Affinage

Showing SPAG9 — JIP4 is a alias.

SPAG9

C-Jun-amino-terminal kinase-interacting protein 4 · UniProt O60271

Length
1321 aa
Mass
146.2 kDa
Annotated
2026-06-10
63 papers in source corpus 25 papers cited in narrative 26 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 9/9 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

SPAG9 (JLP) is a multifunctional scaffolding protein that physically tethers stress-activated MAPK signaling modules — JNK and p38 — to upstream kinases (MKK4, MEKK3) and downstream transcription factors (Max, c-Myc), assembling pathway-specific signaling complexes (PMID:12391307). It binds JNK isoforms (JNK3 > JNK2 > JNK1) through a defined JNK-binding domain, with its leucine-zipper/coiled-coil and transmembrane regions governing dimerization and subcellular localization (PMID:15693750). A second major role is in microtubule-based transport: SPAG9 binds kinesin light chain 1 (KLC1) and kinesin-1 heavy chain to form motor-cargo complexes, with this interaction required for cytoplasmic localization and for ternary complex assembly with JNK (PMID:15987681). Through these motor linkages SPAG9, together with the redundant scaffold JSAP1, drives kinesin-1-dependent axonal transport whose loss causes cargo accumulation, JNK hyperactivation, and progressive neuronal death (PMID:25571974, PMID:26320416), delivers active ARF6 to the midbody during cytokinesis (PMID:25130574), controls endosome-to-TGN trafficking of furin cargo via PIKfyve (PMID:19056739), and positions lysosomes to support autophagosome-lysosome fusion (PMID:32023558, PMID:31787236). SPAG9 couples cell-surface receptors to intracellular cascades, partnering with Cdo to enhance p38α/β activation during myoblast differentiation (PMID:17074887) and mediating CD40 internalization in B lymphocytes through Rab5- and dynein-dependent vesicle transport (PMID:25586186). Upon extracellular stimulation it recruits PLK1 to the centrosome in a p38- and dynein-dependent manner to drive centrosome maturation, microtubule array formation, and microtubule-based NF-κB p65 nuclear translocation (PMID:31803841). In immune and developmental contexts SPAG9 also supports TCR-initiated Ca2+ influx and NF-AT activation in CD4+ T cells (PMID:28521278) and retinoic-acid-induced endodermal differentiation downstream of Gα13 (PMID:16619266). SPAG9 localizes to the sperm acrosome and relocates to the equatorial segment after the acrosome reaction, and its disruption in mice causes male subfertility (PMID:15693750, PMID:18574703). In disease, SPAG9 sustains MAPK-driven proliferation and invasion across multiple cancers (PMID:27655714, PMID:27042099, PMID:30720101), and a SPAG9::JAK2 fusion from t(9;17) constitutively activates JAK2 to phosphorylate STAT1/3/5 and confer growth-factor-independent proliferation (PMID:35879405); its transcriptional repression by TGF-β1/β-catenin/LEF1 contributes to renal fibrosis (PMID:41505216).

Mechanistic history

Synthesis pass · year-by-year structured walk · 15 steps
  1. 2002 High

    Established SPAG9/JLP as a MAPK scaffold, answering how JNK/p38 modules are spatially organized with upstream kinases and target transcription factors.

    Evidence Co-immunoprecipitation and interaction assays in transfected cells

    PMID:12391307

    Open questions at the time
    • Stoichiometry and structure of the assembled scaffold complex not resolved
    • Physiological signals triggering complex assembly not defined
  2. 2005 High

    Mapped the JNK-binding domain and the domains controlling dimerization/localization, defining how SPAG9 selectively engages JNK over p38α or ERK, and identified an acrosomal pool linked to sperm-egg binding.

    Evidence Deletion mutants, CD spectroscopy, immunofluorescence, and hemizona/egg penetration antibody-inhibition assays

    PMID:15693750

    Open questions at the time
    • How a single scaffold reconciles JNK-selectivity with reported p38 binding not resolved
    • Molecular basis of acrosomal targeting unknown
  3. 2005 High

    Identified KLC1 as a kinesin-1 link, explaining how SPAG9 achieves cytoplasmic localization and couples motor transport to JNK signaling.

    Evidence Yeast two-hybrid, reciprocal co-IP, site-directed mutagenesis, and dominant-negative mislocalization assays

    PMID:15987681

    Open questions at the time
    • Cargoes transported by the SPAG9-KLC1-JNK complex not defined here
    • Regulation of motor engagement unknown
  4. 2006 High

    Connected SPAG9 to cell-surface receptor signaling and developmental differentiation, showing it relays Cdo and Gα13 inputs into p38/JNK cascades.

    Evidence Co-IP, p38 activity assays, Cdo-/- primary myoblasts with MKK6 rescue, and siRNA in P19 cells

    PMID:16619266 PMID:17074887

    Open questions at the time
    • Direct vs indirect Cdo-SPAG9 contact regions not fully mapped
    • Gα13-SPAG9 interaction is single-lab
  5. 2008 Medium

    Demonstrated an in vivo physiological requirement, with male subfertility on Jlp disruption establishing relevance to spermatogenesis.

    Evidence Mouse gene knockout, immunohistochemistry, and fertility testing

    PMID:18574703

    Open questions at the time
    • Molecular defect in knockout spermatids not defined
    • Single lab
  6. 2008 High

    Defined a cargo-specific trafficking function, showing SPAG9-PIKfyve is needed for microtubule-based endosome-to-TGN transport of furin but not microtubule-independent cargo.

    Evidence Yeast two-hybrid, pulldown, co-IP, siRNA depletion with ectopic rescue, and defined cargo-trafficking assays

    PMID:19056739

    Open questions at the time
    • How cargo specificity is encoded not resolved
    • Link to JNK scaffolding role unclear
  7. 2014 High

    Extended the motor-scaffold role to cytokinesis and UV stress, showing SPAG9 delivers ARF6 to the midbody and is required for UVB-induced p38-dependent keratinocyte apoptosis.

    Evidence Knockout MEFs/mice with binding-deficient mutant rescue, co-IP, cytokinesis assays, and p38 inhibitor treatment in skin

    PMID:24520900 PMID:25130574

    Open questions at the time
    • Functional redundancy with JSAP1 complicates single-protein attribution
    • Upstream activator of midbody localization not defined
  8. 2015 High

    Resolved the consequences of transport failure in neurons and the receptor-internalization role, showing kinesin-1-dependent axonal transport requires SPAG9/JSAP1 and that its loss drives dynein-mediated nuclear JNK relocalization and neuronal death, while CD40 internalization needs SPAG9 with Rab5/dynein.

    Evidence Conditional double-knockout mice, primary neuron rescue, kinesin-1 interaction assays, jlp-deficient B cells, and dynein inhibitor (ciliobrevin D)

    PMID:25571974 PMID:25586186 PMID:26320416

    Open questions at the time
    • Cargo selectivity among kinesin-1 substrates incompletely defined
    • How the same scaffold engages both kinesin and dynein not mechanistically resolved
  9. 2016 Medium

    Linked SPAG9-JNK complex assembly and translocation to oncogenic signaling, showing LPA-stimulated perinuclear relocalization drives cancer proliferation and invasion, with ELK1 placed downstream in HCC.

    Evidence Co-IP, co-localization, shRNA/siRNA knockdown with ELK1 rescue, and xenograft models

    PMID:27042099 PMID:27655714

    Open questions at the time
    • Single-lab cancer-context findings
    • Direct vs indirect ELK1 regulation not established
  10. 2018 Medium

    Established a cytoprotective JNK-dependent function, showing the JNK-binding domain is required for SPAG9-mediated protection against oxidative cell death.

    Evidence siRNA knockdown with wild-type vs JNK-binding-domain mutant rescue, viability and ROS assays

    PMID:29753743

    Open questions at the time
    • Single lab
    • Mechanism linking JNK scaffolding to ROS control not defined
  11. 2019 High

    Unified the centrosome, lysosome, and pathway-specificity roles, showing SPAG9 tethers PLK1 to the centrosome for microtubule array formation and NF-κB nuclear import, positions lysosomes via kinesin-1 to enable autophagy, and engages an MKK3/p38 (not MKK6) axis in liver cancer.

    Evidence Live imaging, domain deletion, p38/dynein inhibitors, NF-κB transport assays, autophagy flux with mutant rescue, and co-IP with pathway Western blots

    PMID:30720101 PMID:31497238 PMID:31787236 PMID:31803841 PMID:32023558

    Open questions at the time
    • How SPAG9 selects MKK3 over MKK6 not defined
    • Prostate cancer MAPK link is Western-blot only (Low confidence)
  12. 2017 Medium

    Defined a non-MAPK immune role, showing SPAG9 supports TCR-initiated Ca2+ influx and NF-AT activation independently of MAPK/NF-κB/AP-1.

    Evidence jlp-deficient CD4+ T cells, proliferation/cytokine assays, Ca2+ flux, NF-AT reporter, and pathway Western blots

    PMID:28521278

    Open questions at the time
    • Mechanism coupling SPAG9 to Ca2+/NF-AT unknown
    • Single lab
  13. 2023 Medium

    Identified a transcription-factor-tethering anti-EMT function, showing SPAG9 sequesters Foxk1 to relieve repression of N-cadherin and counteract TGF-β1-induced partial EMT.

    Evidence TGF-β1 treatment, JLP knockdown/overexpression, Foxk1 modulation, and N-cadherin expression assays

    PMID:37013185

    Open questions at the time
    • Direct Foxk1-SPAG9 binding interface not mapped
    • Single lab
  14. 2022 High

    Revealed an oncogenic fusion mechanism, showing SPAG9::JAK2 constitutively activates JAK2 to drive STAT1/3/5 phosphorylation and a STAT1-BCL-2/MCL-1 survival axis conferring cytokine-independent growth.

    Evidence Ba/F3 reconstitution, IL-3-independent growth, kinase-domain mutagenesis, and ruxolitinib treatment

    PMID:35879405

    Open questions at the time
    • Contribution of SPAG9 dimerization domains to fusion activation not dissected
    • Prevalence in patients not addressed by these experiments
  15. 2026 Medium

    Placed SPAG9 as a transcriptionally regulated brake on fibrosis-associated autophagy, showing TGF-β1/β-catenin/LEF1 represses SPAG9 to unleash autophagy and renal fibrosis, reversible by β-catenin/LEF1 inhibition.

    Evidence CKD patient tissue, murine fibrosis models, HK-2 cells, iCRT3/LEF1 silencing, and JLP expression/autophagy assays

    PMID:41505216

    Open questions at the time
    • Direct LEF1 binding at the SPAG9 promoter not fully characterized
    • Single lab

Open questions

Synthesis pass · forward-looking unresolved questions
  • How a single scaffold integrates seemingly contradictory roles — selective JNK vs p38 binding, simultaneous engagement of kinesin and dynein, and pro- vs anti-survival outputs — across cell types remains unresolved.
  • No structural model of the full SPAG9 scaffold or its cargo/motor interfaces
  • Context-determinants selecting among partners not defined
  • Relative contributions of SPAG9 vs redundant JSAP1 in vivo incompletely partitioned

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0008092 cytoskeletal protein binding 4 GO:0098772 molecular function regulator activity 4 GO:0060090 molecular adaptor activity 3
Localization
GO:0005764 lysosome 2 GO:0005768 endosome 2 GO:0005829 cytosol 2 GO:0005856 cytoskeleton 2 GO:0005815 microtubule organizing center 1
Pathway
R-HSA-162582 Signal Transduction 4 R-HSA-5653656 Vesicle-mediated transport 4 R-HSA-9612973 Autophagy 3 R-HSA-1266738 Developmental Biology 2 R-HSA-168256 Immune System 2 R-HSA-1640170 Cell Cycle 1 R-HSA-1643685 Disease 1
Partners
ARF6CDOJNK3KLC1MEKK3MKK4PIKFYVEPLK1

Evidence

Reading pass · 26 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2002 JLP (SPAG9) acts as a scaffolding protein that simultaneously tethers JNK and p38MAPK signaling modules with transcription factors Max and c-Myc, as well as their upstream kinases MKK4 and MEKK3, bringing MAPKs and their target transcription factors together for specific signaling pathways. Co-immunoprecipitation, protein interaction assays, transient transfection Proceedings of the National Academy of Sciences of the United States of America High 12391307
2005 SPAG9 interacts with JNK isoforms (JNK3 > JNK2 > JNK1) through its JNK-binding domain; both the leucine zipper/coiled-coil domain and transmembrane domain are required for dimerization and proper subcellular localization. No interaction was detected with p38alpha or ERK pathways. Co-immunoprecipitation, deletion mutant expression, CD spectroscopy, indirect immunofluorescence The Biochemical journal High 15693750
2005 SPAG9 protein localizes to the acrosomal compartment of intact human spermatozoa and relocates to the equatorial segment after acrosome reaction; anti-SPAG9 antibodies inhibit binding of human spermatozoa to intact human oocytes and hemizona, implicating SPAG9 in sperm-egg interaction. Indirect immunofluorescence, hemizona assay, zona-free hamster egg penetration test The Biochemical journal Medium 15693750
2005 JLP (SPAG9) interacts with kinesin light chain 1 (KLC1) through JLP's second leucine zipper domain and a novel leucine zipper-like domain within KLC1's tetratripeptide repeat; mutations at Leu-280, Leu-287, Val-294, and Leu-301 in KLC1 disrupted the interaction. The JLP-KLC1 association is required for formation of a ternary complex with JNK1 and for cytoplasmic localization of JLP. Yeast two-hybrid screening, co-immunoprecipitation, site-directed mutagenesis, immunofluorescence, dominant-negative KLC1 mislocalization assay The Journal of biological chemistry High 15987681
2006 JLP (SPAG9) physically interacts with the intracellular region of cell-surface receptor Cdo, forming a complex with p38alpha/beta MAPK in differentiating myoblasts; Cdo and JLP cooperate to enhance active p38alpha/beta levels. Primary myoblasts from Cdo-/- mice show deficient p38alpha/beta activity that can be rescued by activated MKK6, placing JLP-Cdo at the top of this differentiation-linked p38 signaling cascade. Co-immunoprecipitation, transfection-based p38 activity assays, Cdo-/- primary myoblasts, activated MKK6 rescue The Journal of cell biology High 17074887
2006 JLP (SPAG9) physically associates with Galpha13 in RA-stimulated P19 cells and is required for retinoic acid-induced endodermal differentiation; siRNA silencing of JLP abrogates RA-mediated endodermal differentiation analogously to silencing of Galpha13 or JNK. Co-immunoprecipitation, siRNA knockdown, differentiation assays in P19 cells Journal of cellular biochemistry Medium 16619266
2008 Ablation of JLP (Jlp gene disruption) in male mice causes subfertility, with highest JLP expression in elongated spermatids of the testis, indicating a physiological role for JLP in production of functionally normal spermatozoa. Gene knockout in mice, immunohistochemistry, fertility testing Transgenic research Medium 18574703
2008 JLP (SPAG9) interacts with the phosphoinositide kinase PIKfyve through PIKfyve's cpn60_TCP1 consensus sequence and the last 75 residues of the JLP C-terminus; both proteins are required for microtubule-based endosome-to-TGN transport of furin cargo, but not for microtubule-independent Tac-TGN38 trafficking. Yeast two-hybrid, pulldown, co-immunoprecipitation, siRNA depletion, ectopic rescue, cargo-trafficking assays (Tac-furin, Tac-TGN38) The Journal of biological chemistry High 19056739
2014 JSAP1 and JLP (SPAG9) are required for ARF6 localization to the midbody during cytokinesis; both proteins form a tripartite complex with active ARF6 and kinesin-1 heavy chain (KHC). Mutants of JSAP1 or JLP unable to interact with active ARF6 or KHC failed to rescue ARF6 midbody localization and cytokinesis defects in double-knockout MEFs. Gene knockout in MEFs, rescue experiments with wild-type and binding-deficient mutants, co-immunoprecipitation, cytokinesis assays Genes to cells : devoted to molecular & cellular mechanisms High 25130574
2014 JLP (SPAG9) scaffold protein is required for UVB-induced p38 MAPK activation in skin keratinocytes; Jlp knockout mice show impaired p38 activation and resistance to UVB-induced epidermal apoptosis, while topical p38 inhibitor suppresses apoptosis in wild-type but not Jlp KO skin. Jlp knockout mice (whole-body and keratinocyte-specific), p38 MAPK activity assays, p38 inhibitor treatment, apoptosis assays Genes to cells : devoted to molecular & cellular mechanisms High 24520900
2015 JLP (SPAG9) and JSAP1 are essential regulators of kinesin-1-dependent axonal transport with functional redundancy; double-KO of Jsap1 and Jlp in dorsal telencephalon causes progressive neuron loss. JLP binding to kinesin-1 heavy chain is crucial for kinesin-1–microtubule interactions. Defective transport in dKO neurons leads to JNK hyperactivation via increased intra-axonal Ca2+, and axonal JNK relocalizes to the nucleus via dynein to activate c-Jun and cause neuronal death. Conditional double-knockout mice, primary neuron culture with inducible gene disruption, gene rescue experiments, kinesin-1 interaction assays Cell death and differentiation High 25571974
2015 JLP (SPAG9) scaffold protein is required in B lymphocytes for CD40 internalization upon CD154 engagement; JLP deficiency impairs CD40 internalization through a process dependent on Rab5 and dynein-mediated vesicle transport. JLP deficiency also diminishes CD40-dependent MAPK and JNK activation but not NF-κB. jlp-deficient mice B lymphocytes, dynein inhibitor (ciliobrevin D), co-localization and interaction assays for CD40/JLP/dynein/Rab5, flow cytometry The Journal of biological chemistry High 25586186
2015 JSAP1 and JLP (SPAG9) play critical and redundant roles in kinesin-1-dependent axonal transport in cerebellar Purkinje cells; double-KO causes PC axonal dystrophy with selective accumulation of kinesin-1 cargoes in swollen axons and progressive neuronal loss. Conditional double-knockout mice, immunohistochemistry, axonal transport cargo analysis FEBS letters Medium 26320416
2016 JLP (SPAG9) interacts with JNK in vivo in ovarian cancer cells, and this interaction is stimulated by lysophosphatidic acid (LPA); LPA stimulates translocation of the JLP-JNK complex to the perinuclear region. JLP knockdown abrogates LPA-stimulated JNK activation as well as LPA-stimulated proliferation and invasive migration. Co-immunoprecipitation, co-localization, shRNA knockdown, xenograft mouse model Oncotarget Medium 27655714
2016 SPAG9 knockdown in hepatocarcinoma cells suppresses migration and invasion, and inhibits ELK1 expression and p38 signaling; overexpression of ELK1 reverses the inhibitory effects of SPAG9 siRNA, placing SPAG9 upstream of ELK1 in HCC metastatic signaling. siRNA knockdown, ELK1 overexpression rescue, wound healing/invasion assays, Western blot OncoTargets and therapy Medium 27042099
2018 JLP (SPAG9) protects cancer cells from H2O2-induced cell death through the JLP-JNK signaling pathway; JLP knockdown increases cell death and intracellular ROS levels and attenuates H2O2-induced JNK activation. A JLP mutant lacking the JNK-binding domain failed to rescue cell viability or JNK activation. siRNA knockdown, rescue with wild-type vs JNK-binding domain mutant JLP, cell viability assays, ROS measurement Biochemical and biophysical research communications Medium 29753743
2019 JLP (SPAG9) tethers PLK1 to the centrosome upon extracellular stimulation in a p38MAPK- and dynein-dependent manner, leading to centrosome maturation and microtubule array formation; the JLP centrosome localization domain is required for binding the centrosome and forming the JLP focus and microtubule array. This JLP-centrosome function is essential for NF-κB p65 nuclear translocation via microtubule-based transport. Live imaging, domain deletion analysis, p38 inhibitor treatment, dynein inhibition, NF-κB p65 nuclear transport assays Science advances High 31803841
2019 SPAG9 promotes prostate cancer cell proliferation, migration, and cell cycle progression via the MAPK signaling pathway; knockout of SPAG9 suppresses tumor growth in nude mice. siRNA/overexpression, in vitro proliferation/migration assays, xenograft model, Western blot for MAPK pathway components American journal of translational research Low 31497238
2019 SPAG9 interacts with JNK by co-immunoprecipitation in HepG2 liver cancer cells; SPAG9 depletion decreases expression of JNK, p38, and MKK3 but not MKK6, defining a SPAG9/MKK3/p38 signaling axis in liver cancer cell proliferation. Co-immunoprecipitation, siRNA knockdown, Western blot for pathway components, cell proliferation and apoptosis assays Oncology reports Medium 30720101
2019 JLP (SPAG9) regulates lysosome localization (preventing peripheral redistribution) and autophagosome-lysosome fusion; rescue experiments show this requires the JLP-kinesin-1 heavy chain interaction domain but not the dynactin p150Glued or lysosomal transmembrane protein 55b binding regions. JLP knockdown, lysosome localization imaging, autophagy flux assays, rescue with wild-type and deletion mutants Drug discoveries & therapeutics Medium 32023558
2019 JLP (SPAG9) mediates protection against curcumin-induced cancer cell death by regulating lysosome positioning and p38 MAPK signaling to induce autophagy; JLP knockdown impairs perinuclear lysosomal accumulation and autophagosome-lysosome fusion. A JLP mutant lacking the p38 MAPK-binding domain failed to rescue cell viability or p38 activation. JLP knockdown, rescue with wild-type vs p38-binding domain mutant JLP, lysosome localization assays, autophagy assays, cell viability Biochemical and biophysical research communications Medium 31787236
2022 The SPAG9::JAK2 fusion protein (from t(9;17)(p24;q21)) constitutively activates JAK2 kinase domain phosphorylation, phosphorylating STAT3/STAT5/STAT1, and confers IL-3-independent proliferation in Ba/F3 cells. Mutation of JAK2 kinase domain tyrosines abolishes IL-3 independence. STAT1 activates BCL-2 and MCL-1, contributing to aberrant growth; the JAK2-STAT1-BCL-2/MCL-1 axis mediates SPAG9::JAK2-driven proliferation. Retroviral transduction of Ba/F3 cells, IL-3-independent growth assay, phospho-Western blot, kinase domain mutagenesis, gene expression analysis, JAK inhibitor (ruxolitinib) treatment Cancer gene therapy High 35879405
2023 JLP (SPAG9) tethers Foxk1 transcriptional repressor, preventing Foxk1-mediated repression of N-cadherin; TGF-β1 induces Foxk1 expression, but this is counteracted by JLP. Loss of JLP results in Foxk1 induction and N-cadherin downregulation, promoting partial EMT in renal tubular epithelial cells. TGF-β1 treatment, JLP knockdown/overexpression, Foxk1 modulation, N-cadherin expression assays, transcription factor binding analysis iScience Medium 37013185
2023 HLA-G functions in an HLA-G/SPAG9/STAT3 axis to promote the establishment of brain metastatic lesions; premetastatic brain metastasis initiating cells exhibit increased HLA-G and SPAG9 expression as part of a shared transcriptomic signature. RNA sequencing of premetastatic vs macrometastatic brain metastasis initiating cells, functional studies of the HLA-G/SPAG9/STAT3 axis Proceedings of the National Academy of Sciences of the United States of America Low 36780531
2026 TGF-β1 promotes nuclear translocation of β-catenin and LEF1, which together suppress JLP (SPAG9) transcription, leading to unchecked autophagy and renal fibrosis; inhibition of β-catenin/LEF1 with iCRT3 or LEF1 silencing restores JLP expression and attenuates autophagy and fibrosis in murine models. CKD patient kidney analysis, murine fibrosis models, HK-2 cell culture, β-catenin/LEF1 inhibitor treatment, LEF1 silencing, JLP expression/autophagy assays JCI insight Medium 41505216
2017 JLP (SPAG9) is required for CD4+ T cell TCR-initiated proliferation, IL-2 production, and CD154 induction; JLP deficiency impairs Ca2+ influx and NF-AT activation but not MAPK, NF-κB, or AP-1 signaling pathways upon TCR stimulation. jlp-deficient mice CD4+ T cells, TCR stimulation, proliferation assay, cytokine measurement, Ca2+ flux assay, NF-AT reporter assay, pathway signaling Western blots Molecular immunology Medium 28521278

Source papers

Stage 0 corpus · 63 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2002 JLP: A scaffolding protein that tethers JNK/p38MAPK signaling modules and transcription factors. Proceedings of the National Academy of Sciences of the United States of America 118 12391307
2006 Activation of p38alpha/beta MAPK in myogenesis via binding of the scaffold protein JLP to the cell surface protein Cdo. The Journal of cell biology 103 17074887
2005 Characterization of a novel human sperm-associated antigen 9 (SPAG9) having structural homology with c-Jun N-terminal kinase-interacting protein. The Biochemical journal 74 15693750
2019 LncRNA H19 promotes the committed differentiation of stem cells from apical papilla via miR-141/SPAG9 pathway. Cell death & disease 53 30755596
2005 JLP associates with kinesin light chain 1 through a novel leucine zipper-like domain. The Journal of biological chemistry 51 15987681
2016 MicroRNA-200a-3p suppresses tumor proliferation and induces apoptosis by targeting SPAG9 in renal cell carcinoma. Biochemical and biophysical research communications 44 26797273
2015 JSAP1/JIP3 and JLP regulate kinesin-1-dependent axonal transport to prevent neuronal degeneration. Cell death and differentiation 37 25571974
2013 Down regulation of SPAG9 reduces growth and invasive potential of triple-negative breast cancer cells: possible implications in targeted therapy. Journal of experimental & clinical cancer research : CR 35 24330581
2019 LncRNA NEAT1 enhances the resistance of anaplastic thyroid carcinoma cells to cisplatin by sponging miR‑9‑5p and regulating SPAG9 expression. International journal of oncology 34 31485599
2013 Clinical significance and biological roles of SPAG9 overexpression in non-small cell lung cancer. Lung cancer (Amsterdam, Netherlands) 32 23711689
2008 Ablation of the scaffold protein JLP causes reduced fertility in male mice. Transgenic research 31 18574703
2008 Kinesin adapter JLP links PIKfyve to microtubule-based endosome-to-trans-Golgi network traffic of furin. The Journal of biological chemistry 31 19056739
2013 SPAG9 is overexpressed in human astrocytoma and promotes cell proliferation and invasion. Tumour biology : the journal of the International Society for Oncodevelopmental Biology and Medicine 30 23696027
2009 Small interfering RNA-mediated down-regulation of SPAG9 inhibits cervical tumor growth. Cancer 30 19813278
2005 Sperm associated antigen 9 (SPAG9): a new member of c-Jun NH2 -terminal kinase (JNK) interacting protein exclusively expressed in testis. The Keio journal of medicine 23 16077255
2016 The expression of DAMP proteins HSP70 and cancer-testis antigen SPAG9 in peripheral blood of patients with HCC and lung cancer. Cell stress & chaperones 20 28028759
2015 Scaffold protein JLP is critical for CD40 signaling in B lymphocytes. The Journal of biological chemistry 20 25586186
2006 Endodermal differentiation of murine embryonic carcinoma cells by retinoic acid requires JLP, a JNK-scaffolding protein. Journal of cellular biochemistry 20 16619266
2005 Characterization of immune response in mice to plasmid DNA encoding human sperm associated antigen 9 (SPAG9). Vaccine 20 16061308
2016 SPAG9 is involved in hepatocarcinoma cell migration and invasion via modulation of ELK1 expression. OncoTargets and therapy 18 27042099
2014 SPAG9 is overexpressed in human prostate cancer and promotes cancer cell proliferation. Tumour biology : the journal of the International Society for Oncodevelopmental Biology and Medicine 18 24740566
2004 Isolation and characterization of a haploid germ cell specific sperm associated antigen 9 (SPAG9) from the baboon. Molecular reproduction and development 18 15293220
2005 Immunogenicity and contraceptive potential of recombinant human sperm associated antigen (SPAG9). Journal of reproductive immunology 17 16143403
2016 Sperm-associated antigen 9 (SPAG9) promotes the survival and tumor growth of triple-negative breast cancer cells. Tumour biology : the journal of the International Society for Oncodevelopmental Biology and Medicine 16 27449044
2016 Cancer testis antigen SPAG9 is a promising marker for the diagnosis and treatment of lung cancer. Oncology reports 15 26934841
2015 Identification of SPAG9 as a novel JAK2 fusion partner gene in pediatric acute lymphoblastic leukemia with t(9;17)(p24;q21). Genes, chromosomes & cancer 15 25951811
2019 SPAG9 promotes prostate cancer proliferation and metastasis via MAPK signaling pathway. American journal of translational research 14 31497238
2014 SPAG9 expression is increased in human prostate cancer and promotes cell motility, invasion and angiogenesis in vitro. Oncology reports 14 25310386
2018 JLP-JNK signaling protects cancer cells from reactive oxygen species-induced cell death. Biochemical and biophysical research communications 12 29753743
2018 MiR-524 inhibits cell proliferation and induces cell apoptosis in thyroid cancer via targeting SPAG9. European review for medical and pharmacological sciences 12 29949157
2016 Aberrant expression of JNK-associated leucine-zipper protein, JLP, promotes accelerated growth of ovarian cancer. Oncotarget 12 27655714
2023 An HLA-G/SPAG9/STAT3 axis promotes brain metastases. Proceedings of the National Academy of Sciences of the United States of America 11 36780531
2023 The bioceramic sealer iRoot SP promotes osteogenic differentiation of human stem cells from apical papilla via miR-141-3p/SPAG9/MAPK signalling pathway. International endodontic journal 11 37357722
2021 ZSCAN16-AS1 expedites hepatocellular carcinoma progression via modulating the miR-181c-5p/SPAG9 axis to activate the JNK pathway. Cell cycle (Georgetown, Tex.) 11 34097562
2018 Sperm associated antigen 9 (SPAG9) a promising therapeutic target of ovarian carcinoma. Tumour biology : the journal of the International Society for Oncodevelopmental Biology and Medicine 11 29745297
2018 SPAG9 regulates HEF1 expression and drives EMT in bladder transitional cell carcinoma via rac1 signaling pathway. American journal of cancer research 11 30662804
2014 JSAP1 and JLP are required for ARF6 localization to the midbody in cytokinesis. Genes to cells : devoted to molecular & cellular mechanisms 11 25130574
2019 SPAG9/MKK3/p38 axis is a novel therapeutic target for liver cancer. Oncology reports 10 30720101
2020 Functional role of c-Jun NH2-terminal kinase-associated leucine zipper protein (JLP) in lysosome localization and autophagy. Drug discoveries & therapeutics 9 32023558
2012 A novel role of the scaffolding protein JLP in tuning CD40-induced activation of dendritic cells. Immunobiology 9 23182713
2005 Molecular cloning and characterization of the macaque sperm associated antigen 9 (SPAG9): an orthologue of human SPAG9 gene. Molecular reproduction and development 9 15736136
2014 Sperm associated antigen 9 (SPAG9) expression and humoral response in benign and malignant salivary gland tumors. Oncoimmunology 8 25941602
2019 Protective role of c-Jun NH2-terminal kinase-associated leucine zipper protein (JLP) in curcumin-induced cancer cell death. Biochemical and biophysical research communications 7 31787236
2019 JLP-centrosome is essential for the microtubule-mediated nucleocytoplasmic transport induced by extracellular stimuli. Science advances 7 31803841
2023 JLP/Foxk1/N-cadherin axis fosters a partial epithelial-mesenchymal transition state in epithelial tubular cells. iScience 6 37013185
2022 The combination of ruxolitinib and Bcl-2/Mcl-1 inhibitors has a synergistic effect on leukemic cells carrying a SPAG9::JAK2 fusion. Cancer gene therapy 6 35879405
2016 SPAG9 is overexpressed in osteosarcoma, and regulates cell proliferation and invasion through regulation of JunD. Oncology letters 6 27698841
2015 SPAG9 controls the cell motility, invasion and angiogenesis of human osteosarcoma cells. Experimental and therapeutic medicine 6 26893659
2014 The scaffold protein JLP plays a key role in regulating ultraviolet B-induced apoptosis in mice. Genes to cells : devoted to molecular & cellular mechanisms 5 24520900
2017 Scaffold protein JLP mediates TCR-initiated CD4+T cell activation and CD154 expression. Molecular immunology 4 28521278
2015 Critical role of JSAP1 and JLP in axonal transport in the cerebellar Purkinje cells of mice. FEBS letters 4 26320416
2025 Truncated SPAG9 as a novel candidate gene for a new syndrome: Coarse facial features, albinism, cataract and developmental delay (CACD syndrome). Genetics and molecular biology 3 39846792
2021 c-Jun NH2-terminal kinase (JNK)/stress-activated protein kinase-associated protein 1 (JSAP1) attenuates curcumin-induced cell death differently from its family member, JNK-associated leucine zipper protein (JLP). Drug discoveries & therapeutics 3 33716240
2020 miR-874 inhibits gastric cancer cell proliferation by targeting SPAG9. BMC cancer 3 32503577
2016 SPAG9 promotes endometrial carcinoma cell invasion through regulation of genes related to the epithelial-mesenchymal transition. European journal of gynaecological oncology 3 27352556
2022 Oncolytic adenovirus carrying SPAG9-shRNA enhanced the efficacy of docetaxel for advanced prostate cancer. Anti-cancer drugs 2 34561997
2022 Oncolytic Adenovirus with SPAG9 shRNA Driven by DD3 Promoter Improved the Efficacy of Docetaxil for Prostate Cancer. Journal of oncology 2 35535313
2012 Assessment of SPAG9 Transcript in Fine Needle Aspirates of Thyroid Nodules. European thyroid journal 2 24783006
2022 Significance of cancer testis-associated antigens (SPAG9 and FBXO39) in colon cancer. Indian journal of cancer 1 34380828
2009 [Expression and location of SPAG9 in human ejaculated spermatozoa]. Zhonghua nan ke xue = National journal of andrology 1 19947555
2026 A TGF-β1/LEF1/β-catenin/JLP network motif regulates autophagy and tubule injury in renal fibrosis. JCI insight 0 41505216
2022 [Corrigendum] SPAG9 expression is increased in human prostate cancer and promotes cell motility, invasion and angiogenesis in vitro. Oncology reports 0 35383860
2021 MiR-524 inhibits cell proliferation and induces cell apoptosis in thyroid cancer via targeting SPAG9. European review for medical and pharmacological sciences 0 34919215

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