pathology, research and practice 23 31133441
2017 The miR-599 promotes non-small cell lung cancer cell invasion via SATB2. Biochemical and biophysical research communications 23 28167280
2014 Lentiviral-mediated expression of SATB2 promotes osteogenic differentiation of bone marrow stromal cells in vitro and in vivo. European journal of oral sciences 23 24666017
2019 Deregulation of SATB2 in carcinogenesis with emphasis on miRNA-mediated control. Carcinogenesis 22 30916759
2016 Value of SATB2 immunostaining in the distinction between small intestinal and colorectal adenocarcinomas. Journal of clinical pathology 22 27169755
2012 Satb1 and Satb2 are dispensable for X chromosome inactivation in mice. Developmental cell 22 23079603
2019 MiR-34a inhibits the proliferation, migration, and invasion of oral squamous cell carcinoma by directly targeting SATB2. Journal of cellular physiology 21 31663131
2017 Role of SATB2 in distinguishing the site of origin in glandular lesions of the bladder/urinary tract. Human pathology 21 28711650
2019 Colitis-associated colorectal adenocarcinomas are frequently associated with non-intestinal mucin profiles and loss of SATB2 expression. Modern pathology : an official journal of the United States and Canadian Academy of Pathology, Inc 20 30710095
2017 Bone health and SATB2-associated syndrome. Clinical genetics 20 28787087
2013 Bmp and Shh signaling mediate the expression of satb2 in the pharyngeal arches. PloS one 20 23555697
2021 SATB2 in Neoplasms of Lung, Pancreatobiliary, and Gastrointestinal Origins. American journal of clinical pathology 19 32914850
2019 Loss of SATB2 Expression Is a Biomarker of Inflammatory Bowel Disease-associated Colorectal Dysplasia and Adenocarcinoma. The American journal of surgical pathology 19 31318711
2019 Higher expression of SATB2 in hepatocellular carcinoma of African Americans determines more aggressive phenotypes than those of Caucasian Americans. Journal of cellular and molecular medicine 19 31602781
2023 YAP-Activated SATB2 Is a Coactivator of NRF2 That Amplifies Antioxidative Capacity and Promotes Tumor Progression in Renal Cell Carcinoma. Cancer research 18 36598364
2021 Special AT-rich sequence-binding protein 2 (Satb2) synergizes with Bmp9 and is essential for osteo/odontogenic differentiation of mouse incisor mesenchymal stem cells. Cell proliferation 18 33660290
2021 SATB2 Defect Promotes Colitis and Colitis-associated Colorectal Cancer by Impairing Cl-/HCO3- Exchange and Homeostasis of Gut Microbiota. Journal of Crohn's & colitis 18 34019628
2019 Satb2 regulates proliferation and nuclear integrity of pre-osteoblasts. Bone 18 31325654
2015 SATB2 suppresses gastric cancer cell proliferation and migration. Tumour biology : the journal of the International Society for Oncodevelopmental Biology and Medicine 18 26508023
2020 microRNA-4270-5p inhibits cancer cell proliferation and metastasis in hepatocellular carcinoma by targeting SATB2. Human cell 17 32504285
2021 ZFP451-mediated SUMOylation of SATB2 drives embryonic stem cell differentiation. Genes & development 16 34244292
2019 Maternal regulation of SATB2 in osteo-progeniters impairs skeletal development in offspring. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 16 31908011
2022 SOX9 and SATB2 immunohistochemistry cannot reliably distinguish between osteosarcoma and chondrosarcoma on biopsy material. Human pathology 15 35016891
2022 Chromatin Reorganization during Myoblast Differentiation Involves the Caspase-Dependent Removal of SATB2. Cells 15 35326417
2021 Growth, development, and phenotypic spectrum of individuals with deletions of 2q33.1 involving SATB2. Clinical genetics 15 33381861
2018 SATB2 Shows Different Profiles Between Appendiceal Adenocarcinomas Ex Goblet Cell Carcinoids and Appendiceal/Colorectal Conventional Adenocarcinomas: An Immunohistochemical Study With Comparison to CDX2. Gastroenterology research 15 29915633
2018 miR-34a Inhibits Cell Proliferation by Targeting SATB2 in Hepatocellular Carcinoma. BioMed research international 15 30627547
2017 The crosstalk between p38 and Akt signaling pathways orchestrates EMT by regulating SATB2 expression in NSCLC cells. Tumour biology : the journal of the International Society for Oncodevelopmental Biology and Medicine 15 28937318
2020 SATB2 knockdown decreases hypoxia-induced autophagy and stemness in oral squamous cell carcinoma. Oncology letters 14 32566006
2020 Circ_SATB2 Attenuates the Anti-Tumor Role of Celastrol in Non-Small-Cell Lung Carcinoma Through Targeting miR-33a-5p/E2F7 Axis. OncoTargets and therapy 14 33239891
2019 Genes encoding SATB2-interacting proteins in adult cerebral cortex contribute to human cognitive ability. PLoS genetics 14 30726206
2019 Speech, language, and feeding phenotypes of SATB2-associated syndrome. Clinical genetics 14 31392730
2018 Patients with SATB2-associated syndrome exhibiting multiple odontomas. American journal of medical genetics. Part A 14 30575289
2023 SATB2, coordinated with CUX1, regulates IL-1β-induced senescence-like phenotype in endothelial cells by fine-tuning the atherosclerosis-associated p16INK4a expression. Aging cell 13 36633253
2023 Psammomatoid Ossifying Fibroma Is Defined by SATB2 Rearrangement. Modern pathology : an official journal of the United States and Canadian Academy of Pathology, Inc 13 36788065
2021 Identification of retinal ganglion cell types expressing the transcription factor Satb2 in three primate species. The Journal of comparative neurology 13 33527361
2021 A novel mutation of SATB2 inhibits odontogenesis of human dental pulp stem cells through Wnt/β-catenin signaling pathway. Stem cell research & therapy 13 34863303
2020 The lncRNAs RP1-261G23.7, RP11-69E11.4 and SATB2-AS1 are a novel clinical signature for predicting recurrent osteosarcoma. Bioscience reports 12 31850493
2022 Involvement of the TNF-α/SATB2 axis in the induced apoptosis and inhibited autophagy of osteoblasts by the antipsychotic Risperidone. Molecular medicine (Cambridge, Mass.) 11 35505281
2021 SATB2 induction of a neural crest mesenchyme-like program drives melanoma invasion and drug resistance. eLife 11 33527896
2019 SATB2-associated syndrome (SAS) and associated dental findings. Special care in dentistry : official publication of the American Association of Hospital Dentists, the Academy of Dentistry for the Handicapped, and the American Society for Geriatric Dentistry 11 30648748
2011 Special AT-rich binding protein-2 (SATB2) differentially affects disease-causing p63 mutant proteins. The Journal of biological chemistry 11 21965674
2022 Comprehensive clinicopathologic, molecular, and immunologic characterization of colorectal carcinomas with loss of three intestinal markers, CDX2, SATB2, and KRT20. Virchows Archiv : an international journal of pathology 10 35029777
2022 circ-Iqsec1 induces bone marrow-derived mesenchymal stem cell (BMSC) osteogenic differentiation through the miR-187-3p/Satb2 signaling pathway. Arthritis research & therapy 10 36517907
2021 A eutherian-specific microRNA controls the translation of Satb2 in a model of cortical differentiation. Stem cell reports 10 34019815
2021 Regional Specific Differentiation of Integumentary Organs: Regulation of Gene Clusters within the Avian Epidermal Differentiation Complex and Impacts of SATB2 Overexpression. Genes 10 34440465