Affinage

SPATA2

Spermatogenesis-associated protein 2 · UniProt Q9UM82

Length
520 aa
Mass
58.4 kDa
Annotated
2026-06-10
21 papers in source corpus 12 papers cited in narrative 12 extracted findings
Cross-family judge vs UniProt: tie faithfulness: 5/6 claims corpus-supported (83%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

SPATA2 is a scaffolding adaptor protein that controls deubiquitination within immune receptor signaling complexes, recruiting the deubiquitinase CYLD to regulate inflammatory signaling and cell death (PMID:27307491, PMID:27545878). It is a constitutive component of the TNF receptor signaling complex (TNF-RSC) and NOD2 signaling complex, where its N-terminal PUB domain binds the USP domain of CYLD while a C-terminal PUB-interaction motif (PIM) binds the PUB domain of HOIP, the catalytic subunit of LUBAC; through this bridging, SPATA2 is required for HOIP-dependent recruitment of CYLD to these complexes (PMID:27307491, PMID:27545878, PMID:27458237). Beyond passive scaffolding, SPATA2 acts as an allosteric activator of the K63- and M1-linked deubiquitinase activity of CYLD, and the CYLD–SPATA2 complex preferentially cleaves M1 (linear) ubiquitin chains on RIPK1, thereby restraining NF-κB and MAPK signaling while licensing complex II formation, caspase activation, apoptosis, and necroptosis (PMID:27458237, PMID:28701375). SPATA2 additionally exerts CYLD-independent control of LUBAC by competing with OTULIN for HOIP binding via its PIM and zinc finger, promoting LUBAC autoubiquitylation and limiting linear ubiquitination, with combined loss of SPATA2 and CYLD causing perinatal lethality in mice (PMID:36640323). At the centrosome, SPATA2 directs CYLD-mediated deubiquitination of PLK4 to promote NEK7 phosphorylation and suppress NLRP3 inflammasome activation (PMID:31762063). SPATA2 is also required for male fertility, with knockout mice showing reduced testis size, sperm count, and germ cell proliferation alongside elevated inhibin alpha (PMID:29025062).

Mechanistic history

Synthesis pass · year-by-year structured walk · 7 steps
  1. 2016 High

    Established SPATA2 as a previously unrecognized component of the TNF-RSC that physically bridges CYLD and the LUBAC subunit HOIP, defining how CYLD is recruited to the receptor complex.

    Evidence Quantitative mass spectrometry of TNF-RSC, reciprocal Co-IP with domain mapping, and siRNA knockdown with NF-κB/necroptosis readouts; independently replicated across three concurrent papers

    PMID:27307491 PMID:27458237 PMID:27545878

    Open questions at the time
    • Did not resolve whether SPATA2 affects CYLD catalysis beyond recruitment
    • Structural basis of the PUB/PIM interactions not determined in these studies
  2. 2016 High

    Showed SPATA2 is not merely a scaffold but an allosteric activator of CYLD's K63- and M1-deubiquitinase activity, linking complex assembly to enzymatic output and to apoptosis/NF-κB/MAPK control.

    Evidence In vitro deubiquitinase activity assays plus Co-IP domain mapping and loss-of-function with apoptosis and signaling readouts

    PMID:27458237

    Open questions at the time
    • Mechanism of allosteric activation at the structural level not defined
    • Full substrate range within the TNF-RSC not enumerated
  3. 2017 High

    Defined RIPK1 as a key substrate, showing the CYLD–SPATA2 complex preferentially cleaves M1 chains on RIPK1 to sustain its kinase activity and dictate RIPK1-dependent death versus RIPK1-independent cytokine output.

    Evidence Genetic SPATA2 KO cell lines, in vitro deubiquitinase assays with M1 vs K63 chains, and RIPK1 ubiquitination biochemistry with cell death readouts

    PMID:28701375

    Open questions at the time
    • Did not separate scaffolding from activating contributions to chain specificity in vivo
    • RIPK1-independent JNK/cytokine branch mechanism not fully resolved
  4. 2019 High

    Extended SPATA2 function beyond death-receptor signaling by showing it recruits CYLD to the centrosome to deubiquitinate PLK4 and suppress NLRP3 inflammasome activation.

    Evidence SPATA2 KO macrophages, Co-IP of PLK4/CYLD/NEK7/NLRP3, in vitro deubiquitination assay, NEK7 S204A mutagenesis, and an in vivo peritonitis model

    PMID:31762063

    Open questions at the time
    • How SPATA2 targets CYLD specifically to the centrosome is unclear
    • Relationship between centrosomal and TNF-RSC pools of SPATA2 not addressed
  5. 2023 High

    Revealed a CYLD-independent role: SPATA2 competes with OTULIN for HOIP to restrain LUBAC activity, with genetic epistasis showing combined SPATA2/CYLD loss is lethal and signaling hyperactivation is OTULIN-dependent.

    Evidence Cyld-/-Spata2-/- double KO mice, OTULIN competition binding assays, M1-ubiquitin chain analysis, and pro-inflammatory gene expression across multiple cell types

    PMID:36640323

    Open questions at the time
    • Quantitative balance between CYLD-dependent and OTULIN-competitive functions unresolved
    • Domain requirements for OTULIN competition only partially mapped
  6. 2022 Medium

    Implicated SPATA2/CYLD in deubiquitination of broader substrates, stabilizing or destabilizing STAT1 and NCOA4 to modulate IFN-γ signaling, anti-PD-1 response, and ferritinophagy-driven ferroptosis.

    Evidence Genetic deletion/knockdown in CRC and cardiomyocyte models with STAT1 and NCOA4 ubiquitination assays, functional readouts, and in vivo models

    PMID:36195186 PMID:36531014

    Open questions at the time
    • Direct deubiquitination of these substrates by the SPATA2/CYLD complex not reconstituted in vitro
    • Context-specificity of substrate selection unexplained
  7. 2017 Medium

    Demonstrated a physiological requirement for SPATA2 in spermatogenesis and Sertoli cell function distinct from its inflammatory signaling roles.

    Evidence CRISPR/Cas9n global KO mice with testis histomorphology, sperm counts, BrdU proliferation, and inhibin alpha measurement

    PMID:29025062

    Open questions at the time
    • Molecular pathway linking SPATA2 to inhibin alpha regulation not defined
    • Whether the fertility role depends on CYLD/LUBAC is unknown

Open questions

Synthesis pass · forward-looking unresolved questions
  • How SPATA2's biochemical activities (CYLD recruitment/activation, OTULIN competition, substrate selection) are integrated and spatially partitioned across distinct cellular contexts remains unresolved.
  • No structural model of the SPATA2–CYLD–HOIP assembly
  • Unclear how distinct substrate engagements (RIPK1, PLK4, STAT1, p53, NCOA4, DVL1) are determined
  • Connection between reproductive and immune signaling functions unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 3 GO:0098772 molecular function regulator activity 2
Localization
GO:0005634 nucleus 1 GO:0005815 microtubule organizing center 1
Pathway
R-HSA-168256 Immune System 3 R-HSA-162582 Signal Transduction 2 R-HSA-5357801 Programmed Cell Death 2
Complex memberships
LUBACNOD2 signaling complexTNF-RSC

Evidence

Reading pass · 12 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2016 SPATA2 was identified as a novel component of the TNF receptor signaling complex (TNF-RSC). The N-terminal PUB domain of SPATA2 interacts with the USP domain of CYLD, while the C-terminus of SPATA2 interacts with HOIP (the catalytically active component of LUBAC). SPATA2 is required for recruitment of CYLD to the TNF-RSC. Loss of SPATA2 augments NF-κB transcriptional activation and inhibits TNF-α-induced necroptosis. Quantitative mass spectrometry of TNF-RSC composition, Co-IP/pulldown domain mapping, siRNA knockdown with NF-κB and necroptosis readouts The EMBO journal High 27307491
2016 SPATA2 is a constitutive direct binding partner of HOIP that bridges the interaction between CYLD and HOIP, enabling CYLD recruitment to TNFR1- and NOD2-signaling complexes. SPATA2 recruitment to these complexes is dependent on HOIP. Loss of SPATA2 abolishes CYLD recruitment and diminishes TNF-induced necroptosis, resembling loss of CYLD. Co-IP, pulldown, CRISPR/siRNA loss-of-function with necroptosis and NF-κB readouts in multiple signaling complexes Cell reports High 27545878
2016 SPATA2 interacts with CYLD via its PUB domain, while a PUB interaction motif (PIM) of SPATA2 interacts with the PUB domain of HOIP. Beyond scaffolding, SPATA2 acts as an allosteric activator of the K63- and M1-deubiquitinase activity of CYLD. SPATA2 is required for TNF-induced complex II formation, caspase activation, and apoptosis, and substantially attenuates TNF-induced NF-κB and MAPK signaling. Mass spectrometry screen, Co-IP domain mapping, in vitro deubiquitinase activity assay, loss-of-function with NF-κB/MAPK/apoptosis readouts EMBO reports High 27458237
2017 SPATA2-deficient cells show resistance to RIPK1-dependent apoptosis and necroptosis, and partial protection against RIPK1-independent apoptosis. SPATA2 deficiency promotes M1 ubiquitination of RIPK1, thereby inhibiting RIPK1 kinase activity. Biochemical in vitro evidence shows that the CYLD USP domain complexed with the SPATA2 PUB domain preferentially deubiquitinates M1 (linear) ubiquitin chains. SPATA2 deficiency also promotes MKK4 and JNK activation and cytokine production independently of RIPK1 kinase activity. SPATA2-deficient cell lines (genetic KO), in vitro deubiquitinase assay with M1 and K63 ubiquitin chains, RIPK1 ubiquitination biochemistry, cell death assays Genes & development High 28701375
2019 SPATA2 recruits CYLD to the centrosome to deubiquitinate PLK4 (polo-like kinase 4). Deubiquitination of PLK4 facilitates its phosphorylation of NEK7 at Ser204, which attenuates the NEK7–NLRP3 interaction required for NLRP3 inflammasome activation. SPATA2 deficiency enhances NLRP3 inflammasome activity in macrophages and in a peritonitis animal model. SPATA2 KO macrophages, Co-IP of PLK4/CYLD/NEK7/NLRP3, in vitro ubiquitination/deubiquitination assay, phosphorylation site mutagenesis (NEK7 S204A), shRNA knockdown, animal peritonitis model The EMBO journal High 31762063
2023 SPATA2 has CYLD-independent functions: SPATA2 competes with OTULIN for binding to HOIP via its PIM and zinc finger domain, thereby promoting LUBAC autoubiquitylation and attenuating LUBAC activity. Double knockout (Cyld-/-Spata2-/-) mice show highly penetrant perinatal lethality absent in single knockouts. Cyld-/-Spata2-/- cells show increased M1-linked TNFR1-SC ubiquitylation and elevated pro-inflammatory gene expression compared to either single KO, and this increased signaling depends on OTULIN. Double KO mice (CRISPR), OTULIN competition binding assay, M1-ubiquitin chain analysis, pro-inflammatory gene expression in fibroblasts/macrophages/intestinal epithelial cells, genetic epistasis with OTULIN Cell reports High 36640323
2022 SPATA2 and CYLD inhibit STAT1 accumulation and activation (and subsequent CXCL10 expression) in colorectal cancer cells. At steady state, STAT1 is highly ubiquitinated in a SPATA2/CYLD-dependent manner, suggesting SPATA2/CYLD deubiquitinate STAT1 to regulate IFN-γ signaling. Tumor-specific deletion of SPATA2 or CYLD enhances anti-PD-1 response in vivo. Specific deletion of SPATA2/CYLD in human and mouse CRC cell lines, STAT1 ubiquitination analysis, CXCL10 expression measurement, in vivo anti-PD-1 response assay Frontiers in oncology Medium 36531014
2022 The SPATA2/CYLD pathway contributes to doxorubicin-induced cardiomyocyte ferroptosis by enhancing ferritinophagy. SPATA2 knockdown reduced NCOA4 deubiquitination, attenuated ferritinophagy, and decreased ferroptosis. The pathway promotes NCOA4 deubiquitination (via CYLD), which stabilizes NCOA4 and drives ferritin degradation and intracellular iron overload. SPATA2 knockdown in cardiomyocytes, NCOA4 ubiquitination assay, Co-IP of SPATA2/CYLD interaction, ferroptosis markers (iron, ACSL4, LC3), in vivo mouse doxorubicin model Chemico-biological interactions Medium 36195186
2025 SPATA2/CYLD pathway-dependent deubiquitylation of p53 promotes ferroptosis in rat heart after ischemia/reperfusion by stabilizing p53, which suppresses SLC7A11 transcription, reducing GPX4 levels. SPATA2 knockdown restored p53 ubiquitination, increased SLC7A11 and GPX4, and reduced ferroptosis. SPATA2 knockdown in H9c2 cells and rat I/R model, p53 ubiquitination assay, SLC7A11/GPX4 expression, lurasidone as SPATA2 inhibitor Heart, lung & circulation Medium 40389334
2001 SPATA2 protein tagged with GFP is located in the nucleus when expressed in HLtat transfected cells. SPATA2 mRNA is expressed in testis and Sertoli cells, and FSH treatment of Sertoli cells induces changes in steady-state spata2 mRNA in a time-dependent manner. GFP-fusion protein expression and fluorescence localization, RT-PCR, FSH treatment of Sertoli cells Biochemical and biophysical research communications Low 11322771
2017 SPATA2 global knockout in mice (via CRISPR/Cas9n) results in approximately 40% decrease in testis size/weight, 40% decrease in sperm count, 28% decrease in germ cell proliferation, and approximately 70% increase in inhibin alpha-subunit mRNA and protein in testes, indicating SPATA2 ensures normal secretory function of Sertoli cells and is required for male fertility. CRISPR/Cas9n global KO mice, histomorphology, sperm count, BrdU proliferation assay, RT-PCR and Western blot for inhibin alpha Biology of reproduction Medium 29025062
2023 SPATA2 suppresses β-catenin signaling by attenuating DVL1 ubiquitination. Ectopic overexpression of SPATA2 inhibited EMT, impaired motility and invasiveness in vitro, reduced metastasis in vivo, and increased radiosensitivity in NSCLC cells. SPATA2 overexpression/KD in NSCLC cell lines, DVL1 ubiquitination assay, β-catenin signaling reporter, in vitro invasion/migration assays, in vivo metastasis model Thoracic cancer Low 36814090

Source papers

Stage 0 corpus · 21 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2016 SPATA2 links CYLD to the TNF-α receptor signaling complex and modulates the receptor signaling outcomes. The EMBO journal 121 27307491
2016 SPATA2-Mediated Binding of CYLD to HOIP Enables CYLD Recruitment to Signaling Complexes. Cell reports 116 27545878
2016 SPATA2 promotes CYLD activity and regulates TNF-induced NF-κB signaling and cell death. EMBO reports 100 27458237
2019 PLK4 deubiquitination by Spata2-CYLD suppresses NEK7-mediated NLRP3 inflammasome activation at the centrosome. The EMBO journal 81 31762063
2017 SPATA2 regulates the activation of RIPK1 by modulating linear ubiquitination. Genes & development 58 28701375
2022 The SPATA2/CYLD pathway contributes to doxorubicin-induced cardiomyocyte ferroptosis via enhancing ferritinophagy. Chemico-biological interactions 30 36195186
2017 SPATA2: more than a missing link. Cell death and differentiation 22 28282038
2022 The role of SPATA2 in TNF signaling, cancer, and spermatogenesis. Cell death & disease 21 36402749
2017 Deletion of Spata2 by CRISPR/Cas9n causes increased inhibin alpha expression and attenuated fertility in male mice. Biology of reproduction 17 29025062
2014 Investigating the genetic association of HCP5, SPATA2, TNIP1, TNFAIP3 and COG6 with psoriasis in Chinese population. International journal of immunogenetics 16 25264125
2001 Evidence for FSH-dependent upregulation of SPATA2 (spermatogenesis-associated protein 2). Biochemical and biophysical research communications 16 11322771
2021 Spata2 Knockdown Exacerbates Brain Inflammation via NF-κB/P38MAPK Signaling and NLRP3 Inflammasome Activation in Cerebral Ischemia/Reperfusion Rats. Neurochemical research 15 34075523
2009 The Story of SPATA2 (Spermatogenesis-Associated Protein 2): From Sertoli Cells to Pancreatic Beta-Cells. Current genomics 10 20119533
2007 Zebrafish spata2 is expressed at early developmental stages. The International journal of developmental biology 8 17486545
2023 SPATA2 suppresses epithelial-mesenchymal transition to inhibit metastasis and radiotherapy sensitivity in non-small cell lung cancer via impairing DVL1/β-catenin signaling. Thoracic cancer 7 36814090
2003 Cloning and characterization of the promoter region of human spata2 (spermatogenesis-associated protein 2) gene. Biochimica et biophysica acta 7 12531478
2016 SPATA2 - Keeping the TNF signal short and sweet. The EMBO journal 6 27470000
2023 SPATA2 restricts OTULIN-dependent LUBAC activity independently of CYLD. Cell reports 5 36640323
2022 SPATA2 and CYLD inhibit T cell infiltration into colorectal cancer via regulation of IFN-γ/STAT1 axis. Frontiers in oncology 2 36531014
2025 SPATA2/CYLD Pathway-Dependent Deubiquitylation of p53 Promotes Ferroptosis in Rat Heart After Ischaemia/Reperfusion Through Suppression of SLC7A11. Heart, lung & circulation 1 40389334
2025 Clinical and functional significance of SPATA2 in cancer particularly in LIHC. Scientific reports 0 40069269

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