Affinage

STK33

Serine/threonine-protein kinase 33 · UniProt Q9BYT3

Length
514 aa
Mass
57.8 kDa
Annotated
2026-06-10
34 papers in source corpus 23 papers cited in narrative 22 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 8/8 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

STK33 is a CAMK-group serine/threonine protein kinase whose best-established physiological role is in male germ cell differentiation, where it is required for spermatid elongation and assembly of the sperm tail (PMID:29155043, PMID:37146716). STK33 protein localizes to the spermatid cytoplasm and the caudal end of the manchette during spermatid elongation, and its loss produces malformed, immotile spermatozoa with disordered tail structures including defective mitochondrial sheath, fibrous sheath, outer dense fiber, and axoneme (PMID:29155043, PMID:37146716). Mechanistically it phosphorylates the fibrous sheath A-kinase anchoring proteins AKAP3 and AKAP4 in vitro, and loss-of-function mutations in STK33 cause non-obstructive azoospermia and multiple morphological abnormalities of the flagella (MMAF) in humans (PMID:34155512, PMID:37146716). Beyond the testis, STK33 directly binds and phosphorylates the non-alpha-helical amino-terminal domain of the intermediate filament vimentin (PMID:18811945). STK33 also phosphorylates the tyrosine-metabolic enzyme HPD at T382, a mark that recruits the E3 ligase PELI1 to drive HPD polyubiquitylation and degradation; this STK33 activity is restrained by TTC36, defining a TTC36–STK33–PELI1 axis controlling tyrosine catabolism (PMID:31537781). STK33 protein is held stable by the HSP90/CDC37 chaperone complex and is degraded via the proteasome upon HSP90 inhibition (PMID:22451720). In tumor contexts STK33 binds c-Myc to enhance its transcriptional activity and modulates HIF-1α/VEGF and mTORC1/S6K1 signaling (PMID:25398772, PMID:29100402, PMID:42157211). The early proposal that STK33 kinase activity is selectively required for survival of KRAS-mutant cancer cells was refuted by multiple orthogonal approaches across independent laboratories (PMID:21742770, PMID:22323609, PMID:23256033).

Mechanistic history

Synthesis pass · year-by-year structured walk · 14 steps
  1. 2001 Medium

    Established STK33 as a distinct gene product by identifying it as a novel serine/threonine kinase and placing it phylogenetically in the CAMK group, framing expectations for its catalytic class.

    Evidence Comparative genome analysis, cDNA sequencing and phylogenetic classification

    PMID:11738831

    Open questions at the time
    • No direct enzymatic activity demonstrated
    • Lacks the canonical calcium/calmodulin binding domain, leaving regulation unknown
    • No substrate identified at this stage
  2. 2005 Medium

    Defined a non-ubiquitous expression pattern enriched in testis and other epithelia, pointing toward specialized roles in spermatogenesis and organ development rather than a housekeeping function.

    Evidence Immunofluorescence, Western blot and RNA analysis across tissues

    PMID:16176263

    Open questions at the time
    • Expression pattern does not establish molecular function
    • No causal link to a cellular process yet
  3. 2008 High

    Provided the first direct substrate by showing STK33 binds and phosphorylates the amino-terminal domain of vimentin, the first concrete biochemical activity assigned to the kinase.

    Evidence In vitro kinase assay with recombinant vimentin, reciprocal Co-IP, co-sedimentation and colocalization

    PMID:18811945

    Open questions at the time
    • Functional consequence of vimentin phosphorylation not defined
    • Phosphosite on vimentin not mapped
  4. 2009 Medium

    Proposed a synthetic lethal role in which STK33 kinase activity sustains KRAS-mutant cancer cell viability via S6K1/BAD, motivating it as a drug target.

    Evidence High-throughput RNAi screen with viability readout and pathway analysis

    PMID:19490892

    Open questions at the time
    • No in vitro reconstitution of the proposed kinase activity
    • Subsequently contradicted by independent labs
  5. 2011 High

    Refuted the KRAS synthetic-lethality model by showing STK33 kinase activity is dispensable for KRAS-dependent survival, correcting the field's target rationale.

    Evidence RNAi, dominant-mutant overexpression, small-molecule inhibitors and synthetic-lethal siRNA screening across a broad cell panel, replicated by independent labs

    PMID:21742770 PMID:22323609 PMID:23256033

    Open questions at the time
    • Does not address whether STK33 has KRAS-independent oncogenic roles
    • Leaves the physiological substrate question open
  6. 2012 High

    Identified the mechanism controlling STK33 protein stability, showing HSP90/CDC37 chaperones STK33 and that HSP90 inhibition drives its proteasomal degradation.

    Evidence MS-based interaction screen, Co-IP, pharmacologic HSP90 inhibition with in vitro and in vivo rescue

    PMID:22451720

    Open questions at the time
    • Does not resolve the contested KRAS dependency mechanism
    • Phosphorylation targets downstream of stabilized STK33 not defined here
  7. 2017 High

    Established STK33 as essential for spermatid differentiation, linking its manchette localization to a defined structural sperm phenotype upon knockout.

    Evidence Constitutive and germ-cell conditional knockout mice with immunofluorescence localization and histology

    PMID:29155043

    Open questions at the time
    • Substrates driving the manchette/tail phenotype not yet identified
    • Whether kinase activity is required in vivo not tested here
  8. 2017 Medium

    Connected STK33 to tumor angiogenesis and a hypoxia transcriptional circuit, showing it regulates HIF-1α/VEGF-A and is itself a direct HIF-1α target gene.

    Evidence Co-IP and rescue for HIF-1α/VEGF; ChIP and reporter assays for HIF-1α binding to the STK33 promoter in PDAC

    PMID:29038348 PMID:29100402

    Open questions at the time
    • Direct kinase substrate within the HIF axis not defined
    • Single-lab findings for each arm
  9. 2019 High

    Defined a physiological substrate and signaling axis outside the testis: STK33 phosphorylates HPD at T382 to trigger PELI1-mediated degradation, with TTC36 as a negative regulator controlling tyrosine metabolism.

    Evidence Co-IP, in vitro kinase assay mapping T382, ubiquitylation assay, and Ttc36 knockout mouse with biochemical/behavioral phenotyping

    PMID:31537781

    Open questions at the time
    • Tissue scope of the TTC36-STK33-PELI1 axis beyond the KO model not defined
    • How STK33 activity is regulated in this context unknown
  10. 2019 Low

    Extended STK33's oncogenic interactome with reports that it binds and activates ERK2 and drives PI3K/AKT/mTOR signaling in tumor cells.

    Evidence In vitro kinase assay and Co-IP for ERK2 in colorectal cells; knockdown/overexpression with pathway readout in PanNET cells

    PMID:30760631 PMID:31261148

    Open questions at the time
    • ERK2 finding is from a single lab with a related withdrawn paper, not independently confirmed
    • Direct kinase activity on pathway components in the PI3K/AKT/mTOR arm not measured
  11. 2021 Medium

    Validated the spermatogenic role in humans and added a transcriptional regulator, showing STK33 loss-of-function causes MMAF and that NFYB drives STK33 expression in lymphoma.

    Evidence Whole-exome sequencing with sperm ultrastructure analysis; ChIP and pathway rescue for NFYB/Hedgehog in DLBCL

    PMID:34155512 PMID:34536775

    Open questions at the time
    • MMAF mutation effect on kinase activity not dissected
    • NFYB-Hedgehog link is single-lab
  12. 2023 High

    Pinpointed AKAP3/AKAP4 as fibrous sheath substrates explaining the sperm tail defect and confirmed STK33 loss causes human non-obstructive azoospermia, unifying the spermiogenesis mechanism.

    Evidence Differential phosphoproteomics, in vitro kinase assays, mouse KO/KI models, human mutation identification and sperm structural analysis

    PMID:37146716

    Open questions at the time
    • Direct in vivo demonstration that AKAP3/4 phosphorylation alone drives the phenotype not isolated
    • Full set of germ-cell substrates not enumerated
  13. 2023 Medium

    Added chemical-biology tractability and a renal autophagy role, showing STK33 can be degraded by KLHDC2-recruiting PROTACs and that its knockdown activates mTOR/ULK1-dependent autophagy.

    Evidence AdPROM E3 ligase screen with endogenously tagged STK33 and PROTAC proof-of-concept; siRNA knockdown with autophagy markers in RCC

    PMID:37101009 PMID:37591251

    Open questions at the time
    • Autophagy finding is low-confidence single-lab without direct kinase measurement
    • Whether degrader-based STK33 loss phenocopies genetic loss in vivo untested
  14. 2025 Medium

    Revealed a potential non-catalytic scaffolding role in germ cells, where STK33 is recruited to TSKS foci but does not phosphorylate TSKS or YBX2 in vitro.

    Evidence IP/MS, reciprocal Co-IP, proximity ligation assay and in vitro kinase assay (negative for TSKS/YBX2)

    PMID:39909973

    Open questions at the time
    • Functional role of the STK33-TSKS interaction not established
    • Whether TSKS recruitment is kinase-independent in vivo unknown

Open questions

Synthesis pass · forward-looking unresolved questions
  • How STK33 kinase activity is regulated in the absence of a canonical calcium/calmodulin binding domain, and which subset of substrates and scaffolding interactions account for its distinct roles across spermiogenesis, tyrosine metabolism, and tumors, remains unresolved.
  • No structural model of activation defined in the corpus
  • Catalytic versus scaffolding contributions not separated across contexts
  • Mechanism integrating mTORC1/S6K1, HIF-1α, and c-Myc effects not unified

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0016740 transferase activity 3 GO:0140096 catalytic activity, acting on a protein 3
Localization
GO:0005856 cytoskeleton 2 GO:0005829 cytosol 1
Pathway
R-HSA-1474165 Reproduction 2 R-HSA-392499 Metabolism of proteins 2 R-HSA-1430728 Metabolism 1
Partners
AKAP3AKAP4CDC37HPDHSP90MYCTSKSVIM

Evidence

Reading pass · 22 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2009 STK33 promotes cancer cell viability in a kinase activity-dependent manner by regulating suppression of mitochondrial apoptosis through S6K1-induced inactivation of the death agonist BAD, selectively in mutant KRAS-dependent cells. High-throughput RNAi screen, knockdown with viability readout, pathway analysis Cell Medium 19490892
2011 STK33 kinase activity is NOT required for survival of KRAS-dependent cancer cells: STK33 knockdown, dominant mutant overexpression, and small-molecule kinase inhibitors all failed to affect KRAS signaling or cell survival, refuting the proposed synthetic lethal interaction. RNAi, dominant mutant overexpression, small-molecule inhibitors, synthetic lethal siRNA screen across broad panel Cancer research High 21742770 22323609 23256033
2001 STK33 was identified as a novel serine/threonine protein kinase encoded on human chromosome 11p15.3, with phylogenetic analysis placing it in the calcium/calmodulin-dependent protein kinase (CAMK) group, though lacking the canonical calcium/calmodulin binding domain. Comparative genome analysis, cDNA sequencing, phylogenetic analysis, RT-PCR Gene Medium 11738831
2005 STK33/Stk33 protein is highly expressed in testis (particularly in spermatogenic epithelial cells), lung epithelia, alveolar macrophages, retinal horizontal cells, and embryonic organs, establishing a non-ubiquitous expression pattern consistent with roles in spermatogenesis and organ ontogenesis. Immunofluorescence, Western blot, RNA analysis across tissues The FEBS journal Medium 16176263
2008 Stk33 directly binds to vimentin and phosphorylates the non-alpha-helical amino-terminal domain of vimentin in vitro; co-immunoprecipitation from cultured cell extracts confirmed in vivo association, and co-sedimentation assay showed direct binding without additional mediating proteins. In vitro kinase assay with recombinant vimentin, co-immunoprecipitation, co-sedimentation assay, immunofluorescence colocalization BMC biochemistry High 18811945
2012 The HSP90/CDC37 chaperone complex binds to and stabilizes STK33 protein in human cancer cells; pharmacologic HSP90 inhibition induces proteasome-mediated degradation of STK33, triggering apoptosis preferentially in KRAS mutant cells in an STK33-dependent manner. Mass spectrometry-based protein interaction screen, co-immunoprecipitation, pharmacologic HSP90 inhibition, in vitro and in vivo tumor models The Journal of experimental medicine High 22451720
2013 Stk33 co-localizes with vimentin in hypothalamic tanycytes in rodent and higher mammalian brains, and Stk33 expression in tanycytes is regulated by photoperiod, mirroring vimentin regulation, suggesting involvement in photoperiodic endocrine regulation. Immunofluorescence, double-immunostaining, co-immunoprecipitation, Western blot across photoperiod conditions Cell and tissue research Medium 24057876
2014 STK33 directly binds to c-Myc and increases its transcriptional activity, promoting hepatocellular carcinoma cell proliferation; the C-terminus of STK33 blocks this STK33/c-Myc association and downregulates HCC cell proliferation. Co-immunoprecipitation, domain-mapping with C-terminus constructs, in vitro and in vivo HCC proliferation assays, TAM-inducible transgenic and knockout mouse models Gut Medium 25398772
2017 Stk33 is essential for spermatid differentiation: constitutive Stk33 deletion in mice results in severely malformed and immotile spermatozoa with disordered structural tail elements; Stk33 protein localizes to the cytoplasm and partially co-localizes with the caudal end of the manchette in elongating spermatids, and its loss leads to an abnormal tight, straight, elongated manchette. Constitutive and conditional knockout mice, immunofluorescence localization, histological analysis of spermatogenesis Developmental biology High 29155043
2017 HSP90-stabilized STK33 interacts with and regulates hypoxia-driven accumulation and activation of HIF-1α, as well as secretion of VEGF-A in hypoxic cancer cells, promoting tumor angiogenesis; ectopic STK33 restored blood vessel formation in vivo after HSP90 inhibition. Co-immunoprecipitation, genetic STK33 abrogation/overexpression, tumor xenograft vascularization assay, VEGF-A secretion assay Oncotarget Medium 29100402
2017 HIF-1α directly transcriptionally upregulates STK33 by binding to a hypoxia response element in its promoter, establishing STK33 as a downstream mediator of HIF1α in pancreatic ductal adenocarcinoma. Chromatin immunoprecipitation (ChIP), reporter assay, knockdown/overexpression in PDAC cells and xenografts Cancer research Medium 29038348
2019 TTC36 binds HPD and reduces STK33 binding to HPD, thereby inhibiting STK33-mediated phosphorylation of HPD at T382; this phosphorylation recruits PELI1, which polyubiquitylates HPD leading to its degradation. Deficiency of TTC36 enhances STK33-mediated HPD T382 phosphorylation and PELI1-mediated HPD downregulation, causing tyrosinemia and neurological damage in Ttc36-/- mice. Co-immunoprecipitation, in vitro kinase assay (identifying T382 phosphorylation site), ubiquitylation assay, Ttc36 knockout mouse model with biochemical and behavioral phenotyping Nature communications High 31537781
2019 STK33 phosphorylates ERK2 in vitro, binds ERK2 in cells, and acts as an upstream kinase to increase ERK2 activity, promoting tumorigenesis of colorectal cancer cells. In vitro kinase assay, co-immunoprecipitation, knockdown/overexpression in HCT15 cells, in vivo xenograft Bioscience reports Low 30760631
2019 STK33 promotes growth and progression of pancreatic neuroendocrine tumors via activation of the PI3K/AKT/mTOR pathway, as demonstrated by knockdown and overexpression experiments. siRNA knockdown, overexpression, Western blot for PI3K/AKT/mTOR pathway components, in vitro proliferation/invasion assays, in vivo xenograft Neuroendocrinology Medium 31261148
2021 A homozygous frameshift mutation (c.1235del, p.T412Kfs*14) in STK33 causes MMAF (multiple morphological abnormalities of the flagella) phenotype in humans, establishing STK33 as an MMAF-related gene required for normal sperm flagellar structure and motility. Whole-exome sequencing, mRNA analysis, sperm morphology and ultrastructure analysis (electron microscopy) Human molecular genetics Medium 34155512
2021 NFYB transcription factor binds to the STK33 promoter and promotes STK33 expression, which in turn activates the Hedgehog signaling pathway to promote cisplatin resistance in diffuse large B-cell lymphoma. Chromatin immunoprecipitation, promoter binding assay, knockdown of STK33 and NFYB, Hedgehog pathway inhibitor experiments Leukemia research Medium 34536775
2023 STK33 phosphorylates fibrous sheath components AKAP3 and AKAP4 (A-kinase anchoring proteins 3 and 4) in vitro; STK33 deletion in mice results in defects in mitochondrial sheath, fibrous sheath, outer dense fiber, and axoneme assembly, with AKAP3/4 expression decreased in testis. Loss-of-function mutations in STK33 in humans cause non-obstructive azoospermia. Differential phosphoproteomic analysis, in vitro kinase assay, Stk33 knockout and knockin mouse models, human mutation identification, sperm structural analysis Molecular & cellular proteomics High 37146716
2023 STK33 affects autophagy in renal cell carcinoma cells by activating the mTOR/ULK1 pathway: STK33 knockdown leads to decreased p-mTOR and P62 and increased Beclin1, LC3, and p-ULK1, promoting autophagy and inhibiting cell proliferation and migration. siRNA knockdown, Western blot for mTOR/ULK1 pathway components, LC3 fluorescence assay, proliferation and migration assays Molecular biology reports Low 37101009
2023 KLHDC2 (CUL2 diGly receptor) was identified as an efficient E3 ubiquitin ligase capable of degrading STK33 via proximity-induced degradation (AdPROM system), demonstrating that STK33 can be targeted for proteasomal degradation by KLHDC2-recruiting molecules. AdPROM E3 ligase screen with GFP-tagged endogenous STK33, Western blot for degradation, peptide-based PROTAC proof-of-concept Cell chemical biology Medium 37591251
2025 STK33 is identified as a binding partner of TSKS (Testis Specific Serine Kinase Substrate) in testicular germ cells by co-immunoprecipitation; STK33 is recruited to TSKS foci through this interaction. STK33 was unable to phosphorylate TSKS or YBX2 in vitro, suggesting a non-catalytic scaffolding role in this context. Immunoprecipitation/mass spectrometry, co-immunoprecipitation, proximity ligation assay, in vitro phosphorylation assay (negative result for TSKS and YBX2) Reproductive sciences Medium 39909973
2026 STK33 promotes glycolysis and PanNET growth via the mTORC1/S6K1/HIF-1α signaling axis: STK33 knockdown reduces S6K1 phosphorylation and S6K1 inhibition reverses STK33-driven glycolysis; HIF-1α transcriptionally upregulates STK33 while STK33 promotes HIF-1α protein levels via mTORC1/S6K1, enhancing LDHA expression. shRNA knockdown, overexpression, S6K1 pharmacologic inhibition, Western blot for pathway components, glucose/lactate/ATP assays, in vivo xenograft, immunochemistry in patient specimens Journal of translational medicine Medium 42157211
2025 STK33 is implicated as a novel kinase in acrosomal exocytosis during sperm maturation, identified through phosphoproteomic analysis of epididymal sperm maturation; knockout mouse model validation confirmed functional relevance for sperm motility and fertilization capacity. High-resolution mass spectrometry phosphoproteomics, knockout mouse model, sperm motility and fertilization assays bioRxivpreprint Low

Source papers

Stage 0 corpus · 34 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2009 Synthetic lethal interaction between oncogenic KRAS dependency and STK33 suppression in human cancer cells. Cell 461 19490892
2011 STK33 kinase activity is nonessential in KRAS-dependent cancer cells. Cancer research 98 21742770
2012 STK33 kinase inhibitor BRD-8899 has no effect on KRAS-dependent cancer cell viability. Proceedings of the National Academy of Sciences of the United States of America 64 22323609
2012 Targeting of KRAS mutant tumors by HSP90 inhibitors involves degradation of STK33. The Journal of experimental medicine 58 22451720
2012 A Potent and Selective Quinoxalinone-Based STK33 Inhibitor Does Not Show Synthetic Lethality in KRAS-Dependent Cells. ACS medicinal chemistry letters 58 23256033
2001 A novel serine/threonine kinase gene, STK33, on human chromosome 11p15.3. Gene 39 11738831
2014 STK33 promotes hepatocellular carcinoma through binding to c-Myc. Gut 33 25398772
2005 Differential expression pattern of the novel serine/threonine kinase, STK33, in mice and men. The FEBS journal 33 16176263
2017 STK33 Promotes Growth and Progression of Pancreatic Cancer as a Critical Downstream Mediator of HIF1α. Cancer research 32 29038348
2019 HPD degradation regulated by the TTC36-STK33-PELI1 signaling axis induces tyrosinemia and neurological damage. Nature communications 29 31537781
2021 Novel frameshift mutation in STK33 is associated with asthenozoospermia and multiple morphological abnormalities of the flagella. Human molecular genetics 28 34155512
2008 The Serine/threonine kinase Stk33 exhibits autophosphorylation and phosphorylates the intermediate filament protein Vimentin. BMC biochemistry 27 18811945
2017 Stk33 is required for spermatid differentiation and male fertility in mice. Developmental biology 25 29155043
2023 Identification of KLHDC2 as an efficient proximity-induced degrader of K-RAS, STK33, β-catenin, and FoxP3. Cell chemical biology 23 37591251
2015 STK33 overexpression in hypopharyngeal squamous cell carcinoma: possible role in tumorigenesis. BMC cancer 20 25603720
2018 STK33 alleviates gentamicin-induced ototoxicity in cochlear hair cells and House Ear Institute-Organ of Corti 1 cells. Journal of cellular and molecular medicine 19 30256516
2015 STK33 plays an important positive role in the development of human large cell lung cancers with variable metastatic potential. Acta biochimica et biophysica Sinica 19 25662617
2017 STK33 participates to HSP90-supported angiogenic program in hypoxic tumors by regulating HIF-1α/VEGF signaling pathway. Oncotarget 17 29100402
2024 Click-chemistry mediated synthesis of OTBN-1,2,3-Triazole derivatives exhibiting STK33 inhibition with diverse anti-cancer activities. Bioorganic chemistry 15 38824700
2021 NFYB potentiates STK33 activation to promote cisplatin resistance in diffuse large B-cell lymphoma. Leukemia research 15 34536775
2023 STK33 Phosphorylates Fibrous Sheath Protein AKAP3/4 to Regulate Sperm Flagella Assembly in Spermiogenesis. Molecular & cellular proteomics : MCP 13 37146716
2020 miR-107 inhibited malignant biological behavior of non-small cell lung cancer cells by regulating the STK33/ERK signaling pathway in vivo and vitro. Journal of thoracic disease 13 32395291
2019 STK33/ERK2 signal pathway contribute the tumorigenesis of colorectal cancer HCT15 cells. Bioscience reports 13 30760631
2019 STK33 Promotes the Growth and Progression of Human Pancreatic Neuroendocrine Tumour via Activation of the PI3K/AKT/mTOR Pathway. Neuroendocrinology 12 31261148
2013 Co-localization of serine/threonine kinase 33 (Stk33) and vimentin in the hypothalamus. Cell and tissue research 11 24057876
2016 STK33 potentiates the malignancy of hypopharyngeal squamous carcinoma: Possible relation to calcium. Cancer biology & therapy 8 27414193
2023 Interaction between STK33 and autophagy promoted renal cell carcinoma metastasis by regulating mTOR/ULK1 signaling pathway. Molecular biology reports 6 37101009
2016 Serine/threonine-kinase 33 (Stk33) - Component of the neuroendocrine network? Brain research 6 27840186
2013 The STK33-linked SNP rs4929949 is associated with obesity and BMI in two independent cohorts of Swedish and Greek children. PloS one 6 23967198
2024 STK33 as the functional substrate of miR-454-3p for suppression and apoptosis in neuroblastoma. Molecules and cells 1 39515612
2026 The STK33/mTORC1/S6K1/HIF-1α signalling axis drives glycolysis and progression in pancreatic neuroendocrine tumours. Journal of translational medicine 0 42157211
2026 Piceatannol attenuates thoracic aortic aneurysm progression by targeting STK33. Biology direct 0 42265708
2025 Identification of YBX2 and TSKS As STK33 Interacting Proteins in Testicular Germ Cells. Reproductive sciences (Thousand Oaks, Calif.) 0 39909973
2019 WITHDRAWN: STK33/ERK2 signal pathway contribute the tumorigenesis of colorectal cancer HCT15 cells. Anti-cancer agents in medicinal chemistry 0 31272359

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