Affinage

SP4

Transcription factor Sp4 · UniProt Q02446

Length
784 aa
Mass
82.0 kDa
Annotated
2026-04-28
89 papers in source corpus 34 papers cited in narrative 34 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

SP4 is a GC-box-binding zinc-finger transcription factor enriched in neurons that functions as both an activator and repressor of target genes to control dendritic patterning, synaptic receptor composition, and neuronal excitability. It activates transcription through an N-terminal glutamine-rich domain and binds the same GC-rich elements as SP1/SP3, yet displays distinct target selectivity — directly regulating genes encoding NMDA, AMPA, and GABAA receptor subunits, Na⁺/K⁺-ATPase subunits, neurotrophin-3, TRPV1, and serine racemase — and uniquely synergizes with the photoreceptor factor CRX on rod-specific promoters (PMID:7559627, PMID:15781457, PMID:24219545, PMID:24576410, PMID:26469128, PMID:30811405, PMID:32603878). SP4 protein stability is tightly regulated by STIM1/store-operated calcium entry–driven ubiquitination and proteasomal degradation, NMDA receptor–dependent dephosphorylation at S770 (which opposes a degradation signal), calpain cleavage, O-GlcNAcylation, and lithium-mediated stabilization (PMID:24894994, PMID:24475768, PMID:17316402, PMID:26431879, PMID:22017217). Sp4-null mice exhibit postnatal lethality, impaired hippocampal dentate gyrus development with reduced dendritic arborization, defective sensorimotor gating, cardiac conduction abnormalities, and loss of persistent pain sensitization, while SP4 protein is reduced in postmortem cerebellum of bipolar disorder subjects (PMID:8660867, PMID:16899055, PMID:15558077, PMID:16430881, PMID:30811405, PMID:22017217).

Mechanistic history

Synthesis pass · year-by-year structured walk · 23 steps
  1. 1995 High

    Establishing that SP4 is a transcriptional activator with shared DNA-binding specificity but distinct functional properties relative to SP1/SP3 resolved how three related factors could co-occupy GC-boxes yet exert non-redundant effects — SP4 activates via its glutamine-rich domain but, unlike SP1, cannot synergize through tandem sites, and SP3 competitively represses SP4 activity.

    Evidence Cotransfection in SP-factor-null Drosophila SL2 cells with deletion mutants and superactivation assay

    PMID:7559627

    Open questions at the time
    • Structural basis for failure of SP4 to synergize through adjacent sites unknown
    • No neuronal context tested
  2. 1996 High

    Demonstrating that Sp4-null mice suffer postnatal lethality, growth retardation, and male reproductive failure established that SP4 has essential, non-redundant physiological roles despite sharing DNA-binding specificity with ubiquitous SP1/SP3.

    Evidence Gene targeting by homologous recombination in mice; phenotypic characterization; in situ hybridization

    PMID:11532028 PMID:8660867

    Open questions at the time
    • Specific target genes mediating the knockout phenotypes unidentified at this point
    • Cell-type-autonomous versus systemic contributions unclear
  3. 2002 High

    Identification of the rod cGMP-phosphodiesterase β-subunit promoter as the first SP4-specific target — activated by SP4 but not SP1 or SP3 — answered how a factor with identical DNA-binding specificity achieves gene-selective transcriptional output.

    Evidence Promoter deletion/mutagenesis with transfection in Y79 retinoblastoma cells; EMSA

    PMID:11943774

    Open questions at the time
    • Mechanism of SP4-specific activation through the β/GC element not resolved
    • Photoreceptor in vivo confirmation pending
  4. 2005 High

    Discovery that SP4 physically interacts with CRX via their zinc-finger and homeodomain, respectively, and synergizes on the rod opsin promoter in vivo (ChIP) established a combinatorial mechanism for photoreceptor-specific gene regulation and explained SP4's unique role in retinal cells.

    Evidence Co-immunoprecipitation; ChIP on retinal chromatin; cotransfection assays; in situ hybridization

    PMID:15781457

    Open questions at the time
    • Whether the SP4–CRX interaction is direct or bridged by additional factors not fully resolved
    • Impact of SP4 loss on photoreceptor function in vivo not tested
  5. 2005 High

    Showing that Sp4 hypomorphic mice develop hippocampal vacuolization, reduced NT-3 expression, and sensorimotor gating deficits — all rescued by Cre-dependent Sp4 restoration — linked SP4 to specific neuropsychiatric-relevant circuits and identified neurotrophin-3 as an in vivo downstream target.

    Evidence Hypomorphic knock-in with Cre-dependent rescue; behavioral assays; in situ hybridization

    PMID:15558077

    Open questions at the time
    • Whether NT-3 reduction is cause or consequence of hippocampal pathology not yet tested
    • Direct SP4 binding to NT-3 promoter not demonstrated in this study
  6. 2006 High

    Cell-type-specific conditional knockouts revealed that SP4 is required in both cardiomyocytes and neural crest cells for cardiac conduction system development, expanding SP4's physiological roles beyond the CNS and identifying Cx40 and TrkC as lineage-specific downstream targets.

    Evidence Cre-Lox conditional knockouts (ventricular-specific, neural crest–specific); electrophysiology; immunohistochemistry

    PMID:16430881

    Open questions at the time
    • Whether SP4 directly binds Cx40 and TrkC promoters not established
    • Mechanistic link between TrkC deficiency and atrial dysfunction unclear
  7. 2006 High

    SP4 was shown to be essential for postnatal hippocampal dentate gyrus development, controlling granule cell proliferation and dendritic arborization — connecting the knockout phenotype to a specific developmental process.

    Evidence SP4 null knockout; BrdU incorporation; hippocampal neuronal cultures; immunohistochemistry

    PMID:16899055

    Open questions at the time
    • Specific transcriptional targets mediating proliferation vs. dendrite growth not distinguished
  8. 2007 High

    Demonstrating that SP4 is required for activity-dependent dendritic pruning in cerebellar granule neurons — and that this requires its DNA-binding domain — established SP4 as a transcription-dependent mediator of neuronal activity-to-morphology coupling.

    Evidence shRNA knockdown and overexpression of WT vs. DNA-binding-domain-deleted SP4 in dissociated cerebellar cultures and in vivo

    PMID:17535924

    Open questions at the time
    • Direct transcriptional targets mediating pruning not identified in this study
    • Whether SP4 acts cell-autonomously or via secreted factors unclear
  9. 2007 High

    Discovery that glutamate receptor–activated calpain cleaves SP4 (and SP3) into fragments retaining DNA-binding activity identified a rapid, calcium-dependent post-translational mechanism for altering SP4 function during excitotoxicity and ischemia.

    Evidence In vitro reconstitution with purified calpain I; EMSA supershift; calpain inhibitor rescue; in vivo ischemia model

    PMID:17316402

    Open questions at the time
    • Functional consequences of the truncated, DNA-binding-competent cleavage products unknown
    • Calpain cleavage site(s) on SP4 not mapped
  10. 2009 High

    Establishing that SP4 directly represses NT-3 transcription in cerebellar neurons (ChIP-confirmed) and that NT-3 sequestration blocks excess branching caused by SP4 loss placed SP4 in a specific transcriptional circuit controlling dendrite branch limitation versus pruning.

    Evidence ChIP; luciferase reporter; NT-3 sequestration rescue; SP4 overexpression/knockdown in cerebellar neurons

    PMID:17059557 PMID:19555762

    Open questions at the time
    • Whether SP4 switches from activator to repressor on NT-3 promoter depending on neuronal maturation state not tested
    • Chromatin mechanisms of SP4-mediated repression not identified
  11. 2011 High

    Showing that SP4 protein (but not mRNA) is reduced in bipolar disorder cerebellum and that SP4 undergoes proteasomal degradation stabilized by lithium provided a molecular link between SP4 turnover and psychiatric disease pathophysiology.

    Evidence Western blot of postmortem brain; polyubiquitination assay; proteasome inhibitor and lithium treatment in primary neurons

    PMID:22017217

    Open questions at the time
    • E3 ligase responsible for SP4 ubiquitination not identified
    • Whether lithium's effect is direct or through upstream kinase/phosphatase changes unclear
  12. 2013 High

    Comprehensive demonstration that SP4 directly binds and activates Na⁺/K⁺-ATPase subunit gene promoters (Atp1a1, Atp1a3, Atp1b1) in neurons, with activity-dependent regulation, expanded the SP4 target repertoire to ion homeostasis machinery central to neuronal excitability.

    Evidence EMSA; supershift; ChIP; promoter mutagenesis; overexpression/RNAi in primary neurons

    PMID:24219545

    Open questions at the time
    • Whether altered Na⁺/K⁺-ATPase levels contribute to SP4-knockout phenotypes not tested
  13. 2014 High

    Identification of STIM1-mediated store-operated calcium entry as the upstream signal driving SP4 ubiquitination and proteasomal degradation under resting conditions solved a key question about how neuronal activity state is translated into SP4 protein levels — depolarization blocks SOCE and stabilizes SP4.

    Evidence Ca²⁺ imaging; STIM1 knockdown and constitutively active STIM1; SOCE pharmacological inhibition; ubiquitylation assay in cerebellar granule neurons

    PMID:24894994

    Open questions at the time
    • E3 ubiquitin ligase downstream of SOCE/STIM1 not identified
    • Whether SOCE-dependent SP4 degradation occurs in non-cerebellar neurons unknown
  14. 2014 High

    Mapping NMDA receptor–dependent dephosphorylation of SP4 at S770 (via PP1/2A) and showing that phosphomimetic S770 impairs dendritic maturation while non-phosphorylatable S770A behaves as wild type identified a specific phosphosite integrating synaptic activity with SP4 transcriptional function.

    Evidence Phospho-specific antibody; phosphomimetic/non-phosphorylatable mutants; PP1/2A inhibitors; NMDA receptor pharmacology; neuronal morphometry

    PMID:24475768

    Open questions at the time
    • Kinase responsible for basal S770 phosphorylation not identified
    • Whether S770 phosphorylation affects DNA binding, protein stability, or cofactor recruitment not distinguished
  15. 2014 High

    Discovery that SP4 transcriptionally activates Nwk2 (Fchsd1), and that Nwk2 mediates SP4-dependent control of dendrite number and NR1 surface expression, established a direct transcriptional effector pathway linking SP4 to NMDA receptor trafficking and dendritic morphogenesis.

    Evidence ChIP; luciferase reporter; shRNA epistasis and rescue; NR1 surface biotinylation; dendritic morphometry

    PMID:25045015

    Open questions at the time
    • How Nwk2 mechanistically controls NR1 surface expression not resolved
    • Additional SP4 targets contributing to dendrite number regulation likely exist
  16. 2014 High

    Demonstrating that SP4 directly activates GluA2 (but not GluA1/3/4) expanded the SP4 target gene repertoire to include AMPA receptor subunit-selective regulation, showing SP4 coordinates multiple glutamate receptor systems.

    Evidence EMSA; supershift; ChIP; promoter mutagenesis; qRT-PCR; Western blot in neurons

    PMID:24576410

    Open questions at the time
    • Whether SP4-dependent GluA2 regulation affects synaptic AMPA receptor composition or plasticity not tested
  17. 2015 High

    Demonstrating that SP4 directly activates GABAA receptor subunit genes Gabra1 and Gabra2 (but not Gabra3) with functional effects on GABAergic currents established SP4 as a master regulator coordinating expression across excitatory and inhibitory receptor systems.

    Evidence EMSA; supershift; ChIP; promoter mutagenesis; overexpression/shRNA; electrophysiology

    PMID:26469128

    Open questions at the time
    • Whether SP4 coordinates excitatory/inhibitory balance through coordinated regulation of these targets remains untested
  18. 2015 Medium

    Cell-type-specific SP4 restoration in forebrain GABAergic (but not excitatory) neurons rescued ketamine-induced hyperlocomotion in SP4 hypomorphic mice, establishing that SP4's role in NMDA receptor-mediated behavioral responses is cell-autonomously required in inhibitory interneurons.

    Evidence Cre-LoxP cell-type-specific restoration; behavioral testing

    PMID:26037489

    Open questions at the time
    • Which SP4 target genes in GABAergic neurons mediate the behavioral rescue not identified
    • PPI rescue was not achieved, suggesting additional circuits involved
  19. 2015 Medium

    Establishing that O-GlcNAcylation of SP4's transactivation domain negatively regulates its transcriptional activity added a metabolic sensing layer to SP4 regulation and identified a post-translational modification distinct from ubiquitination and phosphorylation.

    Evidence Co-expression with OGT in E. coli; O-GlcNAc antibody; deletion mutagenesis; luciferase reporter

    PMID:26431879

    Open questions at the time
    • O-GlcNAc modification sites on SP4 not mapped
    • Functional impact in neurons not demonstrated
    • Interplay with other PTMs (phosphorylation, ubiquitination) not examined
  20. 2015 Medium

    Linking S770 phosphorylation elevation in bipolar disorder cerebellum and its correlation with negative symptoms in schizophrenia to SP4 protein instability connected the SP4 post-translational regulatory circuit to human psychiatric disease at a specific molecular level.

    Evidence Phospho-specific antibody on postmortem brain; phosphomimetic/non-phosphorylatable SP4 stability assay in cerebellar granule neurons

    PMID:26049820

    Open questions at the time
    • Causal relationship between S770 phosphorylation and disease not established
    • Sample sizes limited
    • Kinase responsible not identified
  21. 2019 Medium

    Showing that SP4 haploinsufficiency reduces TRPV1 expression in DRG neurons and prevents persistent inflammatory and neuropathic pain states extended SP4's physiological roles to peripheral pain circuits and identified TRPV1 as an SP4-regulated gene outside the CNS.

    Evidence Sp4+/- heterozygous mice; RT-PCR; calcium imaging; inflammatory and neuropathic pain behavioral models

    PMID:30811405

    Open questions at the time
    • Direct SP4 binding to TRPV1 promoter not confirmed by ChIP
    • Whether SP4 regulation of TRPV1 is direct or indirect uncertain
  22. 2024 Medium

    Discovery that SRSF3-dependent alternative splicing of SP4 exon 3 produces a long isoform (L-SP4) with the full transactivation domain that activates SMAD4 and suppresses renal cell carcinoma malignancy, while the short isoform lacks this capacity, revealed that SP4 functional output is regulated at the splicing level.

    Evidence SRSF3 overexpression; RT-PCR for alternative splicing; isoform-specific overexpression; SMAD4 reporter assay; cell malignancy assays

    PMID:39222664

    Open questions at the time
    • Prevalence of L-SP4 vs S-SP4 in neurons and other tissues unknown
    • Whether disease-associated SP4 changes reflect isoform switching not examined
  23. 2025 Medium

    Demonstration that SP4 recruits HDAC class IV (HD11) to the anti-apoptotic BCL-W promoter, causing histone H3 hypo-acetylation and BCL-W repression during copper-induced neurotoxicity, identified a specific epigenetic effector mechanism for SP4-mediated transcriptional repression.

    Evidence ChIP; co-immunoprecipitation; siRNA knockdown (single and double); Western blot; cell viability assay

    PMID:40185277

    Open questions at the time
    • Whether SP4–HDAC11 interaction occurs under physiological (non-toxic) conditions unknown
    • Generalizability of SP4-HDAC11 repressive mechanism to other SP4-repressed genes not tested

Open questions

Synthesis pass · forward-looking unresolved questions
  • The E3 ubiquitin ligase(s) mediating SOCE-dependent SP4 degradation, the kinase(s) phosphorylating S770, the structural basis for SP4's inability to synergize through tandem GC-boxes, and the genome-wide direct SP4 target repertoire in defined neuronal subtypes remain unresolved.
  • E3 ligase identity unknown
  • S770 kinase identity unknown
  • No genome-wide ChIP-seq or CUT&RUN in neurons published
  • Structural basis for SP4 vs SP1 functional divergence not determined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 14 GO:0003677 DNA binding 10
Localization
GO:0005634 nucleus 6
Pathway
R-HSA-112316 Neuronal System 7 R-HSA-1266738 Developmental Biology 4 R-HSA-392499 Metabolism of proteins 4
Partners
CRXHDAC11NRF1SP1SP3SRSF3STIM1

Evidence

Reading pass · 34 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1995 SP4 is a transcriptional activator that functions through its N-terminal glutamine-rich region, binds GC-rich DNA elements via three zinc fingers with the same specificity as SP1 and SP3, but unlike SP1 cannot act synergistically through adjacent binding sites. SP4 can serve as a target for SP1 activation domains in a superactivation assay, and SP3 can repress SP4-mediated transcriptional activation by competing for the same binding sites. Cotransfection in Drosophila SL2 cells lacking endogenous Sp factors; deletion analysis; superactivation assay The Journal of biological chemistry High 7559627
1996 SP4 is required for normal murine growth, viability, and male reproductive behavior; Sp4 null mice display postnatal lethality, growth retardation, and failure of males to copulate despite histologically intact testes. SP4 is expressed restrictively in the developing CNS, testes, and other tissues. Gene targeting by homologous recombination in mice; in situ hybridization; Northern blot Developmental biology High 8660867
1999 SP3 and SP4 zinc finger domains bind the same core GC-rich cis-elements as SP1 on the ADH5/FDH minimal promoter but cannot activate transcription in SP-factor-null Drosophila SL2 cells; both repress SP1-dependent transcriptional activation by competing for the same binding sites. Mutagenesis of promoter cis-elements; cotransfection in Drosophila SL2 cells; EMSA The Journal of biological chemistry High 9867805
2001 SP4 null mice (deletion of exons encoding N-terminal activation domains) show complete loss of SP4 expression, postnatal lethality in two-thirds of homozygotes, growth retardation, male breeding failure, and smaller thymus, spleen, and uterus in females with delayed sexual maturation, demonstrating that SP4's N-terminal glutamine-rich activation domains are required for its physiological functions. Gene targeting/knockout; Western blot for SP4 absence; phenotypic characterization Genes to cells High 11532028
2002 SP4, but not SP1 or SP3, specifically activates the rod cGMP-phosphodiesterase beta-subunit (beta-PDE) promoter through the beta/GC element (-59/-49), defining the first specific SP4 target gene and demonstrating that differential activation by SP4 versus other SP family members is possible despite shared DNA binding specificity. Promoter deletion and mutagenesis; transfection in Y79 retinoblastoma cells; EMSA The Journal of biological chemistry High 11943774
2005 SP4 activates the rod opsin and beta-PDE promoters in photoreceptors; SP1 and SP3 competitively repress SP4-mediated activation of the beta-PDE promoter. SP4 physically interacts with the photoreceptor transcription factor CRX via its zinc finger domain and CRX homeodomain, and synergizes with CRX on the rod opsin promoter. Both SP4 and CRX occupy the rod opsin and beta-PDE promoters in retinal chromatin (ChIP). Cotransfection assays; co-immunoprecipitation; chromatin immunoprecipitation; in situ hybridization and immunohistochemistry The Journal of biological chemistry High 15781457
2005 Reduced Sp4 expression in hypomorphic mice causes hippocampal vacuolization, age-dependent decrease in neurotrophin-3 expression in dentate granule cells, deficits in sensorimotor gating and contextual memory; all abnormalities are fully rescued by Cre-dependent restoration of Sp4 expression. Hypomorphic allele knock-in with Cre-dependent rescue; in situ hybridization; behavioral assays; histology Molecular psychiatry High 15558077
2005 COX-2 inhibitors (celecoxib, nimesulide, NS-398) decrease SP1 and SP4 protein levels in colon cancer cells via enhanced proteasome-dependent ubiquitination and degradation, without affecting SP1 or SP4 mRNA levels, thereby reducing VEGF transcription from the proximal GC-rich promoter region. Western blot; RT-PCR; proteasome inhibitor rescue (gliotoxin); ubiquitination assay; promoter-luciferase deletion analysis Molecular pharmacology High 15883203
2006 SP4 is required for postnatal development of the hippocampal dentate gyrus: SP4 null mice show restricted reduction in cell proliferation in hippocampus (but not cerebellum) during the first postnatal week, decreased dendritic growth and arborization of dentate granule cells in vitro, reduced dentate granule cell density, and reduced synaptophysin expression in the molecular layer. SP4 null knockout; BrdU proliferation assay; hippocampal neuronal cultures; immunohistochemistry; Western blot Genes, brain, and behavior High 16899055
2006 HF-1b/SP4 is required in both cardiomyogenic and neural crest cell lineages for cardiac conduction system development: ventricular-specific conditional deletion disrupts Cx40 expression, while neural crest-specific deletion causes atrial and atrioventricular dysfunction linked to deficiency in the neurotrophin receptor TrkC. Cre-Lox tissue-restricted conditional knockout; Cx40 immunohistochemistry; electrophysiology Developmental biology High 16430881
2007 SP4 knockdown in cerebellar granule neurons leads to increased dendritic branching and failure to resorb transient dendrites; depolarization-induced dendritic remodeling (pruning) requires SP4 and is blocked by SP4 knockdown. Overexpression of wild-type but not DNA-binding-domain-deleted SP4 promotes dendritic pruning in non-depolarizing conditions, establishing SP4's DNA-binding-dependent role in activity-dependent dendritic patterning. shRNA knockdown; overexpression of mutant SP4; dissociated cerebellar cultures and in vivo cerebellar cortex; morphometric dendrite analysis Proceedings of the National Academy of Sciences of the United States of America High 17535924
2007 Glutamate receptor activation causes calpain-mediated cleavage of SP3 and SP4 (the predominant SP-family members in neurons) into products that retain DNA-binding activity; this is calcium-dependent and blocked by calpain inhibitors. In vivo, SP4 activity is lost after cerebral ischemia/reperfusion. In vitro calpain cleavage assay with purified calpain I; Western blot; EMSA supershift; immunofluorescence; calpain inhibitor treatment Journal of neurochemistry High 17316402
2006 SP4 (and SP3) bind GC-boxes within neurotrophin-3 (NT-3) promoter B in cortical neurons; overexpression of SP4 increases NT-3 promoter activity more than other SP family members; dominant-negative SP3 and SP4 siRNA reduce NT-3 promoter activity, identifying NT-3 as a direct SP4 target gene in neurons. EMSA; ChIP; luciferase reporter assay; siRNA knockdown; overexpression in cortical neurons Journal of neurochemistry High 17059557
2009 SP4 directly represses neurotrophin-3 (NT3) transcription in cerebellar granule neurons: SP4 overexpression reduces NT3 promoter activity and mRNA; SP4 knockdown increases NT3 promoter activity and mRNA; SP4 binds NT3 promoter in vivo (ChIP). NT3 promotes dendritic branching, and sequestering NT3 blocks the increased branching caused by SP4 knockdown but not dendrite pruning, placing SP4-dependent NT3 repression specifically in the branch-limitation pathway. Luciferase reporter assay; RT-PCR; ChIP; NT3 sequestration; SP4 overexpression/knockdown in cerebellar neurons Molecular and cellular neurosciences High 19555762
2009 SP4 is constitutively bound to the GC-box of the argininosuccinate synthetase (AS) promoter in melanoma cells regardless of arginine availability. Under arginine depletion, interplay between c-Myc (positive), HIF-1alpha (negative), and SP4 (positive) regulates AS expression; SP4 ChIP confirmed direct promoter binding. ChIP assay; transfection/overexpression; promoter analysis Molecular cancer therapeutics Medium 19934275
2011 SP4 protein, but not mRNA, is reduced in the cerebellum of bipolar disorder subjects. In rat cerebellar granule neurons under non-depolarizing conditions, SP4 (but not SP1) is polyubiquitinated and degraded by the proteasome. Lithium stabilizes SP4 protein, providing a mechanism for lithium's effects on neuronal gene expression. Immunoblot of postmortem brain; RT-PCR; polyubiquitination assay; proteasome inhibitor treatment; lithium treatment in primary neurons Bipolar disorders High 22017217
2013 SP4 functionally regulates the Na+/K+-ATPase subunit genes Atp1a1, Atp1a3, and Atp1b1 in neurons: SP4 binds their promoters (EMSA, supershift, ChIP), promoter mutations abolish SP4-dependent activation, SP4 overexpression up-regulates these genes, and SP4 silencing blocks KCl-induced up-regulation and prevents tetrodotoxin-induced suppression from being rescued. SP4 binds conserved sites across mouse, rat, and human. EMSA; supershift assay; ChIP; promoter mutagenesis; overexpression; RNAi; RT-PCR; Western blot in primary neurons The European journal of neuroscience High 24219545
2014 Store-operated calcium entry (SOCE), maximally activated under resting/hyperpolarizing conditions via STIM1 redistribution into puncta at ER-plasma membrane junctions, promotes ubiquitylation and proteasomal degradation of SP4 in cerebellar granule neurons. SOCE blockers prevent SP4 degradation; STIM1 knockdown blocks SP4 degradation; constitutively active STIM1 decreases SP4 abundance even under depolarizing conditions. Ca2+ imaging; STIM1 knockdown/constitutively-active expression; SOCE pharmacological inhibition; ubiquitylation assay; immunoblot in cerebellar granule neurons Science signaling High 24894994
2014 SP4 activates transcription of Nervous Wreck 2 (Nwk2/Fchsd1) in cerebellar granule neurons; Nwk2 mediates SP4-dependent regulation of primary dendrite number and surface (but not total) NR1 cell-surface expression. Nwk2 knockdown phenocopies SP4 knockdown; exogenous Nwk2 rescues SP4 depletion; NR1 expression suppresses the increased dendrite number caused by SP4 or Nwk2 depletion. Luciferase reporter assay; ChIP; shRNA knockdown; overexpression rescue; NR1 surface biotinylation; dendritic morphometry Developmental neurobiology High 25045015
2014 SP4 is phosphorylated at serine 770 under basal conditions; membrane depolarization decreases this phosphorylation via protein phosphatase 1/2A. NMDA receptor stimulation reduces SP4 S770 phosphorylation (dependent on PP1/2A), while NMDA receptor inhibition increases it. A phosphomimetic S770 substitution impairs SP4-dependent maturation of cerebellar granule neuron primary dendrites, whereas a non-phosphorylatable mutant behaves like wild type. Phospho-specific antibody; phosphomimetic and non-phosphorylatable SP4 mutants; PP1/2A inhibitors; NMDA receptor pharmacology; neuronal morphometry Journal of neurochemistry High 24475768
2014 SP4 specifically regulates transcription of the AMPA receptor subunit GluA2 (Gria2) but not GluA1, 3, or 4, in neurons; SP4 binds the Gria2 promoter (EMSA, supershift, ChIP), and promoter mutations abolish SP4-dependent activation. SP4 and NRF-1 regulate Gria2 in a concurrent and parallel manner. SP4 overexpression up-regulates and SP4 shRNA down-regulates Gria2 mRNA and protein. EMSA; supershift; ChIP; promoter mutagenesis; qRT-PCR; Western blot; overexpression and shRNA in neurons Biochimica et biophysica acta High 24576410
2015 SP4 (the predominant neuronal SP1-like factor) binds and represses the 5-HT1A receptor gene promoter via the conserved -681 CpG site within a Sp1-like element. DNA methylation of this site attenuates SP4-induced repression. Chronic mild stress increases DNA methylation at -681, antagonizing SP4 repression to increase 5-HT1A expression despite increased SP4 levels. Luciferase reporter assay with SP4 overexpression; site-directed mutagenesis; bisulfite sequencing; ChIP; Western blot in prefrontal cortex and midbrain Neurobiology of disease High 26188176
2015 SP4 transcriptionally regulates GABAA receptor subunit genes Gabra1 and Gabra2 (but not Gabra3) in neurons: SP4 binds their promoters (EMSA, supershift, ChIP), promoter mutations reduce activity, SP4 overexpression up-regulates and SP4 shRNA down-regulates subunit expression, and functional GABA receptor currents are affected. EMSA; supershift; ChIP; promoter mutagenesis; qRT-PCR; Western blot; overexpression/shRNA; electrophysiology Biochimica et biophysica acta High 26469128
2015 O-GlcNAc modification is present on SP3 and SP4 (but not SP2), primarily in their N-terminal transactivation domains, as demonstrated by co-expression with O-GlcNAc transferase (OGT) in E. coli and O-GlcNAc antibody detection. O-GlcNAcylation of SP4 negatively regulates its transcriptional activity in reporter gene assays. Co-expression with OGT in E. coli; O-GlcNAc-specific antibody; GlcNAc-positive protein fraction enrichment; deletion mutagenesis; luciferase reporter assay; co-immunoprecipitation Biochemical and biophysical research communications Medium 26431879
2015 SP4 phosphorylation at S770 is increased in the cerebellum of bipolar disorder subjects and positively correlates with negative symptoms in schizophrenia. SP4 S770 phosphorylation inversely correlates with SP4 protein levels. A phosphomimetic mutation in truncated SP4 decreases its steady-state levels, while a non-phosphorylatable mutant shows increased stability, identifying S770 phosphorylation as a degradation signal. Phospho-specific antibody; Western blot of postmortem human brain; phosphomimetic/non-phosphorylatable SP4 expression in cerebellar granule neurons; pulse-chase/stability assay European neuropsychopharmacology Medium 26049820
2017 In methylmercury (MeHg)-treated neurons, p38 MAPK activation increases SP1 and SP4 protein expression, which in turn up-regulate HDAC4. SP4 (with SP1) drives HDAC4 expression; HDAC4 binds the BDNF promoter IV, reducing BDNF mRNA. siRNA knockdown of p38, SP1, SP4, or HDAC4 each partly prevent MeHg-induced neuronal death. Pharmacological p38 blockade reverses SP4 protein increase. siRNA knockdown; Western blot; ChIP (HDAC4 on BDNF promoter IV); p38 inhibitor; RT-PCR; cell viability assay Frontiers in neuroscience Medium 28154524
2019 SP4 positively regulates TRPV1 expression in dorsal root ganglion (DRG) neurons; Sp4+/- mice have reduced DRG TRPV1 mRNA and reduced neuronal responses to capsaicin. Sp4+/- mice fail to develop persistent inflammatory thermal hyperalgesia, persistent mechanical hypersensitivity to NGF, and persistent cold/mechanical hypersensitivity to oxaliplatin, identifying SP4 as a critical transcriptional regulator of persistent pain states. Sp4+/- heterozygous mice; RT-PCR; calcium imaging (capsaicin response); inflammatory and neuropathic pain behavioral models; IHC co-localization PloS one Medium 30811405
2020 SP4 controls constitutive expression of serine racemase (SR) in neurons by directly binding SP-binding elements in the Srr promoter (ChIP confirmed); mutagenesis of SP-binding elements reduces promoter activity. SP4 overexpression increases and SP4 knockdown decreases SR mRNA and protein. Valproic acid induces SR via Nrf2 binding to AREs, operating in parallel to SP4. Dual luciferase reporter assay; site-directed mutagenesis; ChIP; overexpression; siRNA knockdown; RT-PCR; Western blot Biochimica et biophysica acta. Gene regulatory mechanisms Medium 32603878
2023 EZH2 epigenetically activates SP4 expression by promoting methylation of the -170 bp CpG site in the SP4 promoter via DNMT3B recruitment in the IL-6/sIL-6R signaling pathway; SP4 then drives VEGF production and angiogenesis. EZH2 silencing/inhibition reduces DNMT3B and SP4 expression and VEGF levels. EZH2 overexpression/silencing/inhibition; bisulfite sequencing; Western blot; ELISA (VEGF); tube formation assay; in vivo rat peritoneal dialysis model International journal of medical sciences Medium 36619221
2024 SP4 activates PHF14 transcription by binding the PHF14 promoter region, thereby promoting PHF14 expression and activating Wnt/β-catenin signaling in esophageal squamous cell carcinoma; SP4 knockdown inhibits and PHF14 overexpression rescues proliferation and cell-cycle progression. ChIP; promoter luciferase assay; SP4 knockdown and PHF14 rescue; cell proliferation assay; in vivo xenograft Molecular cancer research Medium 37768180
2024 SRSF3 binding to SP4 exon 3 promotes inclusion of exon 3, producing a long SP4 isoform (L-SP4) that suppresses RCC cell malignancy by transcriptionally activating SMAD4; the short isoform (S-SP4), lacking the transactivation domain, lacks this function. L-SP4 specifically participates in SRSF3-mediated anti-proliferative effects. SRSF3 overexpression; RT-PCR for alternative splicing; L-SP4 vs S-SP4 overexpression; SMAD4 reporter assay; cell malignancy assays Biochimica et biophysica acta. Molecular cell research Medium 39222664
2025 In CuCl2-treated neurons, SP4 forms a complex with HDAC class IV (HD11) and co-localizes on the BCL-W (anti-apoptotic) promoter, causing histone H3 hypo-acetylation and BCL-W repression; simultaneously, SP1 forms a complex with HAT-p300 on the BAX promoter causing H4 hyper-acetylation and BAX activation. SP4 siRNA prevents HD11 binding to BCL-W and its down-regulation. Double knockdown of SP4+SP1 completely reverts CuCl2-induced neuronal death. ChIP; siRNA knockdown (single and double); Western blot; co-immunoprecipitation; cell viability assay Neurochemistry international Medium 40185277
2015 Restoration of SP4 expression specifically in forebrain GABAergic inhibitory neurons (but not excitatory neurons) rescues ketamine-induced hyperlocomotion in SP4 hypomorphic mice, demonstrating that SP4 function in GABAergic neurons is required for normal NMDA receptor-mediated behavioral responses. Cre-LoxP cell-type-specific SP4 restoration; behavioral testing (locomotion, PPI) The international journal of neuropsychopharmacology Medium 26037489
2001 SP4 and RORalpha exhibit negative cooperativity in regulating the prosaposin gene promoter in vivo; SP4 binds Sp1 sites within 310 bp upstream of the transcription start site. In SP4 knockout mice, the Sp1/U cluster deletion reduces reporter activity in the cerebellum of RORalpha-deficient staggerer mice but not in controls, indicating SP4 and RORalpha interact at the prosaposin locus. Transgenic reporter mice in SP4 KO and staggerer (RORalpha null) backgrounds; promoter deletion analysis DNA and cell biology Low 11879571

Source papers

Stage 0 corpus · 89 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1995 Functional analyses of the transcription factor Sp4 reveal properties distinct from Sp1 and Sp3. The Journal of biological chemistry 192 7559627
1996 Sp4, a member of the Sp1-family of zinc finger transcription factors, is required for normal murine growth, viability, and male fertility. Developmental biology 156 8660867
2009 Resistance to arginine deiminase treatment in melanoma cells is associated with induced argininosuccinate synthetase expression involving c-Myc/HIF-1alpha/Sp4. Molecular cancer therapeutics 112 19934275
2005 Cyclooxygenase-2 inhibitors decrease vascular endothelial growth factor expression in colon cancer cells by enhanced degradation of Sp1 and Sp4 proteins. Molecular pharmacology 101 15883203
1999 Sp3 and Sp4 can repress transcription by competing with Sp1 for the core cis-elements on the human ADH5/FDH minimal promoter. The Journal of biological chemistry 99 9867805
2016 Specificity protein (Sp) transcription factors Sp1, Sp3 and Sp4 are non-oncogene addiction genes in cancer cells. Oncotarget 81 26967243
1993 A novel, tissue-restricted zinc finger protein (HF-1b) binds to the cardiac regulatory element (HF-1b/MEF-2) in the rat myosin light-chain 2 gene. Molecular and cellular biology 77 8321243
2022 Solvent Precipitation SP3 (SP4) Enhances Recovery for Proteomics Sample Preparation without Magnetic Beads. Analytical chemistry 76 35848328
2005 Reduced expression of the Sp4 gene in mice causes deficits in sensorimotor gating and memory associated with hippocampal vacuolization. Molecular psychiatry 72 15558077
2004 Peroxisome proliferator-activated receptor gamma-dependent activation of p21 in Panc-28 pancreatic cancer cells involves Sp1 and Sp4 proteins. Endocrinology 69 15345676
1994 Positive regulatory elements (HF-1a and HF-1b) and a novel negative regulatory element (HF-3) mediate ventricular muscle-specific expression of myosin light-chain 2-luciferase fusion genes in transgenic mice. Molecular and cellular biology 66 8289802
2007 Transcription factor Sp4 regulates dendritic patterning during cerebellar maturation. Proceedings of the National Academy of Sciences of the United States of America 65 17535924
2006 Impaired postnatal development of hippocampal dentate gyrus in Sp4 null mutant mice. Genes, brain, and behavior 58 16899055
2014 Store-operated calcium entry promotes the degradation of the transcription factor Sp4 in resting neurons. Science signaling 54 24894994
2015 Chronic mild stress and antidepressant treatment alter 5-HT1A receptor expression by modifying DNA methylation of a conserved Sp4 site. Neurobiology of disease 52 26188176
2009 Transcription factor SP4 is a susceptibility gene for bipolar disorder. PloS one 52 19401786
2001 Complex phenotype of mice homozygous for a null mutation in the Sp4 transcription factor gene. Genes to cells : devoted to molecular & cellular mechanisms 51 11532028
2007 Glutamate receptor activation evokes calpain-mediated degradation of Sp3 and Sp4, the prominent Sp-family transcription factors in neurons. Journal of neurochemistry 44 17316402
2011 The transcription factor SP4 is reduced in postmortem cerebellum of bipolar disorder subjects: control by depolarization and lithium. Bipolar disorders 43 22017217
2009 Purification and concentration of a rhamnolipid biosurfactant produced by Pseudomonas aeruginosa SP4 using foam fractionation. Bioresource technology 40 19716289
2005 Sp4 is expressed in retinal neurons, activates transcription of photoreceptor-specific genes, and synergizes with Crx. The Journal of biological chemistry 40 15781457
2017 p38/Sp1/Sp4/HDAC4/BDNF Axis Is a Novel Molecular Pathway of the Neurotoxic Effect of the Methylmercury. Frontiers in neuroscience 38 28154524
2010 Reduced NMDAR1 expression in the Sp4 hypomorphic mouse may contribute to endophenotypes of human psychiatric disorders. Human molecular genetics 34 20634195
2005 Regulation of expression of the chorionic gonadotropin/luteinizing hormone receptor gene in the human myometrium: involvement of specificity protein-1 (Sp1), Sp3, Sp4, Sp-like proteins, and histone deacetylases. The Journal of clinical endocrinology and metabolism 33 15788387
2002 The rod cGMP-phosphodiesterase beta-subunit promoter is a specific target for Sp4 and is not activated by other Sp proteins or CRX. The Journal of biological chemistry 33 11943774
2022 Coexisting autoantibodies against transcription factor Sp4 are associated with decreased cancer risk in patients with dermatomyositis with anti-TIF1γ autoantibodies. Annals of the rheumatic diseases 32 36008132
2015 GlyT-1 Inhibition Attenuates Attentional But Not Learning or Motivational Deficits of the Sp4 Hypomorphic Mouse Model Relevant to Psychiatric Disorders. Neuropsychopharmacology : official publication of the American College of Neuropsychopharmacology 32 25907107
2014 From pre-DP, post-DP, SP4, and SP8 Thymocyte Cell Counts to a Dynamical Model of Cortical and Medullary Selection. Frontiers in immunology 30 24592261
1991 Isolation of cDNA for a Xenopus sperm-specific basic nuclear protein (SP4) and evidence for expression of SP4 mRNA in primary spermatocytes. Experimental cell research 29 2015853
2013 Regulation of Na(+)/K(+)-ATPase by neuron-specific transcription factor Sp4: implication in the tight coupling of energy production, neuronal activity and energy consumption in neurons. The European journal of neuroscience 28 24219545
2014 Increased SP4 and SP1 transcription factor expression in the postmortem hippocampus of chronic schizophrenia. Journal of psychiatric research 27 25175639
2013 Reduced expression of SP1 and SP4 transcription factors in peripheral blood mononuclear cells in first-episode psychosis. Journal of psychiatric research 27 23941741
2007 Sp3 and sp4 transcription factor levels are increased in brains of patients with Alzheimer's disease. Neuro-degenerative diseases 27 17934324
2009 Sp4-dependent repression of neurotrophin-3 limits dendritic branching. Molecular and cellular neurosciences 26 19555762
2006 Regulation of neurotrophin-3 gene transcription by Sp3 and Sp4 in neurons. Journal of neurochemistry 26 17059557
2013 Prolonged Ketamine Effects in Sp4 Hypomorphic Mice: Mimicking Phenotypes of Schizophrenia. PloS one 24 23823008
2006 Distinct roles of HF-1b/Sp4 in ventricular and neural crest cells lineages affect cardiac conduction system development. Developmental biology 24 16430881
2006 R5020 and RU486 act as progesterone receptor agonists to enhance Sp1/Sp4-dependent gene transcription by an indirect mechanism. Molecular endocrinology (Baltimore, Md.) 24 17192405
2001 Characterization and promoter analysis of the mouse gene for transcription factor Sp4. Gene 24 11245974
2006 lin-35/Rb and the CoREST ortholog spr-1 coordinately regulate vulval morphogenesis and gonad development in C. elegans. Developmental biology 22 17070797
2005 Knockout of the neural and heart expressed gene HF-1b results in apical deficits of ventricular structure and activation. Cardiovascular research 21 15907824
2014 Transcription factor Sp4 regulates expression of nervous wreck 2 to control NMDAR1 levels and dendrite patterning. Developmental neurobiology 20 25045015
2014 Specificity protein 4 (Sp4) regulates the transcription of AMPA receptor subunit GluA2 (Gria2). Biochimica et biophysica acta 18 24576410
2021 Over-representation of potential SP4 target genes within schizophrenia-risk genes. Molecular psychiatry 17 34750502
2014 Phosphorylation of the transcription factor Sp4 is reduced by NMDA receptor signaling. Journal of neurochemistry 17 24475768
2022 Metabolomics-Based Profiling, Antioxidant Power, and Uropathogenic Bacterial Anti-Adhesion Activity of SP4TM, a Formulation with a High Content of Type-A Proanthocyanidins. Antioxidants (Basel, Switzerland) 14 35883725
2020 Sp4 controls constitutive expression of neuronal serine racemase and NF-E2-related factor-2 mediates its induction by valproic acid. Biochimica et biophysica acta. Gene regulatory mechanisms 13 32603878
2015 Transcription factor SP4 phosphorylation is altered in the postmortem cerebellum of bipolar disorder and schizophrenia subjects. European neuropsychopharmacology : the journal of the European College of Neuropsychopharmacology 12 26049820
1999 A PEA3 site flanked by SP1, SP4, and GATA sites positively regulates the differentiation-dependent expression of Brachyury in embryonal carcinoma P19 cells. Biochemical and biophysical research communications 12 9920775
2015 Specificity protein 4 (Sp4) transcriptionally regulates inhibitory GABAergic receptors in neurons. Biochimica et biophysica acta 11 26469128
2005 SP4, a novel anti-cyclin D1 rabbit monoclonal antibody, is a highly sensitive probe for identifying mantle cell lymphomas bearing the t(11;14)(q13;q32) translocation. Applied immunohistochemistry & molecular morphology : AIMM 11 16280660
2019 Transcription factor Sp4 is required for hyperalgesic state persistence. PloS one 10 30811405
2015 O-GlcNAc modification of Sp3 and Sp4 transcription factors negatively regulates their transcriptional activities. Biochemical and biophysical research communications 10 26431879
2024 Differences in Protein Capture by SP3 and SP4 Demonstrate Mechanistic Insights of Proteomics Cleanup Techniques. Journal of proteome research 8 39161190
2023 EZH2 promotes angiogenesis in peritoneal dialysis by epigenetically activating SP4 expression in the IL-6/sIL-6R signalling pathway. International journal of medical sciences 8 36619221
2021 Thalidomide Suppresses Angiogenesis Through the Signal Transducer and Activator of Transcription 3/SP4 Signaling Pathway in the Peritoneal Membrane. Frontiers in physiology 8 34539435
2012 Genetics and expression analysis of the specificity protein 4 gene (SP4) in patients with Alzheimer's disease and frontotemporal lobar degeneration. Journal of Alzheimer's disease : JAD 8 22614877
2020 Characterisation of pectin and optimization of pectinase enzyme from novel Streptomyces fumigatiscleroticus VIT-SP4 for drug delivery and concrete crack-healing applications: An eco-friendly approach. Saudi journal of biological sciences 7 33304164
2007 Association of the Asn306Ser variant of the SP4 transcription factor and an intronic variant in the beta-subunit of transducin with digenic disease. Molecular vision 7 17356515
2024 The Novel A-Type Proanthocyanidin-Rich Phytocomplex SP4™ Acts as a Broad-Spectrum Antiviral Agent against Human Respiratory Viruses. International journal of molecular sciences 6 39000477
2022 Dual bio-degradative pathways of di-2-ethylhexyl phthalate by a novel bacterium Burkholderia sp. SP4. World journal of microbiology & biotechnology 6 36526923
2018 Ketamine independently modulated power and phase-coupling of theta oscillations in Sp4 hypomorphic mice. PloS one 6 29513708
1999 New phenolic antioxidants of PYA and PYE class increase the resistance S. cerevisiae strain SP4, its SOD- and catalase-deficient mutants to lipophilic oxidants. Folia microbiologica 6 11097024
2025 [Electroacupuncture of "Gongsun" (SP4) alleviates oxidative stress injury and promotes normal follicle development by regulating keap1/Nrf2/HO-1 signaling in rats with premature ovarian insufficiency]. Zhen ci yan jiu = Acupuncture research 5 39961757
2015 Restoration of Sp4 in Forebrain GABAergic Neurons Rescues Hypersensitivity to Ketamine in Sp4 Hypomorphic Mice. The international journal of neuropsychopharmacology 5 26037489
2015 Role played by the SP4 gene in schizophrenia and major depressive disorder in the Han Chinese population. The British journal of psychiatry : the journal of mental science 5 26450579
2009 [Association of Sp4 gene polymorphism with pathological tortuosity of internal carotid arteries]. Kardiologiia 5 19656107
2021 Overlapping and non-overlapping roles of the class-I histone deacetylase-1 corepressors LET-418, SIN-3, and SPR-1 in Caenorhabditis elegans embryonic development. Genes & genomics 4 33740234
1995 Structure of genes for sperm-specific nuclear basic protein (SP4) in Xenopus laevis. Biochimica et biophysica acta 4 8541323
2024 SP4 Facilitates Esophageal Squamous Cell Carcinoma Progression by Activating PHF14 Transcription and Wnt/Β-Catenin Signaling. Molecular cancer research : MCR 3 37768180
2022 The FOXP4-AS1/miR-3130-3p/SP4 feedback loop is associated with prostate cancer. Cellular and molecular biology (Noisy-le-Grand, France) 3 37114256
2021 Truncation of 3' CCND1 by t(11;22) leads to negative SP4 CCND1 immunohistochemistry in blastoid mantle cell lymphoma. Blood advances 3 33570637
1981 Placenta-associated plasma protein-A (PAPP-A, SP4) in trophoblastic tumours. Acta pathologica et microbiologica Scandinavica. Section A, Pathology 3 6164245
2025 Sp4/HD11 and Sp1/HAT-p300 complexes induce apoptotic cell death in CuCl2-treated neurons by modulating histone acetylation on BCL-W and BAX promoters. Neurochemistry international 2 40185277
2024 SRSF3 suppresses RCC tumorigenesis and progression via regulating SP4 alternative splicing. Biochimica et biophysica acta. Molecular cell research 2 39222664
2023 SPR-1/CoREST facilitates the maternal epigenetic reprogramming of the histone demethylase SPR-5/LSD1. Genetics 2 36655746
2022 The genetic polymorphisms in the SP4 gene and the risk of gastric cancer. Future oncology (London, England) 2 36346067
2017 Cloning, expression and identification of KTX-Sp4, a selective Kv1.3 peptidic blocker from Scorpiops pococki. Cell & bioscience 2 29142737
2001 In vivo roles of RORalpha and Sp4 in the regulation of murine prosaposin gene. DNA and cell biology 2 11879571
1995 The occurrence of a gene-encoded variant of nuclear basic protein (SP4) in sperm of Xenopus laevis. Biochemical and biophysical research communications 2 7677774
2025 NXF1 suppresses progression of endometrial cancer by interacting with the SRSF3 to regulate SP4 splicing. iScience 1 40792019
2024 The Relationship Between Anti-Cell Division Cycle and Apoptosis Regulator 1 Autoantibodies, Anti-Sp4 Autoantibodies, and Cancer in Anti-Transcription Intermediary Factor 1γ-Positive Dermatomyositis. ACR open rheumatology 1 39370373
2026 Differential SP4 expression and HSP60 abundance in buccal swabs from patients with schizophrenia. Science advances 0 41779859
2025 Anti-Sp4 and anti-CCAR1 autoantibodies in UK vs US patients with adult and juvenile-onset anti-TIF1γ-positive myositis. Rheumatology (Oxford, England) 0 39514366
2025 Partial rescue of schizophrenia-related phenotypes in young adult Sp4 hypomorphic mice. Journal of psychiatric research 0 40339225
2024 Differences in Protein Capture by SP3 and SP4 Demonstrate Mechanistic Insights of Proteomics Clean-up Techniques. bioRxiv : the preprint server for biology 0 38559195
2024 Efficacy of the combination of water aerobics and metacognitive training on psychological and physical health variables and their relationship with SP1 and SP4 biomarkers in people with psychosis: a study protocol. Frontiers in psychology 0 38919799
2022 Sp4 Regulates PTTG1IP Gene Transcription and Expression. DNA and cell biology 0 36383136
1996 Relative amounts of basic nuclear proteins SP4 and SP5 in Xenopus laevis sperm correlate with gene copy number. Development, growth & differentiation 0 37281109