Affinage

SP2

Transcription factor Sp2 · UniProt Q02086

Length
613 aa
Mass
64.9 kDa
Annotated
2026-06-10
100 papers in source corpus 24 papers cited in narrative 24 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 8/8 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

SP2 is a Sp/KLF-family transcription factor that controls genes governing proliferation, metabolism, and cell-fate transitions, and is essential for early mammalian development (PMID:22684502, PMID:20221402). Unlike its paralogs, SP2 engages chromatin largely through protein-protein recruitment rather than autonomous zinc-finger DNA binding: its glutamine-rich N-terminal region, acting through an Sp-box motif, recruits the histone-fold factor Nf-y to CCAAT motifs genome-wide and reciprocally potentiates Nf-y binding (PMID:25793500, PMID:30337366). At composite regulatory elements SP2 is tethered to a TALE homeoprotein complex of Pbx1:Prep1 together with Nf-y, again with mutual potentiation of chromatin occupancy (PMID:30337366). SP2 also binds GC-box/Sp consensus sites directly (consensus 5'-GGGCGGGAC-3'), where it occupies proximal promoters of genes for transcription, replication, and metabolism, activating some targets while repressing others such as the cholesterol-synthesis battery (PMID:22684502, PMID:14726517). Direct repression can proceed through recruitment of histone deacetylase activity, as shown at the CEACAM1 promoter (PMID:15126343). SP2 localizes to nuclear-matrix-associated subnuclear foci via a sequence spanning its first zinc finger that also encodes a bipartite nuclear localization signal (PMID:16467376). Genetically, SP2 is required for autonomous proliferation of fibroblasts and for the neurogenic (but not expansive) divisions of cortical neural stem cells, where its loss prolongs mitosis (PMID:20221402, PMID:32001437). Its 9aaTAD transactivation domain was inactivated early in chordate evolution by valine accumulation, rendering SP2 a relatively weak trans-activator in mammals (PMID:14726517, PMID:31375868).

Mechanistic history

Synthesis pass · year-by-year structured walk · 11 steps
  1. 2000 Medium

    Established that SP2 acts on GC-rich cis-elements with context-dependent outcomes, raising the question of whether it activates or represses transcription.

    Evidence transient transfection, promoter mutation, and EMSA comparing Drosophila SL2 and mammalian cells on the CTα gene

    PMID:10744779

    Open questions at the time
    • Mechanism underlying cell-context switch between repression and activation not defined
    • No genome-wide target mapping
  2. 2004 High

    Defined SP2's high-affinity DNA consensus and showed its intrinsic trans-activation is weak and negatively regulated, distinguishing it from strong activators like Sp1.

    Evidence PCR-based consensus selection, kinetic binding assays, chimeric Sp1/Sp2 constructs, and DHFR reporter assays in vitro and in mammalian cells

    PMID:14726517

    Open questions at the time
    • Identity of the negative regulator of DNA-binding/activation unknown
    • Endogenous direct targets not defined
  3. 2004 High

    Provided a concrete repressive mechanism, showing SP2 silences a target promoter by recruiting HDAC activity.

    Evidence reciprocal ChIP, in vitro binding, overexpression/knockdown, and HDAC activity assay at the CEACAM1 promoter in prostate epithelial cells

    PMID:15126343

    Open questions at the time
    • Specific HDAC isoform and corepressor adaptors not identified
    • Generality of HDAC recruitment across other SP2 targets not tested
  4. 2006 High

    Localized SP2 to the nuclear matrix and mapped the domains responsible, linking its first zinc finger to subnuclear targeting and nuclear import.

    Evidence nuclear fractionation, live-cell imaging, and GFP deletion-chimera analysis

    PMID:16467376

    Open questions at the time
    • Functional consequence of nuclear-matrix association for transcription not established
    • Matrix-anchoring partner proteins unknown
  5. 2006 High

    Showed SP2 participates in a multiprotein repressor complex regulated by hormone signaling, framing it as a node in inducible gene control.

    Evidence siRNA, reporter, EMSA, and ChIP on the MMP-9 promoter with FXR/SHP/KLF6 perturbation

    PMID:17071613

    Open questions at the time
    • Direct SP2-KLF6 interaction surface not mapped
    • Whether SP2 is the DNA-anchoring subunit of the complex unclear
  6. 2010 High

    Established SP2 as non-redundant and essential, demonstrating an organismal and cell-autonomous proliferative requirement distinct from other Sp factors.

    Evidence constitutive and conditional knockout mouse models with MEF proliferation assays and in situ hybridization

    PMID:20221402

    Open questions at the time
    • Molecular targets mediating the proliferation defect not pinpointed
    • Cause of pre-E9.5 lethality at the pathway level undefined
  7. 2012 High

    Mapped the genome-wide GC-box occupancy and transcriptional output of SP2, revealing a dual activator/repressor role and direct control of cholesterol synthesis genes.

    Evidence ChIP-seq, RNAi, in vitro binding, and transcriptome profiling

    PMID:22684502

    Open questions at the time
    • How activation versus repression is selected at each promoter not resolved
    • Coactivator/corepressor partners genome-wide not catalogued
  8. 2015 High

    Resolved that SP2 binds CCAAT motifs not via its zinc fingers but through its glutamine-rich N-terminus recruiting Nf-y, redefining its mode of chromatin engagement.

    Evidence ChIP-seq with zinc-finger mutant rescue in matched KO MEFs and co-occupancy analysis

    PMID:25793500

    Open questions at the time
    • Direct vs. bridged contact between SP2 N-terminus and Nf-y not structurally defined
    • Functional output of CCAAT co-occupancy on target genes incomplete
  9. 2018 High

    Extended the cofactor model to composite TALE+CCAAT elements, showing SP2 forms and potentiates a Pbx1:Prep1-Nf-y complex via its Sp-box.

    Evidence ChIP-exo, reciprocal Co-IP, and domain deletion/mutation in MEFs

    PMID:30337366

    Open questions at the time
    • Stoichiometry and architecture of the SP2-Pbx1:Prep1-Nf-y assembly unknown
    • Target genes regulated by composite sites not enumerated
  10. 2019 Medium

    Explained why mammalian SP2 is a weak activator, attributing it to evolutionary inactivation of its 9aaTAD by valine accumulation.

    Evidence amino-acid substitution analysis, comparative genomics, and transactivation reporter assays

    PMID:31375868

    Open questions at the time
    • Whether intron-encoded 9aaTADs are used in any normal human context untested
    • Link between weak TAD and the recruitment-based mechanism not directly probed
  11. 2020 Medium

    Defined a stage-specific developmental role, showing SP2 is required for neurogenic divisions and timely mitotic progression in cortical progenitors.

    Evidence conditional deletion with MADM clonal analysis, cell-cycle analysis, and neuronal layer quantification in mice

    PMID:32001437

    Open questions at the time
    • Mitotic targets of SP2 driving the prolonged mitosis not identified
    • Why expansive divisions are SP2-independent unexplained

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved how SP2 selects activation versus repression at individual promoters and which corepressor/coactivator and matrix-anchoring partners convert its weak-TAD, recruitment-based binding into specific transcriptional outputs.
  • No structural model of SP2 with Nf-y or Pbx1:Prep1
  • Direct effectors of the proliferation and mitotic phenotypes unknown
  • Many cancer-context targets rest on single-lab ChIP/reporter evidence without reciprocal validation

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 4 GO:0003677 DNA binding 2 GO:0060090 molecular adaptor activity 2
Localization
GO:0005634 nucleus 2 GO:0005654 nucleoplasm 1
Pathway
R-HSA-74160 Gene expression (Transcription) 3 R-HSA-1266738 Developmental Biology 2
Partners
KLF6NFYAPBX1PREP1SHP

Evidence

Reading pass · 24 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2015 Sp2 binds to CCAAT motifs genome-wide in mouse embryonic fibroblasts via its glutamine-rich amino-terminal region, independently of its zinc finger domain, and recruits the histone-fold CCAAT-binding factor Nf-y; Nf-y is critical for recruitment of Sp2 to co-occupied regulatory elements, and reciprocally Sp2 potentiates Nf-y binding at shared sites. ChIP-seq in MEFs; re-expression of Sp2/Sp3 zinc finger mutants in corresponding knockout MEFs; co-occupancy analysis PLoS genetics High 25793500
2018 Sp2 is recruited to composite genomic sites containing a TALE homeoprotein recognition sequence and an Nf-y recognition sequence separated by ~11 bp; a complex of Pbx1:Prep1 and Nf-y recruits Sp2 to co-occupied regulatory elements, and in turn Sp2 potentiates binding of Pbx1:Prep1 and Nf-y. The Sp-box motif near the Sp2 N-terminus is required for this cofactor function. ChIP-exo sequencing in MEFs; co-immunoprecipitation; domain deletion/mutation analysis The Journal of biological chemistry High 30337366
2012 Sp2 binds GC-boxes and occupies proximal promoters of genes required for gene expression, replication, metabolism and signalling; it activates transcription factor and co-activator genes and represses the entire battery of cholesterol synthesis genes. Sp2 is required for cellular proliferation. ChIP-seq, RNA interference, in vitro DNA binding assays, global gene-expression profiling Nucleic acids research High 22684502
2004 Sp2 binds the CEACAM1 promoter in vitro and in vivo, represses CEACAM1 transcription by recruiting histone deacetylase activity to the promoter, and overexpression of Sp2 down-regulates endogenous CEACAM1 in normal prostate epithelial cells. In vitro DNA binding, chromatin immunoprecipitation, transient overexpression, siRNA knockdown, HDAC activity assay Cancer research High 15126343
2006 SP2 and KLF6 form a repressor complex on the MMP-9 promoter to maintain its silenced state; upon FXR activation, SHP interacts with SP2 and KLF6 and disrupts this repressor complex, enabling MMP-9 up-regulation and endothelial cell migration. siRNA silencing, promoter reporter assays, gel shift assays, chromatin immunoprecipitation, gain- and loss-of-function approaches The Journal of biological chemistry High 17071613
2004 Sp2 DNA-binding consensus sequence is 5'-GGGCGGGAC-3' and Sp2 binds this sequence with high affinity (Kd ~225 pM) in vitro; however, Sp2 DNA-binding activity and trans-activation are negatively regulated in mammalian cells, and Sp2 is a relatively weak trans-activator with little capacity for additive or synergistic activation. PCR-based consensus determination, kinetic binding assays (in vitro), chimeric Sp1/Sp2 constructs, transient co-transfection with DHFR promoter reporter The Journal of biological chemistry High 14726517
2006 Sp2 localizes to subnuclear foci associated with the nuclear matrix; a 37 amino acid sequence spanning the first zinc finger is sufficient to direct nuclear matrix association and also encodes a bipartite nuclear localization sequence. A second nuclear matrix targeting sequence is within the Sp2 trans-activation domain. Nuclear fractionation, live-cell imaging, deletion analysis with GFP-tagged chimeric proteins, immunofluorescence Molecular biology of the cell High 16467376
2010 Sp2 is essential for early mouse development (constitutive knockout lethal before E9.5 with severe growth retardation) and for autonomous proliferation of mouse embryonic fibroblasts; conditional ablation of Sp2 in MEFs strongly impairs cell proliferation. Sp2 has a non-redundant function distinct from Sp1, Sp3, and Sp4. Conditional and constitutive knockout mouse models (tri-loxP strategy, Cre-mediated ablation), MEF proliferation assays, whole-mount in situ hybridization PloS one High 20221402
2009 Zebrafish Sp2 is maternally inherited, localizes to the nuclear matrix in a DNA-independent fashion, and is required for completion of gastrulation; knockdown phenotype establishes a role in early embryonic development. Zebrafish morpholino knockdown, RT-PCR expression analysis, GFP reporter transgenic lines, in vitro transcription assays The Journal of biological chemistry Medium 19959469
2010 Transgenic overexpression of Sp2 in epidermal basal keratinocytes in mice causes arrest of the epidermal differentiation program (homozygous animals perish within 2 weeks), increased susceptibility to wound- and carcinogen-induced papillomagenesis, and Sp2 protein levels increase with carcinogen-induced squamous cell carcinoma progression. Transgenic mouse overexpression (hemizygous and homozygous), carcinogen treatment, histological analysis Cancer research Medium 20959487
2016 Sp2 directly binds to GC-box/Sp-binding motifs in the human RORγT promoter and is required for maintaining RORγT expression; Sp2 is necessary for maximal IL-17 expression in in vitro-differentiated Th17 cells. Promoter activity assays, in situ mutagenesis, chromatin immunoprecipitation, siRNA knockdown, real-time RT-PCR in human lymphocytic cell line and differentiated Th17 cells Journal of leukocyte biology Medium 27256574
2020 Loss of Sp2 in cortical progenitors undergoing neurogenic divisions results in prolonged mitosis due to extension of early mitotic stages, and depletion of upper layer neurons; early cortical neural stem cells expand normally without Sp2, indicating a stage-specific requirement. Conditional genetic deletion in mice combined with MADM (mosaic analysis with double markers), cell cycle analysis, neuronal layer quantification Development (Cambridge, England) Medium 32001437
2019 The 9aaTAD transactivation domain in Sp2 was inactivated early in chordate evolution by accumulation of valine residues; Sp2 secondary 9aaTADs are encoded in introns from fish to primates, and in the gibbon genome such intron-9aaTADs are used as exons, converting Sp2 into a transcriptional activator. Amino acid substitutions showed that glutamine residues are dispensable for 9aaTAD function. Amino acid substitution analysis, comparative genomics, transactivation reporter assays Cellular and molecular life sciences : CMLS Medium 31375868
2005 Sp2 protein present in muscle cell nuclear extracts binds a specific site in the mouse Id3 proximal promoter in vitro; mutation of this Sp2 site substantially reduces Id3 promoter activity in proliferating muscle cells, and protein binding to this site is markedly reduced in differentiated muscle cells. DNase I footprinting, EMSA, promoter mutagenesis, nuclear extract binding assays with Sp2 antibody supershift Biochimica et biophysica acta Medium 16216350
2021 Sp2 transcription factor directly binds the promoter regions of MMP2, MMP9, and VE-cadherin to promote their transcription, thereby regulating vasculogenic mimicry formation in glioma cells. Sp2 expression is regulated upstream by miR-651-3p targeting its 3'-UTR. ChIP assay, luciferase reporter assay, siRNA knockdown, transwell/VM formation assays Cell death & disease Medium 33542193
2021 Sp2 directly binds the promoter region of miR-195-5p to repress its expression; reduced Sp2 in bicuspid aortic valve interstitial cells increases SMAD7 expression and attenuates Smad2/3 signaling, promoting osteogenic differentiation through a SMAD-dependent pathway. ChIP assay, cotransfection with SP2 shRNA and miR-195-5p mimic, western blot, immunofluorescence in porcine VICs Experimental cell research Medium 34914964
2023 Sp2, activated downstream of AMPK, directly binds the NEAT1 promoter to induce its expression; this forms a LTF/AMPK/SP2/NEAT1/miR-214-5p positive feedback loop promoting radioresistance via autophagy in lung squamous cell carcinoma. ChIP assay, luciferase reporter assay, co-immunoprecipitation (LTF-AMPK), siRNA knockdown, functional radioresistance assays International journal of biological sciences Medium 37056929
2024 Sp2 promotes the transcription of the Vdac2 gene by directly binding to its promoter; this Sp2/VDAC2 axis mediates DBP-induced ferroptosis in mouse Leydig cells. ChIP assay, promoter reporter assay, siRNA knockdown, western blot, ferroptosis assays Environmental research Low 38246300
2021 SP2 regulates ANLN (Anillin) expression by binding to its promoter; SP2-mediated ANLN upregulation promotes colorectal carcinoma cell proliferation via PI3K/AKT and MAPK pathways. Chromatin immunoprecipitation, luciferase reporter assay, siRNA knockdown, CCK-8, colony formation, mouse xenograft model Journal of investigative surgery Low 33757382
2000 Sp2 represses Sp1- and Sp3-driven transcription of the CTα gene in Drosophila SL2 cells but stimulates transcription in mammalian C3H10T1/2 cells, acting through specific cis-acting GC-rich elements; effectiveness depends on cellular background and organization of three cis-acting elements. Transient transfections, promoter mutation analysis, electromobility gel shifts, immunoblot Journal of lipid research Medium 10744779
2005 Sp1 and Sp2 transcription factors bind to GC-rich elements in the SULT2B1b promoter and 5'-UTR, inducing reporter gene activity; this induction is markedly augmented by histone deacetylase inhibition. Supershift EMSA (identifying Sp1 and Sp2 binding), reporter assays, HDAC inhibitor treatment FEBS letters Low 15953604
2010 Sp2 transcription is regulated by at least two promoters with differing cell-type specificity; the 5'-UTR undergoes alternative splicing; sub-genomic mRNAs capable of encoding a partial-Sp2 protein are produced in a tissue- and cell-type-specific manner; Sp2 is widely expressed in embryonic brain and neurogenic regions of postnatal brain. cDNA cloning, RT-PCR, promoter deletion analysis in cell lines, RNA in situ hybridization Biochimica et biophysica acta Medium 20353838
2024 CD93 interacts with MMRN2 and integrin β1; knockdown of CD93 inhibits activation of integrin β1, thereby suppressing the PI3K/AKT/SP2 signaling pathway and inhibiting breast cancer cell vasculogenic mimicry. Overexpression of SP2 rescues the effects of CD93 knockdown. Co-immunoprecipitation, siRNA knockdown, lentiviral overexpression, western blot, in vivo xenograft model Journal of biochemical and molecular toxicology Low 38511888
2021 Src-1 interacts with SP2 protein (co-immunoprecipitation) and together they regulate EMT-related genes (E-cadherin, N-cadherin, vimentin, ZEB1) and apoptosis/proliferation genes in nasopharyngeal carcinoma cells; SP2 knockdown reverses the pro-tumorigenic effects of Src-1 overexpression. Co-immunoprecipitation, siRNA knockdown, overexpression, western blot, CCK-8, transwell assays Open medicine (Warsaw, Poland) Low 34307888

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2006 Palladium-catalyzed alkylation of sp2 and sp3 C-H bonds with methylboroxine and alkylboronic acids: two distinct C-H activation pathways. Journal of the American Chemical Society 311 17002342
2016 Nanoscale Electrochemistry of sp(2) Carbon Materials: From Graphite and Graphene to Carbon Nanotubes. Accounts of chemical research 93 27501067
2006 Disruption of an SP2/KLF6 repression complex by SHP is required for farnesoid X receptor-induced endothelial cell migration. The Journal of biological chemistry 68 17071613
2004 Identification of Sp2 as a transcriptional repressor of carcinoembryonic antigen-related cell adhesion molecule 1 in tumorigenesis. Cancer research 68 15126343
2013 Fullerene-sp2-iminosugar balls as multimodal ligands for lectins and glycosidases: a mechanistic hypothesis for the inhibitory multivalent effect. Chemistry (Weinheim an der Bergstrasse, Germany) 66 24150869
2020 Conformation and Aromaticity Switching in a Curved Non-Alternant sp2 Carbon Scaffold. Angewandte Chemie (International ed. in English) 61 32815658
1988 Measurement of a breast cancer associated antigen detected by monoclonal antibody SP-2 in sera of cancer patients. Breast cancer research and treatment 61 3164223
2014 miR-638 suppresses cell proliferation in gastric cancer by targeting Sp2. Digestive diseases and sciences 60 24623314
2023 Designing Single-Atom Active Sites on sp2 -Carbon Linked Covalent Organic Frameworks to Induce Bacterial Ferroptosis-Like for Robust Anti-Infection Therapy. Advanced science (Weinheim, Baden-Wurttemberg, Germany) 58 36847061
1995 Allele-specific methylation and expression of an imprinted U2af1-rs1 (SP2) gene. Nucleic acids research 53 7870588
2013 Spatial and temporal control of the diazonium modification of sp2 carbon surfaces. Journal of the American Chemical Society 48 24325138
2015 Zinc finger independent genome-wide binding of Sp2 potentiates recruitment of histone-fold protein Nf-y distinguishing it from Sp1 and Sp3. PLoS genetics 46 25793500
2012 Intramolecular dehydrogenative coupling of sp2 C-H and sp3 C-H bonds: an expeditious route to 2-oxindoles. Organic letters 46 23134301
1994 Elimination of endogenous aberrant kappa chain transcripts from sp2/0-derived hybridoma cells by specific ribozyme cleavage: utility in genetic therapy of HIV-1 infections. Nucleic acids research 46 7816635
2024 Stabilizing atomic Ru species in conjugated sp2 carbon-linked covalent organic framework for acidic water oxidation. Nature communications 45 38926414
2012 Genome-wide localization and expression profiling establish Sp2 as a sequence-specific transcription factor regulating vitally important genes. Nucleic acids research 43 22684502
1999 Four genotypic variants of a Spodoptera exigua Nucleopolyhedrovirus (Se-SP2) are distinguishable by a hypervariable genomic region. Virus research 43 10854166
2010 A Fluorescent sp2-iminosugar with pharmacological chaperone activity for gaucher disease: synthesis and intracellular distribution studies. Chembiochem : a European journal of chemical biology 42 21064079
2009 Effect of artesunate on inhibiting proliferation and inducing apoptosis of SP2/0 myeloma cells through affecting NFkappaB p65. International journal of hematology 40 19728025
2000 Functional significance of Sp1, Sp2, and Sp3 transcription factors in regulation of the murine CTP:phosphocholine cytidylyltransferase alpha promoter. Journal of lipid research 40 10744779
2020 Selective valorization of lignin to phenol by direct transformation of Csp2-Csp3 and C-O bonds. Science advances 39 33158871
2019 Sargassum fusiforme Fucoidan SP2 Extends the Lifespan of Drosophila melanogaster by Upregulating the Nrf2-Mediated Antioxidant Signaling Pathway. Oxidative medicine and cellular longevity 38 31485301
2018 Isolation and characterization of a novel phage Xoo-sp2 that infects Xanthomonas oryzae pv. oryzae. The Journal of general virology 38 30102145
2012 Role of the SP2 domain and its proteolytic cleavage in HIV-1 structural maturation and infectivity. Journal of virology 37 23055560
2003 Estimation of hormone receptor status in fine-needle aspirates and paraffin-embedded sections from breast cancer using the novel rabbit monoclonal antibodies SP1 and SP2. Diagnostic cytopathology 36 14506673
2017 High-Throughput Lipidomic and Transcriptomic Analysis To Compare SP2/0, CHO, and HEK-293 Mammalian Cell Lines. Analytical chemistry 35 27991764
2017 Cytotoxic and Antibacterial Compounds from the Coral-Derived Fungus Aspergillus tritici SP2-8-1. Marine drugs 35 29112138
2004 Sp2 DNA binding activity and trans-activation are negatively regulated in mammalian cells. The Journal of biological chemistry 35 14726517
2017 sp2 -Iminosugar α-glucosidase inhibitor 1-C-octyl-2-oxa-3-oxocastanospermine specifically affected breast cancer cell migration through Stim1, β1-integrin, and FAK signaling pathways. Journal of cellular physiology 34 28145580
2021 Single-atom nickel terminating sp2 and sp3 nitride in polymeric carbon nitride for visible-light photocatalytic overall water splitting. Chemical science 32 34163637
2014 Inhibition of tumor growth and angiogenesis by SP-2, an anti-lectin, galactoside-binding soluble 3 binding protein (LGALS3BP) antibody. Molecular cancer therapeutics 31 24552775
2018 Probing the Inhibitor versus Chaperone Properties of sp²-Iminosugars towards Human β-Glucocerebrosidase: A Picomolar Chaperone for Gaucher Disease. Molecules (Basel, Switzerland) 30 29673163
2008 Potential for false-positive staining with a rabbit monoclonal antibody to progesterone receptor (SP2): findings of the UK National External Quality Assessment Scheme for Immunocytochemistry and FISH highlight the need for correct validation of antibodies on introduction to the laboratory. American journal of clinical pathology 30 18285262
2023 LTF Induces Radioresistance by Promoting Autophagy and Forms an AMPK/SP2/NEAT1/miR-214-5p Feedback Loop in Lung Squamous Cell Carcinoma. International journal of biological sciences 29 37056929
2021 HNRNPD interacts with ZHX2 regulating the vasculogenic mimicry formation of glioma cells via linc00707/miR-651-3p/SP2 axis. Cell death & disease 28 33542193
1997 Aberrant methylation of an imprinted gene U2af1-rs1(SP2) caused by its own transgene. The Journal of biological chemistry 28 9083040
2020 Sp2 promotes invasion and metastasis of hepatocellular carcinoma by targeting TRIB3 protein. Cancer medicine 26 32160655
2013 Carbon-binding designer proteins that discriminate between sp2- and sp3-hybridized carbon surfaces. Langmuir : the ACS journal of surfaces and colloids 26 23510486
1985 Stereospecificity of SP1 and SP2 substance P receptors. Life sciences 25 2579310
2010 Specificity protein 2 (Sp2) is essential for mouse development and autonomous proliferation of mouse embryonic fibroblasts. PloS one 23 20221402
2005 Exploring the universality of unusual conformations of the 17-membered Pt(d(G*pG*)) macrochelate ring. Dependence of conformer formation on a change in bidentate carrier ligand from an sp3 to an sp2 nitrogen donor. Inorganic chemistry 22 16212391
1996 Primary structure of mannuronate lyases SP1 and SP2 from Turbo cornutus and involvement of the hydrophobic C-terminal residues in the protein stability. Journal of protein chemistry 22 9008294
2016 Oxidative coupling between C(sp²)-H and C(sp³)-H bonds of indoles and cyclic ethers/cycloalkanes. Organic & biomolecular chemistry 21 26842403
2006 Development of new rabbit monoclonal antibody to progesterone receptor (Clone SP2): no heat pretreatment but effective for paraffin section immunohistochemistry. Applied immunohistochemistry & molecular morphology : AIMM 21 16785796
2012 Rh-catalyzed intramolecular sp2 C-H bond difluoromethylenation. Chemical communications (Cambridge, England) 20 22327167
2006 Chemistry of HIV-1 virucidal Pt complexes having neglected bidentate sp2 N-donor carrier ligands with linked triazine and pyridine rings. synthesis, NMR spectral features, structure, and reaction with guanosine. Inorganic chemistry 19 16933918
2017 A Tandem Enzymatic sp2 -C-Methylation Process: Coupling in Situ S-Adenosyl-l-Methionine Formation with Methyl Transfer. Chembiochem : a European journal of chemical biology 18 28371017
2016 Ascorbic Acid-Assisted Synthesis of Mesoporous Sodium Vanadium Phosphate Nanoparticles with Highly sp(2) -Coordinated Carbon Coatings as Efficient Cathode Materials for Rechargeable Sodium-Ion Batteries. Chemistry (Weinheim an der Bergstrasse, Germany) 18 27346677
1997 Identification of the tumor antigen 90K domains recognized by monoclonal antibodies SP2 and L3 and preparation and characterization of novel anti-90K monoclonal antibodies. Biochemical and biophysical research communications 18 9125183
1996 Effects of ethanol on lateral and rotational mobility of plasma membrane vesicles isolated from cultured mouse myeloma cell line Sp2/0-Ag14. Biochimica et biophysica acta 18 8664314
2016 Expression of human gene coding RORγT receptor depends on the Sp2 transcription factor. Journal of leukocyte biology 17 27256574
2015 Enantioselective and Synergetic Toxicity of Axial Chiral Herbicide Propisochlor to SP2/0 Myeloma Cells. Journal of agricultural and food chemistry 17 26299567
2006 Oxidative stress is not required for the induction of apoptosis upon glutamine starvation of Sp2/0-Ag14 hybridoma cells. European journal of cell biology 17 16412532
2006 Sp2 localizes to subnuclear foci associated with the nuclear matrix. Molecular biology of the cell 17 16467376
2020 C(sp2)-H Bond Multiple Functionalization in Air for Construction of Tetrahydrocarbazoles with Continuous Quaternary Carbons and Polycyclic Diversification. Organic letters 15 32073283
2010 Fluorescent-tagged sp2-iminosugars with potent β-glucosidase inhibitory activity. Bioorganic & medicinal chemistry 15 20889348
2010 Overexpression of transcription factor sp2 inhibits epidermal differentiation and increases susceptibility to wound- and carcinogen-induced tumorigenesis. Cancer research 15 20959487
2005 Sp1 elements in SULT2B1b promoter and 5'-untranslated region of mRNA: Sp1/Sp2 induction and augmentation by histone deacetylase inhibition. FEBS letters 15 15953604
1996 Azidothymidine induces apoptosis in mouse myeloma cell line Sp2/0. Biochemical pharmacology 15 8781503
2024 CD93 regulates breast cancer growth and vasculogenic mimicry through the PI3K/AKT/SP2 signaling pathway activated by integrin β1. Journal of biochemical and molecular toxicology 14 38511888
2008 Gene analysis of an antiviral protein SP-2 from Chinese wild silkworm, Bombyx mandarina Moore and its bioactivity assay. Science in China. Series C, Life sciences 14 18815751
2021 ANLN, Regulated by SP2, Promotes Colorectal Carcinoma Cell Proliferation via PI3K/AKT and MAPK Signaling Pathway. Journal of investigative surgery : the official journal of the Academy of Surgical Research 13 33757382
2020 Sp2 regulates late neurogenic but not early expansive divisions of neural stem cells underlying population growth in the mouse cortex. Development (Cambridge, England) 13 32001437
2016 TransOmic analysis of forebrain sections in Sp2 conditional knockout embryonic mice using IR-MALDESI imaging of lipids and LC-MS/MS label-free proteomics. Analytical and bioanalytical chemistry 13 26942738
2016 Lysine acetylation stabilizes SP2 protein in the silkworm Bombyx mori. Journal of insect physiology 13 27374983
2016 Engineering the metal sensitive sites in Macrolampis sp2 firefly luciferase and use as a novel bioluminescent ratiometric biosensor for heavy metals. Analytical and bioanalytical chemistry 13 27815607
2013 Identification of a novel gene product expressed by Trichinella spiralis that binds antiserum to Sp2/0 myeloma cells. Veterinary parasitology 13 23465747
2009 Sp2 is a maternally inherited transcription factor required for embryonic development. The Journal of biological chemistry 13 19959469
2008 Easy access to bio-inspired osmium(II) complexes through electrophilic intramolecular C(sp2)-H bond cyclometalation. Inorganic chemistry 13 18459720
2008 Neglected bidentate sp2 N-donor carrier ligands with triazine nitrogen lone pairs: platinum complexes retromodeling cisplatin guanine nucleobase adducts. Inorganic chemistry 13 18800828
1995 Variation in N-linked carbohydrate chains in different batches of two chimeric monoclonal IgG1 antibodies produced by different murine SP2/0 transfectoma cell subclones. Glycoconjugate journal 13 7496147
1991 Cuttlefish spermatid-specific protein T. Molecular characterization of two variants T1 and T2, putative precursors of sperm protamine variants Sp1 and Sp2. The Journal of biological chemistry 13 1894625
2021 Complete genome sequence of the cellulolytic planctomycete Telmatocola sphagniphila SP2T and characterization of the first cellulolytic enzyme from planctomycetes. Systematic and applied microbiology 12 34735803
2021 Transcription factor Sp2 promotes TGFB-mediated interstitial cell osteogenic differentiation in bicuspid aortic valves through a SMAD-dependent pathway. Experimental cell research 12 34914964
2020 H-Bond Surrogate-Stabilized Shortest Single-Turn α-Helices: sp2 Constraints and Residue Preferences for the Highest α-Helicities. ACS omega 12 32566857
2018 Acinetobacter soli SP2 Capable of High-Efficiency Degradation of Food Emulsifier Polysorbate 80. Current microbiology 12 29478241
2014 Hypericin-mediated photodynamic therapy induces apoptosis of myoloma SP2/0 cells depended on caspase activity in vitro. Cancer cell international 12 26074732
2013 Identification of the differentially expressed genes in SP2/0 myeloma cells from Balb/c mice infected with Trichinella spiralis. Veterinary parasitology 12 23473833
2004 Steroidogenic acute regulatory (StAR)-directed immunotherapy protects against tumor growth of StAR-expressing Sp2-0 cells in a rodent adrenocortical carcinoma model. Endocrinology 12 14715709
1987 Action of the SP2-11 and SP3-11 fragments of substance P on rat peritoneal mast cells. Agents and actions 12 2440265
2017 Charge-Transfer-Induced para-Selective sp2 C-H Bond Activation of Arenes by Use of a Hypervalent Iodine Compound: A Theoretical Study. The Journal of organic chemistry 11 28225619
1985 Antigens apparently nonimmunogenic for xid/Y mice cause the development of specific splenic plasmablasts and of fusion partners for SP2 cells giving hybridomas that secrete antibodies. European journal of immunology 11 2412836
2024 Unveiling the enzymatic pathway of UMG-SP2 urethanase: insights into polyurethane degradation at the atomic level. Chemical science 10 39790984
2013 B7-H1 protein vaccine induces protective and therapeutic antitumor responses in SP2/0 myeloma-bearing mice. Oncology reports 10 23970300
2009 Estimation of hormone receptor status and HER2 in cytologic cell blocks from breast cancer using the novel rabbit monoclonal antibodies (SP1, SP2, and SP3). Diagnostic cytopathology 10 19530101
2005 Regulation of inhibitor of differentiation gene 3 (Id3) expression by Sp2-motif binding factor in myogenic C2C12 cells: downregulation of DNA binding activity following skeletal muscle differentiation. Biochimica et biophysica acta 10 16216350
2022 Advances in the site-selective C-5, C-3 and C-2 functionalization of chromones via sp2 C-H activation. RSC advances 9 35425362
2019 The evolution of the 9aaTAD domain in Sp2 proteins: inactivation with valines and intron reservoirs. Cellular and molecular life sciences : CMLS 9 31375868
2018 Transcription factor Sp2 potentiates binding of the TALE homeoproteins Pbx1:Prep1 and the histone-fold domain protein Nf-y to composite genomic sites. The Journal of biological chemistry 9 30337366
2012 A transcriptional survey of the cDNA library of Macrolampis sp2 firefly lanterns (Coleoptera: Lampyridae). Comparative biochemistry and physiology. Part D, Genomics & proteomics 9 23299360
2010 Transcription of mouse Sp2 yields alternatively spliced and sub-genomic mRNAs in a tissue- and cell-type-specific fashion. Biochimica et biophysica acta 9 20353838
2000 sp2-bridged diaryl retinoids: effects of bridge-region substitution on retinoid X receptor (RXR) selectivity. Bioorganic & medicinal chemistry letters 9 10890152
1997 Extension of Sp2/0 hybridoma cell viability through interleukin-6 supplementation. Biotechnology and bioengineering 9 18636502
1997 Expression of exogenous NF-IL6 induces apoptosis in Sp2/0-Ag14 myeloma cells. DNA and cell biology 9 9052733
2024 Melatonin alleviates di-butyl phthalate (DBP)-induced ferroptosis of mouse leydig cells via inhibiting Sp2/VDAC2 signals. Environmental research 8 38246300
2022 Comparative systeomics to elucidate physiological differences between CHO and SP2/0 cell lines. Scientific reports 7 35228567
1992 Xenopus laevis sperm proteins, previously identified as surface proteins with egg coat binding capability, are indeed histone H4, histone H3, and sperm specific protein SP2. The Journal of experimental zoology 7 1500885
2021 Strategy for Selective Csp2-F and Csp2-Csp2 Formations from Organoplatinum Complexes. Inorganic chemistry 6 33397093
2021 Src-1 and SP2 promote the proliferation and epithelial-mesenchymal transition of nasopharyngeal carcinoma. Open medicine (Warsaw, Poland) 6 34307888
2003 Effects of brevetoxins on murine myeloma SP2/O cells: aberrant cellular division. International journal of toxicology 6 12745987

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