Affinage

Showing YBX2MSY2 is a alias.

YBX2

Y-box-binding protein 2 · UniProt Q9Y2T7

Length
364 aa
Mass
38.5 kDa
Annotated
2026-06-11
44 papers in source corpus 27 papers cited in narrative 27 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 8/9 claims corpus-supported (89%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

YBX2 (MSY2/Contrin) is a germ cell-enriched Y-box protein that serves as a global stabilizer and translational repressor of stored mRNAs, packaging them into translationally silent cytoplasmic ribonucleoprotein particles during gametogenesis (PMID:9780336, PMID:10772793, PMID:15031116). It binds mRNA through its cold-shock domain together with C-terminal basic/aromatic islands, with the intact CSD required for cytoplasmic localization and an additional C-terminal island needed for cytoskeletal/Triton-insoluble retention; intact mRNA bridges YBX2 to these cytoplasmic structures (PMID:12297523, PMID:12648488). Binding occurs largely with limited sequence specificity, allowing multiple YBX2 molecules to coat a single transcript, but YBX2 with MSY4 can also recognize a defined 7-8 nt Y-box recognition site in 3'UTRs to repress specific transcripts such as Prm1 in a position-dependent manner (PMID:11564883, PMID:12297523, PMID:26773323). RNA-binding activity is mechanistically required for its stabilizing function: knockout reduces postmeiotic and maternal mRNA pools, and wild-type but not RNA-binding-deficient YBX2 rescues mRNA stability in null oocytes (PMID:17035640, PMID:21613634). Genetic deletion causes male and female infertility, with postmeiotic spermatogenic arrest and oocyte/follicle defects driven by mRNA destabilization rather than transcriptional failure (PMID:15824319, PMID:17035640). YBX2 function is switched by phosphorylation: CDK1-mediated phosphorylation during oocyte maturation converts it from a stabilizer to a trigger of maternal mRNA degradation (PMID:18606161). Beyond germ cells, YBX2 acts as an m5C RNA reader via W100 that promotes target mRNA stability and undergoes liquid-liquid phase separation (PMID:38933916, PMID:42011916), partners with PAIP1 to reactivate translation of masked spermatogenic mRNAs (PMID:30295753), and cooperates with the m6A reader YTHDF2 to destabilize specific transcripts (PMID:37030331). In brown adipose tissue, AMPK/Akt2 phosphorylation stabilizes YBX2 to promote PGC1α mRNA stability and translation of glycolytic enzyme mRNAs (PMID:28970281, PMID:37860762). Transcription of Ybx2 itself is driven by BNC1 and by N-Myc binding to its promoter (PMID:33211273, PMID:16624424).

Mechanistic history

Synthesis pass · year-by-year structured walk · 25 steps
  1. 1998 High

    Establishing that YBX2 is a germ cell-specific Y-box protein with defined developmental expression set the stage for a gamete-specific RNA-handling function.

    Evidence Northern, immunoblot, and in situ analysis of isolated germ cell populations

    PMID:9780336

    Open questions at the time
    • Molecular function not yet defined
    • No direct RNA target identified at this stage
  2. 2000 High

    Identifying that YBX2 forms a complex with MSY4 on non-polysomal mRNPs linked the protein to translationally repressed, stored mRNAs.

    Evidence EMSA, co-IP, polysome analysis, yeast three-hybrid

    PMID:10772793

    Open questions at the time
    • Specific RNA sequence target not yet mapped
    • Mechanism of repression unresolved
  3. 1999 High

    Showing that an associated kinase phosphorylates YBX2 and that dephosphorylation reduces RNA binding introduced phosphorylation as a regulator of its mRNA-binding state.

    Evidence Co-IP, in vitro kinase assay, Northwestern, in vitro translation repression

    PMID:10076007

    Open questions at the time
    • Identity of the kinase not established
    • In vivo relevance of phosphoregulation not yet shown
  4. 2001 High

    Defining the 7-8 nt Y-box recognition site in the Prm1 3'UTR, with in vitro mutagenesis and transgenic validation, established sequence-specific repression alongside general binding.

    Evidence EMSA, competition binding, yeast three-hybrid, transgenic mouse reporter

    PMID:11564883

    Open questions at the time
    • Generality of YRS across the transcriptome unclear
    • Co-factor requirement not yet defined
  5. 2001 High

    Localizing YBX2 to the oocyte cytoplasm/cytoskeletal fraction and documenting maturation-coupled phosphorylation tied its activity to oocyte developmental transitions.

    Evidence Immunoblot, RT-PCR, MSY2-EGFP microinjection, confocal imaging, cytoskeletal fractionation

    PMID:11566752

    Open questions at the time
    • Functional consequence of maturation phosphorylation not yet established
    • Kinase responsible unidentified
  6. 2002 High

    Demonstrating that recombinant YBX2 binds mRNA with limited sequence specificity and represses translation at physiological ratios established its capacity as a global mRNA-coating repressor.

    Evidence Gel shift, filter binding, in vitro translation/luciferase with recombinant protein

    PMID:12297523

    Open questions at the time
    • In vivo target spectrum not addressed
    • Reconciliation of sequence-specific vs non-specific binding modes incomplete
  7. 2003 High

    Mapping the CSD requirement for cytoplasmic localization and the C-terminal island for insoluble retention, with RNase release of YBX2, established that mRNA binding underpins its cytoplasmic anchoring.

    Evidence Domain-mutant MSY2-EGFP microinjection, confocal imaging, Triton extraction, RNase A treatment, RNA EMSA

    PMID:12648488

    Open questions at the time
    • Identity of cytoskeletal anchoring structures not defined here
    • Structural basis of CSD binding not resolved
  8. 2004 High

    RNAi knockdown reducing total mRNA and impairing protein synthesis provided the first in vivo functional evidence that YBX2 stabilizes maternal mRNAs.

    Evidence Transgenic Zp3-driven RNAi, immunoblot, microinjection, Ca2+ imaging, mRNA and protein synthesis assays

    PMID:15031116

    Open questions at the time
    • Residual protein from partial knockdown
    • Direct vs indirect effect on individual targets not separated
  9. 2005 High

    Defining the YBX2-bound germ cell mRNA subset and linking Y-box promoters to cytoplasmic storage connected nuclear transcription to cytoplasmic mRNA masking.

    Evidence IP plus suppressive subtractive hybridization, ChIP, transgenic GFP reporter

    PMID:15665108

    Open questions at the time
    • Mechanism coupling transcription to storage not biochemically defined
    • Direct vs indirect target distinction limited
  10. 2005 High

    Germline knockout producing male and female infertility with reduced postmeiotic mRNA established YBX2 as essential for gametogenesis.

    Evidence Gene targeting, histology, RT-PCR, apoptosis and fertility assays

    PMID:15824319

    Open questions at the time
    • Causal chain from mRNA loss to arrest not yet dissected
    • Stage-specific requirements not separated
  11. 2006 High

    Showing that in null testes mRNAs shift from RNP storage to polysomes and decline established mRNA instability, not transcriptional defects, as the cause of arrest.

    Evidence Polysome/RNP fractionation, real-time RT-PCR, transcription and transport assays in knockout

    PMID:17035640

    Open questions at the time
    • Degradation machinery engaged not identified
    • Selectivity among targets not fully mapped
  12. 2006 Medium

    Identifying N-Myc-driven transcription via an E-box and a C-terminal region controlling cytoplasmic retention added upstream regulation and a nuclear-exclusion determinant.

    Evidence Promoter deletion/reporter, in vivo footprinting, Myc co-transfection, localization analysis in germ cell tumor lines

    PMID:16624424

    Open questions at the time
    • Cell line context may not reflect normal germ cells
    • Physiological role of nuclear translocation unclear
  13. 2008 High

    Demonstrating that CDK1 phosphorylation triggers maturation-associated maternal mRNA degradation defined a phospho-switch converting YBX2 from stabilizer to destabilizer.

    Evidence Cdc2a microinjection, roscovitine inhibition, phosphomutant/phosphomimetic MSY2, immunoblot, mRNA quantification

    PMID:18606161

    Open questions at the time
    • Mechanism linking phosphorylation to degradation machinery unknown
    • Which targets are selected for degradation not defined
  14. 2009 Medium

    Discovering YBX2 association with a class of MIWI-independent small RNAs broadened its RNA-binding repertoire beyond mRNA.

    Evidence RNA-IP, small RNA-seq, fractionation, analysis in Miwi and Msy2 null mice

    PMID:19597149

    Open questions at the time
    • Functional significance of MSY-RNA binding not established
    • Single lab
  15. 2011 High

    Rescue with wild-type but not RNA-binding-deficient YBX2 in null oocytes directly proved RNA-binding activity is required for mRNA stabilization.

    Evidence Knockout analysis, WT and mutant MSY2 microinjection, mRNA stability assays, spindle imaging, transcriptome profiling

    PMID:21613634

    Open questions at the time
    • Stabilization mechanism (sequestration vs decay protection) not resolved
    • Link to spindle defect not mechanistically defined
  16. 2016 High

    Showing that 3'UTR binding-site position dictates repression and that targets are prematurely polysome-loaded in knockouts established position-dependent cooperative repression.

    Evidence Transgenic 3'UTR mutants, polysome analysis in knockout, reporter assays

    PMID:26773323

    Open questions at the time
    • Identity of position-dependent co-factors not defined
    • Single lab
  17. 2016 Medium

    Identifying PADI6 as required to anchor the mRNA-YBX2 complex to cytoplasmic lattices placed YBX2 storage in a defined oocyte structure.

    Evidence Padi6 knockout, Triton fractionation, RNase A treatment, immunostaining

    PMID:27929740

    Open questions at the time
    • Ternary complex inferred indirectly
    • Direct PADI6-YBX2 contact not shown
  18. 2017 High

    Extending YBX2 function to brown adipocytes, where it stabilizes PGC1α mRNA and drives adipogenesis, established a somatic mRNA-stability role.

    Evidence shRNA, overexpression, knockout mice, RIP-seq

    PMID:28970281

    Open questions at the time
    • Mechanism of selective stabilization unclear
    • Direct vs indirect targets not fully separated
  19. 2019 High

    Demonstrating that PAIP1 partnership reactivates translation of YBX2-masked mRNAs defined how stored transcripts are switched on for spermiogenesis.

    Evidence Reciprocal co-IP, sequential RNA-IP, in vitro translation rescue, immunolocalization

    PMID:30295753

    Open questions at the time
    • In vivo requirement of PAIP1 for reactivation not tested
    • Single lab
  20. 2021 High

    Crystal structure identifying W100 as the m5C-binding site and linking m5C RNA to phase separation defined YBX2 as an m5C reader driving LLPS.

    Evidence Crystal structure, m5C RNA-binding assays, in vivo/in vitro LLPS, W100 mutagenesis

    PMID:38933916

    Open questions at the time
    • Physiological consequence of m5C-driven LLPS in vivo not established
    • Single lab
  21. 2022 Medium

    Showing ALYREF exports m5C-marked YBX2 mRNA to increase YBX2 protein, which inhibits adipogenesis, added a post-transcriptional layer to YBX2 regulation.

    Evidence Bisulfite/m5C sequencing, adipogenesis assays, ALYREF perturbation, fractionation

    PMID:35962235

    Open questions at the time
    • Mechanism by which YBX2 inhibits adipogenesis not defined
    • Single lab
  22. 2022 Medium

    Detecting YBX2 at the multi-cilia axoneme interacting with RIBC2, with ciliary defects on knockdown, raised a non-canonical structural/ciliary role.

    Evidence IP/MS, morpholino knockdown, bead flow assay, immunostaining in Xenopus

    PMID:36508087

    Open questions at the time
    • Non-canonical context for YBX2; mechanism unclear
    • Mammalian relevance not tested
  23. 2023 Medium

    Showing YBX2 recruits YTHDF2 to cytoplasmic granules and cooperatively destabilizes HSPA6 mRNA defined a co-reader destabilization mechanism in cancer cells.

    Evidence Ectopic expression, transcriptomics, RNA stability assays, co-IP/co-localization, YTHDF2 knockdown

    PMID:37030331

    Open questions at the time
    • Cancer cell line context
    • Endogenous relevance not established
  24. 2023 Medium

    Identifying AMPK/Akt2 phosphorylation sites that block YBX2 ubiquitination and showing 5'-UTR binding to glycolytic mRNAs defined how β-adrenergic signaling stabilizes YBX2 to drive glycolysis.

    Evidence Phosphosite mutagenesis, ubiquitination assays, 5'-UTR RNA-binding, metabolic assays in brown adipocytes

    PMID:37860762

    Open questions at the time
    • In vivo contribution of each phosphosite not separated
    • Single lab
  25. 2026 Medium

    Demonstrating YBX2 reads NSUN5-installed m5C on SCD1 mRNA to prolong its half-life and suppress ferroptosis extended the m5C-reader stabilizing function to a disease context.

    Evidence Bisulfite sequencing, RIP, mRNA half-life assays, NSUN5 perturbation, ferroptosis assays

    PMID:42011916

    Open questions at the time
    • Generality of m5C-stabilization across targets unclear
    • Single lab, disease model

Open questions

Synthesis pass · forward-looking unresolved questions
  • How phosphorylation, m5C/m6A reader cooperation, and phase separation are integrated to switch individual transcripts between storage, stabilization, and degradation remains unresolved.
  • No unified model of transcript-selective fate control
  • Degradation machinery downstream of CDK1 phosphorylation unidentified
  • Structural basis of cooperative position-dependent repression undefined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003723 RNA binding 6 GO:0045182 translation regulator activity 4 GO:0140313 molecular sequestering activity 3 GO:0003677 DNA binding 1
Localization
GO:0005829 cytosol 3 GO:0005856 cytoskeleton 3 GO:0005929 cilium 1
Pathway
R-HSA-392499 Metabolism of proteins 4 R-HSA-8953854 Metabolism of RNA 4 R-HSA-1474165 Reproduction 3
Complex memberships
YBX2-MSY4 mRNPcytoplasmic lattice mRNP (with PADI6)

Evidence

Reading pass · 27 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1998 MSY2 (YBX2) is a germ cell-specific Y-box protein containing a cold shock domain and four basic/aromatic islands; it is expressed maximally in postmeiotic round spermatids and in diplotene-stage/mature oocytes, and is maternally inherited in the one-cell embryo but degraded by the late two-cell stage coincident with bulk maternal mRNA degradation. Northern blotting, immunoblotting, in situ analysis of isolated germ cell populations Biology of reproduction High 9780336
2000 MSY2 (YBX2) forms a sequence-specific RNA-binding complex with MSY4 on non-polysomal mRNPs; the complex co-immunoprecipitates and is associated with translationally repressed mRNAs, consistent with a role in mRNA storage. Electrophoretic mobility shift assay (EMSA), immunoprecipitation, polysome analysis, yeast three-hybrid system Developmental biology High 10772793
1999 In male germ cells, MSY2 is associated on non-polysomal mRNAs with a co-immunoprecipitating kinase activity that phosphorylates MSY2 in vitro; dephosphorylation of MSY2 decreases its RNA-binding affinity, suggesting phosphorylation state regulates mRNA binding. MSY2-bound non-polysomal RNP proteins repress translation of exogenous mRNAs in vitro. Co-immunoprecipitation, in vitro kinase assay, Northwestern blotting, in vitro translation assay, reconstitution Nucleic acids research High 10076007
2001 MSY2 and MSY4 bind a specific 7-8 nucleotide sequence (Y-box recognition site, YRS: U(AC)C(A)CAU(C)CA) in the 3' UTR of the translationally repressed protamine 1 mRNA; single nucleotide mutations abolish binding in vitro, and disruption of the YRS in transgenic mice relieves Prm1-like translational repression in vivo. EMSA, competition binding, yeast three-hybrid system, transgenic mouse reporter assay Molecular and cellular biology High 11564883
2001 MSY2 protein is primarily cytoplasmic in mouse oocytes (~75% associated with cytoskeletal/Triton-insoluble preparations); it is phosphorylated during oocyte maturation and dephosphorylated following fertilization. MSY2 constitutes ~2% of total oocyte protein. Immunoblotting, reverse transcription-PCR, microinjection of MSY2-EGFP mRNA, confocal microscopy, cytoskeletal fractionation Biology of reproduction High 11566752
2002 Bacterially expressed recombinant MSY2 binds mRNA with limited sequence specificity (multiple molecules bind one Prm1 mRNA) and represses translation in vitro at protein:mRNA ratios approximating those in oocytes. Gel shift assay, filter binding assay, in vitro translation/luciferase assay with recombinant protein Biology of reproduction High 12297523
2003 An intact cold-shock domain (CSD) with its two RNA-binding motifs is required for cytoplasmic localization of MSY2 in oocytes; an additional C-terminal basic/aromatic island is necessary for Triton-insoluble retention. RNase A treatment releases essentially all endogenous MSY2, demonstrating that intact mRNA is required for cytoplasmic retention. Microinjection of mutant MSY2-EGFP mRNAs, laser-scanning confocal microscopy, Triton extraction, RNase A treatment, RNA EMSA Developmental biology High 12648488
2004 RNAi-mediated reduction of MSY2 in growing oocytes (~60% knockdown) reduces total mRNA by ~20-25%, impairs protein synthesis, and prevents maturation-associated changes in protein synthesis, directly supporting MSY2's role in stabilizing maternal mRNAs in vivo. Transgenic RNAi (Zp3 promoter-driven hairpin dsRNA), immunoblotting, microinjection, Ca2+ imaging, total mRNA quantification, protein synthesis assay Developmental biology High 15031116
2005 MSY2 binds a specific subset of germ cell mRNAs enriched for stored/translationally delayed, male gamete-specific transcripts; chromatin precipitation shows most MSY2 target mRNAs are transcribed from promoters containing the Y-box DNA-binding motif; transgenic reporter experiments show Y-box promoters direct mRNAs into the MSY2-bound cytoplasmic fraction, linking nuclear transcription to cytoplasmic mRNA storage. Immunoprecipitation combined with suppressive subtractive hybridization, chromatin precipitation (ChIP), transgenic mouse GFP reporter assay Proceedings of the National Academy of Sciences of the United States of America High 15665108
2005 Genetic deletion of Msy2 in mice causes male and female infertility: males exhibit post-meiotic spermatogenic arrest with multinucleated spermatids and no spermatozoa, accompanied by large reductions in postmeiotic mRNA levels; females show oocyte loss, anovulation, and follicle defects. Gene targeting/knockout, histology, RT-PCR, apoptosis assays, fertility testing Proceedings of the National Academy of Sciences of the United States of America High 15824319
2006 In Msy2-null male mice, mRNAs are redistributed from RNP storage particles to polysomes and show markedly decreased abundance; transcription, mRNA processing, and intracellular transport are normal, indicating that increased mRNA instability (not transcriptional defects) causes spermatogenic arrest. Polysome/RNP fractionation, real-time RT-PCR, transcriptional assays, mRNA transport assays Biology of reproduction High 17035640
2008 CDC2A (CDK1)-mediated phosphorylation of MSY2 triggers the maturation-associated degradation of maternal mRNAs during mouse oocyte maturation: CDK1 activation induces MSY2 phosphorylation and mRNA degradation; CDK1 inhibition blocks both; non-phosphorylatable dominant-negative MSY2 prevents mRNA degradation, while constitutively active MSY2 induces degradation in the absence of maturation. Microinjection of Cdc2a mRNA, CDK1 inhibition with roscovitine, expression of phosphomutant/phosphomimetic MSY2, immunoblotting, mRNA quantification Developmental biology High 18606161
2009 MSY2 binds a class of small RNAs (~25-31 nt, termed MSY-RNAs) that are MIWI-independent, often initiate with 5' adenine, are expressed in germ and somatic cells, map within annotated genes and to piRNA clusters, and are present in nuclei (enriched in chromatin) and cytoplasm (in both RNPs and polysomes). RNA immunoprecipitation, small RNA sequencing, fractionation (chromatin, RNP, polysome), analysis in Miwi and Msy2 null mice Proceedings of the National Academy of Sciences of the United States of America Medium 19597149
2011 In Msy2-null oocytes, total poly(A)-containing RNA is reduced by ~25%; injection of wild-type MSY2 into mutant oocytes rescues mRNA stability, but a RNA-binding-deficient mutant MSY2 does not, directly demonstrating that RNA-binding activity is required for MSY2-mediated mRNA stabilization. Loss of MSY2 also causes failure of transcription quiescence during oocyte growth and aberrant spindle formation. Knockout mouse analysis, microinjection of wild-type and mutant MSY2 mRNA, mRNA stability assays, spindle imaging, transcriptome profiling Biology of reproduction High 21613634
2016 YBX2 binding sites positioned near the 3' ends of Prm1 and Smcp mRNA 3'UTRs are required for translational repression in round spermatids; moving the same binding sites to upstream positions produces negligible repression, demonstrating position-dependent cooperative interactions with co-factors. In Ybx2-knockout round spermatids, Prm1 and Smcp mRNAs are prematurely recruited to polysomes. Transgenic mice with 3'UTR mutations, polysome analysis of Ybx2-knockout mice, transgenic reporter assays Molecular reproduction and development High 26773323
2016 PADI6 is required for anchoring the mRNA-MSY2 complex to oocyte cytoplasmic lattices (CPLs): in Padi6-KO oocytes, the abundance of MSY2 in the Triton-insoluble fraction is markedly decreased; RNase A treatment of permeabilized oocytes releases both PADI6 and MSY2, indicating mRNA bridges the MSY2-CPL interaction. Padi6 knockout mouse analysis, Triton extraction/fractionation, RNase A treatment, immunostaining Cell cycle Medium 27929740
2017 Ybx2 knockdown in brown adipocytes blocks brown adipogenesis and destabilizes PGC1α mRNA; Ybx2 overexpression promotes brown adipose tissue (BAT) marker expression; RNA immunoprecipitation identifies a set of mRNA targets including Pgc1α that are destabilized upon Ybx2 depletion during cold-induced activation, establishing YBX2 as a regulator of mRNA stability in BAT. shRNA knockdown, overexpression, Ybx2-knockout mice, RNA-immunoprecipitation, RNA sequencing Diabetes High 28970281
2019 YBX2 interacts with PAIP1 (a translation enhancer) in vitro and in vivo in mouse testes; PAIP1 co-localizes with YBX2 in round spermatids; sequential RNA immunoprecipitation identifies spermiogenic mRNAs indirectly bound by PAIP1 through YBX2; translation of YBX2-masked mRNAs is blocked by YBX2 alone but reinitiated when YBX2 is co-expressed with PAIP1 in vitro. Co-immunoprecipitation (in vitro and in vivo), sequential RNA-IP, in vitro translation assay, immunolocalization Biology of reproduction High 30295753
2020 BNC1 transcription factor directly binds to BNC1 binding elements in the Ybx2 promoter to activate Ybx2 transcription; demonstrated by chromatin immunoprecipitation on mouse testicular tissue and luciferase reporter assays; Bnc1 mutation or knockdown reduces YBX2 protein levels. Chromatin immunoprecipitation (ChIP), luciferase reporter assay, Bnc1 mutant mouse model, siRNA knockdown, Western blotting Reproductive sciences Medium 33211273
2021 YBX2 serves as a mammalian m5C RNA-binding protein; m5C-marked RNA enhances YBX2-dependent liquid-liquid phase separation (LLPS) in vivo and in vitro; crystal structure identifies W100 as the distinct m5C binding site critical for mediating YBX2 phase separation. Crystal structure determination, m5C RNA-binding assays, LLPS assays in vivo and in vitro, site-directed mutagenesis (W100) Fundamental research High 38933916
2022 YBX2 mRNA bearing m5C modification is recognized by ALYREF, which exports it from nucleus to cytoplasm, increasing YBX2 protein expression; elevated YBX2 protein inhibits adipogenesis. RNA bisulfite sequencing, m5C profiling, in vitro adipogenesis assays, ALYREF knockdown/overexpression, subcellular fractionation Cellular and molecular life sciences Medium 35962235
2023 YBX2 expressed in endometrial cancer cells forms cytoplasmic granules via its mRNA-binding domain; its cold-shock domain recruits m6A reader YTHDF2 to these granules; YBX2 binding to HSPA6 mRNA reduces its stability, and YTHDF2 knockdown rescues HSPA6 mRNA levels, demonstrating synergistic mRNA destabilization by YBX2 and YTHDF2. Ectopic expression, transcriptomic analysis, RNA stability assays, co-immunoprecipitation/co-localization, YTHDF2 knockdown, Western blotting Experimental cell research Medium 37030331
2023 YBX2 is phosphorylated by AMPK at Thr115 and by Akt2 at Ser137 in brown adipocytes in response to β-adrenergic signaling; these phosphorylations reduce ubiquitination-mediated degradation and stabilize YBX2 protein. Stabilized YBX2 promotes glycolysis by directly binding to 5'-UTRs of glycolytic enzyme mRNAs to modulate their translation. Phosphorylation assays, site-directed mutagenesis, ubiquitination assays, RNA-binding assays (5'-UTR binding), metabolic assays in brown adipocytes iScience Medium 37860762
2006 Cell-type-specific expression of dbpC/Contrin (YBX2) in germ cell tumor lines is regulated by N-Myc, which binds a functional E-box element at -272 to -253 relative to the transcription start site; deletion of the C-terminal region causes nuclear translocation of dbpC/Contrin, which is otherwise primarily cytoplasmic. Promoter deletion/reporter assays, in vivo footprint analysis, co-transfection with Myc expression plasmids, Western blotting, subcellular localization analysis Biochimica et biophysica acta Medium 16624424
2022 YBX2 localizes to the axoneme of multi-cilia in Xenopus laevis embryonic epithelium and interacts with RIBC2; knockdown of Ybx2 results in abnormal ciliary beating and reduced cilia-driven fluid flow; Ybx2 axonemal localization is reduced in Ribc2-depleted multi-cilia. Immunoprecipitation/mass spectrometry, antisense morpholino knockdown, fluorescent bead flow assay, immunostaining, Western blotting in Xenopus Genes & genomics Medium 36508087
2025 In testicular germ cells, YBX2 forms a complex with STK33 as detected by proximity ligation assay; however, STK33 cannot phosphorylate YBX2 in vitro, and the interaction was not recapitulated in co-transfected cultured cells. Immunoprecipitation/mass spectrometry, proximity ligation assay, co-transfection, in vitro phosphorylation assay Reproductive sciences Low 39909973
2026 YBX2 acts as an m5C reader for SCD1 mRNA: NSUN5-installed m5C on SCD1 mRNA is bound by YBX2, which prolongs SCD1 mRNA half-life and enhances its stability, thereby suppressing ferroptosis in renal tubular epithelial cells. Bisulfite sequencing, RNA immunoprecipitation, mRNA stability (half-life) assays, YBX2 binding assays, NSUN5 overexpression/knockdown, ferroptosis assays Advanced science Medium 42011916

Source papers

Stage 0 corpus · 44 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1998 Mammalian male and female germ cells express a germ cell-specific Y-Box protein, MSY2. Biology of reproduction 136 9780336
2005 Absence of the DNA-/RNA-binding protein MSY2 results in male and female infertility. Proceedings of the National Academy of Sciences of the United States of America 135 15824319
2001 Expression of MSY2 in mouse oocytes and preimplantation embryos. Biology of reproduction 99 11566752
2005 The DNA/RNA-binding protein MSY2 marks specific transcripts for cytoplasmic storage in mouse male germ cells. Proceedings of the National Academy of Sciences of the United States of America 80 15665108
2011 Absence of MSY2 in mouse oocytes perturbs oocyte growth and maturation, RNA stability, and the transcriptome. Biology of reproduction 79 21613634
2004 Transgenic RNAi-mediated reduction of MSY2 in mouse oocytes results in reduced fertility. Developmental biology 74 15031116
2019 LncRNA HOXA11-AS promotes OSCC progression by sponging miR-98-5p to upregulate YBX2 expression. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 69 31731187
2000 A sequence-specific RNA binding complex expressed in murine germ cells contains MSY2 and MSY4. Developmental biology 68 10772793
2001 MSY2 and MSY4 bind a conserved sequence in the 3' untranslated region of protamine 1 mRNA in vitro and in vivo. Molecular and cellular biology 67 11564883
2002 RNA-binding properties and translation repression in vitro by germ cell-specific MSY2 protein. Biology of reproduction 59 12297523
2006 In the absence of the mouse DNA/RNA-binding protein MSY2, messenger RNA instability leads to spermatogenic arrest. Biology of reproduction 58 17035640
2008 CDC2A (CDK1)-mediated phosphorylation of MSY2 triggers maternal mRNA degradation during mouse oocyte maturation. Developmental biology 49 18606161
2019 LINC00958 regulated miR-627-5p/YBX2 axis to facilitate cell proliferation and migration in oral squamous cell carcinoma. Cancer biology & therapy 43 31161900
2022 mRNA m5C inhibits adipogenesis and promotes myogenesis by respectively facilitating YBX2 and SMO mRNA export in ALYREF-m5C manner. Cellular and molecular life sciences : CMLS 36 35962235
2017 RNA Binding Protein Ybx2 Regulates RNA Stability During Cold-Induced Brown Fat Activation. Diabetes 36 28970281
2006 Deletion of the DNA/RNA-binding protein MSY2 leads to post-meiotic arrest. Molecular and cellular endocrinology 33 16413673
2021 RNA 5-methylcytosine regulates YBX2-dependent liquid-liquid phase separation. Fundamental research 31 38933916
2016 Role for PADI6 in securing the mRNA-MSY2 complex to the oocyte cytoplasmic lattices. Cell cycle (Georgetown, Tex.) 30 27929740
2008 Sequence alterations in the YBX2 gene are associated with male factor infertility. Fertility and sterility 29 18339382
1999 The mouse Y-box protein, MSY2, is associated with a kinase on non-polysomal mouse testicular mRNAs. Nucleic acids research 27 10076007
1999 Contrin, the human homologue of a germ-cell Y-box-binding protein: cloning, expression, and chromosomal localization. Journal of andrology 27 10100484
2016 Position-dependent interactions of Y-box protein 2 (YBX2) with mRNA enable mRNA storage in round spermatids by repressing mRNA translation and blocking translation-dependent mRNA decay. Molecular reproduction and development 24 26773323
2003 Requirement for RNA-binding activity of MSY2 for cytoplasmic localization and retention in mouse oocytes. Developmental biology 24 12648488
2024 Comprehensive pan-cancer analysis of YBX family reveals YBX2 as a potential biomarker in liver cancer. Frontiers in immunology 22 38698857
2009 MIWI-independent small RNAs (MSY-RNAs) bind to the RNA-binding protein, MSY2, in male germ cells. Proceedings of the National Academy of Sciences of the United States of America 21 19597149
2000 MSY2: a slowly evolving minisatellite on the human Y chromosome which provides a useful polymorphic marker in Chinese populations. Gene 20 10689184
2006 Expression of Y-box-binding protein dbpC/contrin, a potentially new cancer/testis antigen. British journal of cancer 18 16479255
2023 YBX2 modulates mRNA stability via interaction with YTHDF2 in endometrial cancer cells. Experimental cell research 17 37030331
2021 YBX2 and cancer testis antigen 45 contribute to stemness, chemoresistance and a high degree of malignancy in human endometrial cancer. Scientific reports 14 33602962
2008 Some single nucleotide polymorphisms of MSY2 gene might contribute to susceptibility to spermatogenic impairment in idiopathic infertile men. Urology 14 18372033
2019 Murine PAIP1 stimulates translation of spermiogenic mRNAs stored by YBX2 via its interaction with YBX2†. Biology of reproduction 13 30295753
2017 Genetic Polymorphisms in PRM1, PRM2, and YBX2 Genes are Associated with Male Factor Infertility. Genetic testing and molecular biomarkers 12 29227750
2008 MSY2 and polypyrimidine tract binding protein 2 stabilize mRNAs in the mammalian testis. International journal of andrology 12 18380784
2015 Evaluation of mRNA Contents of YBX2 and JHDM2A Genes on Testicular Tissues of Azoospermic Men with Different Classes of Spermatogenesis. Cell journal 11 25870841
2020 BNC1 Promotes Spermatogenesis by Regulating Transcription of Ybx2 and Papolb via Direct Binding to Their Promotor Elements. Reproductive sciences (Thousand Oaks, Calif.) 9 33211273
2016 Association of G/T(rs222859) polymorphism in Exon 1 of YBX2 gene with azoospermia, among Iranian infertile males. Andrologia 9 26804374
2015 Comparison of Protamine 1 to Protamine 2 mRNA Ratio and YBX2 gene mRNA Content in Testicular Tissue of Fertile and Azoospermic Men. International journal of fertility & sterility 9 26644857
2023 Phosphorylated YBX2 is stabilized to promote glycolysis in brown adipocytes. iScience 5 37860762
2025 Uremic toxin receptor NR1H3 contributes to hyperlipidemia- and chronic kidney disease-accelerated vascular inflammation, which is partially suppressed by novel YBX2 anti-ROS pathway. Redox biology 3 40505347
2021 Pseudogene RPL32P3 regulates the blood-tumor barrier permeability via the YBX2/HNF4G axis. Cell death discovery 3 34819492
2022 Ckb and Ybx2 interact with Ribc2 and are necessary for the ciliary beating of multi-cilia. Genes & genomics 2 36508087
2006 Expression and cellular localization of dbpC/Contrin in germ cell tumor cell lines. Biochimica et biophysica acta 1 16624424
2026 NSUN5 Attenuates Renal Injury and Ferroptosis in Hyperuricaemic Nephropathy Through YBX2-Dependent Stabilisation of SCD1 m5C Methylation. Advanced science (Weinheim, Baden-Wurttemberg, Germany) 0 42011916
2025 Identification of YBX2 and TSKS As STK33 Interacting Proteins in Testicular Germ Cells. Reproductive sciences (Thousand Oaks, Calif.) 0 39909973

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