Affinage

SPDYA

Speedy protein A · UniProt Q5MJ70

Length
313 aa
Mass
36.5 kDa
Annotated
2026-06-10
42 papers in source corpus 19 papers cited in narrative 19 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

SPDYA (Spy1/RingoA) is an atypical, cyclin-like activator of cyclin-dependent kinases that drives cell-cycle entry and proliferation through a structurally distinct mechanism of CDK2/CDK1 activation (PMID:12972555, PMID:28666995). Crystal structures of the CDK2-Spy1 binary and p27-CDK2-Spy1 ternary complexes show that Spy1 induces conformational changes that activate CDK2 independently of activation-loop phosphorylation, and that Spy1 lacks the cyclin-binding groove, rendering Spy1-CDK2 complexes resistant to p27 inhibition and altering substrate specificity (PMID:28666995). Spy1 directly binds the CDK inhibitor p27Kip1 through residues separable from its CDK2-interaction surface, and stimulates CDK2-mediated phosphorylation of p27 at T187, triggering its degradation to overcome p27-induced G1 arrest and promote S-phase entry (PMID:12972555, PMID:17671428, PMID:26771716). Through CDK2 activation Spy1 suppresses DNA-damage checkpoints and apoptosis in a p53/p21-dependent context, permitting damage-resistant DNA synthesis and mitotic entry while impairing repair fidelity (PMID:12839962, PMID:16951407, PMID:19106603). Spy1 promotes mammary transformation via CDK1 activation and FOXO1 inhibition, activates ERK1/2 in a MEK-independent manner to drive ligand-independent ERα signalling and tamoxifen resistance, and engages epigenetic silencing of CDK inhibitors via EZH2/H3K27me3 to override senescence during reprogramming (PMID:22280365, PMID:28423577, PMID:34486784). Spy1 protein abundance is controlled by proteasomal degradation through the E3 ligases Nedd4 (downstream of p53) and MDM2 (PMID:31829284, PMID:36443440). Beyond cell-cycle control, Spy1 uses its Ringo domain to bind SUN1 at the nuclear envelope and recruit CDK2 to support telomere-LINC complex attachment, homologous pairing and synapsis during meiosis prophase I (PMID:33015044, PMID:34039995), and regulates neural stem-cell exit from quiescence as a CDK activity threshold setter (PMID:36876138).

Mechanistic history

Synthesis pass · year-by-year structured walk · 19 steps
  1. 2003 High

    Established that Spy1 physically binds the CDK inhibitor p27Kip1 and uses this interaction to bypass p27-imposed G1 arrest, defining a mechanism by which Spy1 overrides a key proliferation brake.

    Evidence Yeast two-hybrid, in vitro pulldown, co-IP, CDK2 H1 kinase assay, and p27-null rescue in mammalian cells

    PMID:12972555

    Open questions at the time
    • Did not resolve whether Spy1 phosphorylates p27 directly or via CDK2
    • Structural basis of p27 binding unresolved
  2. 2003 Medium

    Showed Spy1 binds and activates CDK2 to promote survival after genotoxic damage, with dominant-negative CDK2 abolishing the effect, placing CDK2 activation as the required effector of Spy1-mediated cell survival.

    Evidence Clonogenic and comet assays, dominant-negative CDK2, siRNA knockdown across multiple damaging agents

    PMID:12839962

    Open questions at the time
    • Single lab
    • Did not define which checkpoint nodes are bypassed
  3. 2006 Medium

    Demonstrated that Spy1 suppresses UV-induced apoptosis and checkpoint signalling (Chk1, RPA, H2A.X) in a manner dependent on the Speedy/Ringo box, placing CDK2 activation upstream of checkpoint bypass.

    Evidence Inducible Spy1 expression, flow cytometry, checkpoint-marker westerns, Ringo-box point mutant

    PMID:16951407

    Open questions at the time
    • Mechanism of checkpoint suppression not biochemically dissected
    • Single lab
  4. 2007 High

    Reconstituted the biochemical core of Spy1 function by showing Spy1/CDK2 phosphorylates p27 at T187 in vitro and drives T187-dependent p27 degradation in cells, linking Spy1 directly to controlled p27 turnover.

    Evidence In vitro kinase assay with recombinant proteins, inducible expression, synchronized HeLa cells, T187A mutant control

    PMID:17671428

    Open questions at the time
    • Did not identify the ubiquitin ligase acting on phospho-T187 p27 in this context
  5. 2009 Medium

    Defined the genetic context of checkpoint bypass, showing Spy1's anti-apoptotic and checkpoint-overriding effects require functional p53 and p21 and come at the cost of impaired excision repair and increased mutation frequency.

    Evidence siRNA knockdown, p53/p21 requirement analysis, CPD repair and mutation-frequency assays, γH2A.X/Chk1 westerns

    PMID:19106603

    Open questions at the time
    • Mechanistic basis of repair inhibition unresolved
    • Single lab
  6. 2012 Medium

    Extended Spy1's transforming role to CDK1, showing mammary transformation depends on CDK1 activation and FOXO1 inhibition, broadening Spy1's CDK targets beyond CDK2.

    Evidence CDK1 activity assays, FOXO1 analysis, shRNA, in vivo mouse mammary tumor model

    PMID:22280365

    Open questions at the time
    • Direct CDK1 binding not structurally shown
    • FOXO1 regulation mechanism incomplete
  7. 2014 Low

    Linked Spy1 to growth-cone dynamics through CRMP1, implicating Spy1 in CDK5-mediated CRMP1 phosphorylation and Sema3A-induced collapse.

    Evidence Yeast two-hybrid, co-IP, overexpression/shRNA, growth-cone collapse assay, rat sciatic nerve crush

    PMID:25526860

    Open questions at the time
    • Single Co-IP plus limited mechanistic follow-up
    • How Spy1 modulates a CDK5-dependent event not defined
  8. 2015 Medium

    Identified a non-cell-cycle role in which Spy1 binds SCG10 and promotes its phosphorylation and proteasomal degradation, coupling Spy1 to axonal degeneration after nerve injury.

    Evidence Co-IP, in vitro binding, phosphorylation/degradation westerns, Spy1 inhibition with phenotypic rescue

    PMID:25869138

    Open questions at the time
    • Direct kinase identity for SCG10 phosphorylation not fully resolved
    • Single lab
  9. 2015 Medium

    Showed Spy1 sequesters CLIPR-59 away from the deubiquitinase CYLD, controlling RIP1 K63-ubiquitination and TNF-α-induced apoptosis in glioblastoma, revealing a scaffold-type role distinct from CDK activation.

    Evidence Co-IP, siRNA, ubiquitination and caspase activity assays

    PMID:26017671

    Open questions at the time
    • Structural basis of CLIPR-59 binding unknown
    • Single lab
  10. 2016 Medium

    Genetically separated Spy1's CDK2- and p27-binding surfaces, showing both contribute to proliferation and tumorigenesis but only p27 binding is required for p27 destabilisation.

    Evidence Site-directed mutagenesis, co-IP, CDK2 kinase and p27 stability assays, in vivo tumour model

    PMID:26771716

    Open questions at the time
    • Did not provide atomic-resolution view of the two interfaces
    • Single lab
  11. 2017 High

    Provided the structural mechanism: crystal structures of CDK2-Spy1 and p27-CDK2-Spy1 explain phosphorylation-independent activation, p27 resistance, and altered substrate specificity arising from absence of the cyclin-binding groove.

    Evidence X-ray crystallography of two complexes, in vitro kinase assay, mutagenesis, proliferation assay

    PMID:28666995

    Open questions at the time
    • Substrate-specificity rules not exhaustively mapped
    • No structure of Spy1-CDK1 complex
  12. 2017 Medium

    Connected Spy1 to MEK-independent ERK1/2 activation that drives ligand-independent ERα activation and tamoxifen resistance, adding a kinase-signalling output to Spy1 biology.

    Evidence ERK1/2 and ERα phosphorylation assays, MEK inhibitor controls, tamoxifen sensitivity, overexpression/knockdown

    PMID:28423577

    Open questions at the time
    • Direct mechanism of MEK-independent ERK activation undefined
    • Single lab
  13. 2019 Medium

    Defined upstream control of Spy1 abundance, showing p53 directs Nedd4-mediated proteasomal degradation of Spy1 and that p53 loss permits Spy1 accumulation and mammary tumorigenesis.

    Evidence Transgenic mouse, protein stability assays, Nedd4 co-IP/ubiquitination, p53 KO/mutation analysis

    PMID:31829284

    Open questions at the time
    • Nedd4 recognition motif on Spy1 not mapped
    • Single lab
  14. 2020 Medium

    Revealed a meiotic role: SPDYA binds SUN1 at the nuclear envelope through its Ringo domain and recruits CDK2 near the MAJIN-binding site, supporting telomere-nuclear envelope attachment.

    Evidence Co-IP, domain mapping, CDK2 inhibitor treatment, binding-site mutagenesis

    PMID:33015044

    Open questions at the time
    • In vivo meiotic requirement not yet tested at this stage
    • CDK2 substrates at the LINC complex unknown
  15. 2021 High

    Established structurally and genetically that the SUN1-SPDYA-CDK2 complex organises telomere supramolecular architecture and is required for homologous pairing and synapsis in meiosis prophase I.

    Evidence X-ray crystallography of ternary complex, SUN1 mutagenesis, SPDYA-binding-deficient SUN1 knockin mouse, meiotic phenotyping

    PMID:34039995

    Open questions at the time
    • Direct CDK2 phosphorylation targets driving telomere attachment not identified
  16. 2021 Medium

    Showed SPY1 activates EZH2/H3K27me3 to silence CDKN1A/CDKN2A and override senescence, making SPY1 a required factor for efficient somatic-cell reprogramming.

    Evidence iPSC reprogramming assay, EZH2/H3K27me3 westerns, knockdown/overexpression, CDK inhibitor expression analysis

    PMID:34486784

    Open questions at the time
    • Mechanism by which SPY1 activates EZH2 undefined
    • Single lab
  17. 2022 Medium

    Implicated Spy1 in neuroprotection, showing MDM2-mediated degradation of Spy1 in ALS neurons and that Spy1 overexpression suppresses ferroptosis via the GCH1/BH4 axis and reduced TFR1-mediated iron import, delaying disease in mice.

    Evidence Ubiquitination assay, MDM2 manipulation, lipid peroxidation and GCH1/TFR1 readouts, ALS transgenic mouse

    PMID:36443440

    Open questions at the time
    • Direct link between Spy1 and GCH1/TFR1 regulation not biochemically defined
    • Single lab
  18. 2022 Low

    Correlated Spy1 with c-Myc protein stabilisation in triple-negative breast cancer and chemoresistance, but without defining the stabilisation mechanism.

    Evidence shRNA knockdown, c-Myc protein westerns, chemotherapy sensitivity assays, tissue microarray

    PMID:36029387

    Open questions at the time
    • Mechanism of c-Myc stabilisation not biochemically defined
    • Co-regulation observed without direct binding evidence
  19. 2023 Medium

    Demonstrated in vivo that RingoA sets a CDK activity threshold governing exit of adult neural stem cells from quiescence, with deficiency causing quiescent-NSC accumulation and reduced olfactory neurogenesis.

    Evidence RingoA knockout mouse, CDK activity assays, neurogenesis quantification

    PMID:36876138

    Open questions at the time
    • Specific CDK partner and substrates in NSCs not defined
    • Single lab

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved which CDK2 substrates at the SUN1-LINC complex execute telomere-nuclear envelope attachment, and how the same atypical activator selects between cell-cycle, signalling, epigenetic, and scaffolding outputs in different tissues.
  • No meiotic CDK2 substrate identified
  • Mechanism of context-dependent output switching unknown
  • No structure of a Spy1-CDK1 complex

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0098772 molecular function regulator activity 3 GO:0140096 catalytic activity, acting on a protein 3 GO:0060089 molecular transducer activity 2
Localization
GO:0005634 nucleus 2 GO:0005635 nuclear envelope 2
Pathway
R-HSA-1640170 Cell Cycle 3 R-HSA-5357801 Programmed Cell Death 3 R-HSA-1474165 Reproduction 2 R-HSA-73894 DNA Repair 2
Partners
CDK1CDK2CDKN1BCLIP1MDM2NEDD4STMN2SUN1
Complex memberships
CDK2-Spy1 complexSUN1-SPDYA-CDK2 complexp27-CDK2-Spy1 complex

Evidence

Reading pass · 19 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2003 Spy1 directly interacts with the CDK inhibitor p27Kip1 (confirmed by yeast two-hybrid, in vitro pulldown with bacterially expressed proteins, and co-immunoprecipitation of endogenous and transfected proteins in mammalian cells), and this interaction allows Spy1 to overcome p27-induced G1 arrest to promote CDK2-dependent DNA synthesis; proliferative effects of Spy1 require endogenous p27. Yeast two-hybrid screen, in vitro pulldown (bacterially expressed proteins), co-immunoprecipitation, CDK2 histone H1 kinase assay, p27-null cell rescue experiments Molecular biology of the cell High 12972555
2003 Human Spy1 binds and activates CDK2 during the DNA damage response; Spy1 overexpression enhances cell viability after genotoxic damage (camptothecin, cisplatin, hydroxyurea), and a dominant-negative CDK2 overrides Spy1 function, demonstrating that CDK2 activation is required for Spy1-mediated cell survival. Clonogenic survival assays, comet assays, dominant-negative CDK2 expression, siRNA knockdown, endogenous protein level analysis by western blot Cancer research Medium 12839962
2006 Spy1 expression suppresses UV-induced caspase-3 activation and apoptosis, allows UV-irradiation-resistant DNA synthesis and mitotic entry (histone H3 phosphorylation), and inhibits phosphorylation of Chk1, RPA, and histone H2A.X; mutation of the conserved Speedy/Ringo box (CDK2-interaction domain) abrogates these effects, placing CDK2 activation upstream of checkpoint bypass and apoptosis suppression. Inducible Spy1 expression system, flow cytometry, western blot for checkpoint markers, Speedy/Ringo box point mutant The Journal of biological chemistry Medium 16951407
2007 Spy1 activates CDK2 to phosphorylate p27Kip1 at T187 in vitro using recombinant proteins; Spy1/CDK2 added to a preformed inhibited cyclin E/CDK2/p27 complex promotes T187 phosphorylation; in cells, inducible Spy1 expression reduces endogenous p27 and exogenous p27WT but not p27T187A, and synchronised cells show enhanced T187 phosphorylation and p27 degradation in late G1 and S-phase. In vitro kinase assay with recombinant proteins, inducible expression system, synchronized HeLa cell analysis, T187A mutant control, western blot Cell cycle High 17671428
2009 Spy1-mediated inhibition of apoptosis and bypass of UV-induced checkpoint activation requires the presence of functional p53 and the CDK inhibitor p21; Spy1 expression prevents nucleotide excision repair of cyclobutane pyrimidine dimers, increases cellular mutation frequency, reduces cyclin E-induced γH2A.X foci formation, and knockdown of endogenous Spy1 itself leads to γH2A.X foci, Chk1 phosphorylation, and proliferation defects. siRNA knockdown, p53/p21 genetic requirement analysis, CPD repair assay, mutation frequency assay, γH2A.X and Chk1 phosphorylation western blot Cell cycle Medium 19106603
2012 Spy1 facilitates mammary cell transformation in a manner dependent on activation of CDK1, with subsequent inhibition of the anti-apoptotic regulator FOXO1; knockdown of Spy1 impairs breast cancer cell proliferation. CDK1 activity assays, FOXO1 functional analysis, shRNA knockdown, in vivo mouse mammary tumor model BMC cancer Medium 22280365
2015 Spy1 directly binds SCG10 (a microtubule-destabilising protein/axonal maintenance factor) and mediates SCG10 phosphorylation and proteasomal degradation in a partially JNK-dependent manner following sciatic nerve injury; inhibition of Spy1 attenuates SCG10 phosphorylation and delays axonal degeneration. Co-immunoprecipitation, in vitro binding assay, western blot for phosphorylation/degradation, Spy1 inhibition with phenotypic rescue The Journal of biological chemistry Medium 25869138
2015 Spy1 interacts with CLIPR-59 in glioblastoma cells, sequestering CLIPR-59 away from CYLD (a K63-deubiquitinase); knockdown of Spy1 increases CLIPR-59-CYLD association, enhances K63-deubiquitination of RIP1, and promotes caspase-8/caspase-3 activation, thereby sensitising GBM cells to TNF-α-induced apoptosis. Co-immunoprecipitation, siRNA knockdown, ubiquitination assay, caspase activity assays, western blot Cell cycle Medium 26017671
2016 Specific residues in Spy1 mediate direct interaction with CDK2 and separately with p27Kip1; point mutations disrupting each interaction reduce endogenous CDK2 activity and Spy1-mediated proliferation; only the p27-interaction mutation abolishes Spy1-mediated p27 destabilisation, while disruption of either interaction slows Spy1-driven tumorigenesis in vivo. Site-directed mutagenesis, co-immunoprecipitation, CDK2 kinase assay, p27 stability assay, in vivo mouse tumour model Cell cycle Medium 26771716
2017 Crystal structures of the Cdk2-Spy1 binary complex and the p27-Cdk2-Spy1 ternary complex reveal that Spy1 activates Cdk2 by inducing structural changes that bypass the requirement for activation-loop phosphorylation; Spy1 lacks the cyclin-binding groove that mediates p27 inhibition and substrate cyclin-docking, explaining why Cdk-Spy1 complexes are resistant to p27 and have altered substrate specificity; Spy1 mutations that ablate Cdk2 binding also abolish cell proliferation. X-ray crystallography, in vitro CDK2 kinase assay, site-directed mutagenesis, cell proliferation assay The EMBO journal High 28666995
2017 Spy1 activates ERK1/2 in a MEK-independent manner; Spy1-mediated ERK activation increases ligand-independent phosphorylation and activation of estrogen receptor α, correlating with decreased tamoxifen sensitivity in breast cancer cells. ERK1/2 phosphorylation assays, MEK inhibitor controls, ERα phosphorylation assays, tamoxifen sensitivity assays, Spy1 overexpression/knockdown Oncotarget Medium 28423577
2019 p53 controls Spy1 protein levels via proteasomal degradation mediated in part by the E3 ubiquitin ligase Nedd4; loss or mutation of p53 allows Spy1 accumulation, leading to mammary hyperplasia; in a transgenic mouse model, sustained Spy1 elevation results in increased mammary proliferation and tumour susceptibility. Transgenic mouse model, in vitro protein stability assays, Nedd4 co-immunoprecipitation/ubiquitination, p53 knockout/mutation analysis Breast cancer research Medium 31829284
2020 SPDYA (Speedy A) directly interacts with SUN1 at the nuclear envelope via its Ringo domain and recruits CDK2 to SUN1; the SPDYA-binding site on SUN1 is in the N-terminal domain, close to the MAJIN-binding site; CDK2 inhibitors decrease the SUN1-MAJIN interaction, suggesting SPDYA-CDK2 promotes telomere-nuclear envelope attachment during meiosis. Co-immunoprecipitation, domain-mapping experiments, CDK2 inhibitor treatment, binding-site mutagenesis Frontiers in cell and developmental biology Medium 33015044
2021 SPDYA directly interacts with SUN1; the crystal structure of the human SUN1-SPDYA-CDK2 ternary complex was determined; SUN1 mutations that disrupt SPDYA binding in mice impair telomere-LINC complex attachment, abolish the ring-shaped telomere supramolecular architecture at the nuclear envelope, and cause defects in homologous pairing and synapsis during meiosis prophase I. X-ray crystallography (ternary complex), site-directed mutagenesis of SUN1, mouse genetic model (SPDYA-binding-deficient SUN1 knockin), meiosis phenotype analysis Nature communications High 34039995
2021 SPY1 activates the epigenetic regulator EZH2 to promote H3K27me3-mediated repression of CDK inhibitors CDKN1A and CDKN2A, overriding senescence during reprogramming; SPY1 is required for efficient reprogramming of mouse fibroblasts to induced pluripotent stem cells. iPSC reprogramming assay, EZH2 activity/H3K27me3 western blot, Spy1 knockdown/overexpression, CDKN1A/CDKN2A expression analysis Stem cells Medium 34486784
2022 SPY1 protein is degraded via ubiquitination mediated by MDM2 (an E3 ubiquitin ligase) in ALS motor neurons; overexpression of Spy1 suppresses ferroptosis in hSOD1G93A cells by restoring the GCH1/BH4 axis (a ferroptosis resistance pathway) and reducing TFR1-mediated iron import, decreasing lipid peroxidation; neuron-specific Spy1 overexpression delays disease onset and extends survival in ALS transgenic mice. Ubiquitination assay, MDM2 knockdown/overexpression, lipid peroxidation measurement, GCH1/TFR1 western blot and activity assays, transgenic mouse model with neurological scoring Cell death and differentiation Medium 36443440
2022 Spy1 stabilises c-Myc protein in TNBC cells; knockdown of Spy1 reduces c-Myc protein levels and sensitises TNBC cells to chemotherapy; Spy1 protein levels positively correlate with c-Myc protein (not mRNA) levels in TNBC patient samples. shRNA knockdown, western blot for c-Myc protein, chemotherapy sensitivity assays, tissue microarray Breast cancer research and treatment Low 36029387
2023 RingoA (Spy1 ortholog in mouse) regulates exit from quiescence in adult neural stem cells; RingoA expression increases CDK activity and facilitates cell cycle entry; RingoA-deficient mice show reduced olfactory neurogenesis and accumulation of quiescent NSCs, placing RingoA as a CDK activity threshold regulator for NSC quiescence-to-activation transition. RingoA knockout mouse, CDK activity assays, neurogenesis quantification, quiescent NSC accumulation analysis iScience Medium 36876138
2014 CRMP1 interacts with Spy1 (identified by yeast two-hybrid and confirmed by co-IP); Spy1 affects CDK5-mediated CRMP1 phosphorylation, which disrupts CRMP1-actin association and contributes to Sema3A-induced growth cone collapse and impaired regeneration after sciatic nerve crush. Yeast two-hybrid, co-immunoprecipitation, overexpression/shRNA, CRMP1 phosphorylation assay, growth cone collapse assay, rat sciatic nerve crush model Molecular neurobiology Low 25526860

Source papers

Stage 0 corpus · 42 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2022 SPY1 inhibits neuronal ferroptosis in amyotrophic lateral sclerosis by reducing lipid peroxidation through regulation of GCH1 and TFR1. Cell death and differentiation 136 36443440
2003 Spy1 interacts with p27Kip1 to allow G1/S progression. Molecular biology of the cell 44 12972555
2003 Human Spy1 promotes survival of mammalian cells following DNA damage. Cancer research 38 12839962
2000 Spy1, a histidine-containing phosphotransfer signaling protein, regulates the fission yeast cell cycle through the Mcs4 response regulator. Journal of bacteriology 38 10940030
2008 The Spy1/RINGO family represents a novel mechanism regulating mammary growth and tumorigenesis. Cancer research 37 18483240
2017 Structural basis of divergent cyclin-dependent kinase activation by Spy1/RINGO proteins. The EMBO journal 36 28666995
2006 Spy1 expression prevents normal cellular responses to DNA damage: inhibition of apoptosis and checkpoint activation. The Journal of biological chemistry 36 16951407
2014 The cyclin-like protein Spy1 regulates growth and division characteristics of the CD133+ population in human glioma. Cancer cell 34 24434210
2021 The SUN1-SPDYA interaction plays an essential role in meiosis prophase I. Nature communications 31 34039995
2007 Spy1 enhances phosphorylation and degradation of the cell cycle inhibitor p27. Cell cycle (Georgetown, Tex.) 31 17671428
2009 The atypical CDK activator Spy1 regulates the intrinsic DNA damage response and is dependent upon p53 to inhibit apoptosis. Cell cycle (Georgetown, Tex.) 27 19106603
2012 Expression of Spy1 protein in human non-Hodgkin's lymphomas is correlated with phosphorylation of p27 Kip1 on Thr187 and cell proliferation. Medical oncology (Northwood, London, England) 25 22492278
2015 Spy1 induces de-ubiquitinating of RIP1 arrest and confers glioblastoma's resistance to tumor necrosis factor (TNF-α)-induced apoptosis through suppressing the association of CLIPR-59 and CYLD. Cell cycle (Georgetown, Tex.) 22 26017671
2014 CRMP1 Interacted with Spy1 During the Collapse of Growth Cones Induced by Sema3A and Acted on Regeneration After Sciatic Nerve Crush. Molecular neurobiology 21 25526860
2018 Spy1, a unique cell cycle regulator, alters viability in ALS motor neurons and cell lines in response to mutant SOD1-induced DNA damage. DNA repair 18 30630676
2012 The cyclin-like protein Spy1/RINGO promotes mammary transformation and is elevated in human breast cancer. BMC cancer 18 22280365
2017 Receptor activator of nuclear factor kappa B (RANK) expression in primary breast cancer correlates with recurrence-free survival and development of bone metastases in I-SPY1 (CALGB 150007/150012; ACRIN 6657). Breast cancer research and treatment 17 28577080
2014 Serotype-independent protection against pneumococcal infections elicited by intranasal immunization with ethanol-killed pneumococcal strain, SPY1. Journal of microbiology (Seoul, Korea) 17 24682994
2014 The atypical cell cycle regulator Spy1 suppresses differentiation of the neuroblastoma stem cell population. Oncoscience 16 25594028
2013 Cell adhesion to fibronectin down-regulates the expression of Spy1 and contributes to drug resistance in multiple myeloma cells. International journal of hematology 16 24037419
2012 Spy1 is frequently overexpressed in malignant gliomas and critically regulates the proliferation of glioma cells. Journal of molecular neuroscience : MN 16 22447439
2020 Tethering of Telomeres to the Nuclear Envelope Is Mediated by SUN1-MAJIN and Possibly Promoted by SPDYA-CDK2 During Meiosis. Frontiers in cell and developmental biology 15 33015044
2018 Protective Regulatory T Cell Immune Response Induced by Intranasal Immunization With the Live-Attenuated Pneumococcal Vaccine SPY1 via the Transforming Growth Factor-β1-Smad2/3 Pathway. Frontiers in immunology 15 30116243
2014 Mucosal immunization with the live attenuated vaccine SPY1 induces humoral and Th2-Th17-regulatory T cell cellular immunity and protects against pneumococcal infection. Infection and immunity 15 25312946
2015 Spy1 participates in the proliferation and apoptosis of epithelial ovarian cancer. Journal of molecular histology 13 26644004
2019 The atypical cyclin-like protein Spy1 overrides p53-mediated tumour suppression and promotes susceptibility to breast tumourigenesis. Breast cancer research : BCR 12 31829284
2017 The cyclin-like protein, SPY1, regulates the ERα and ERK1/2 pathways promoting tamoxifen resistance. Oncotarget 12 28423577
2008 Peripheral nerve lesion induces an up-regulation of Spy1 in rat spinal cord. Cellular and molecular neurobiology 12 19082704
2016 Direct interactions with both p27 and Cdk2 regulate Spy1-mediated proliferation in vivo and in vitro. Cell cycle (Georgetown, Tex.) 10 26771716
2015 Spy1 Protein Mediates Phosphorylation and Degradation of SCG10 Protein in Axonal Degeneration. The Journal of biological chemistry 8 25869138
2012 Isolation and functional analysis of spy1 responsible for pristinamycin yield in Streptomyces pristinaespiralis. Journal of microbiology and biotechnology 8 22573156
2022 Stabilization of c-Myc by the atypical cell cycle regulator, Spy1, decreases efficacy of breast cancer treatments. Breast cancer research and treatment 7 36029387
2023 The atypical CDK activator RingoA/Spy1 regulates exit from quiescence in neural stem cells. iScience 6 36876138
2016 Attenuated Streptococcus pneumoniae vaccine candidate SPY1 promotes dendritic cell activation and drives a Th1/Th17 response. Immunology letters 6 27609353
2010 Protective roles and Pap1-dependent regulation of the Schizosaccharomyces pombe spy1 gene under nitrosative and nutritional stresses. Molecular biology reports 6 20563653
2017 Heterologous prime-boost immunization with live SPY1 and DnaJ protein of Streptococcus pneumoniae induces strong Th1 and Th17 cellular immune responses in mice. Journal of microbiology (Seoul, Korea) 5 28956354
2012 Temporal-spatial expressions of Spy1 in rat sciatic nerve after crush. Cellular and molecular neurobiology 5 23129232
2021 The cyclin-like protein SPY1 overrides reprogramming induced senescence through EZH2 mediated H3K27me3. Stem cells (Dayton, Ohio) 4 34486784
2023 IL-27 mediates immune response of pneumococcal vaccine SPY1 through Th17 and memory CD4+T cells. iScience 3 37588169
2025 Targeting cyclin-dependent kinase 2 (CDK2) interactions with cyclins and Speedy 1 (Spy1) for cancer and male contraception. Future medicinal chemistry 2 40034037
2015 [Identification of a point mutation in the promoter region of cps operon responsible for capsular polysaccharide deficiency in Streptococcus pneumniae SPY1]. Wei sheng wu xue bao = Acta microbiologica Sinica 1 26563004
2014 SPY1-mediated symmetric cell division promotes BTIC self-renewal. Cancer discovery 0 24596205

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