Affinage

Showing DDX4 — VASA is a alias.

DDX4

Probable ATP-dependent RNA helicase DDX4 · UniProt Q9NQI0

Length
724 aa
Mass
79.3 kDa
Annotated
2026-06-09
100 papers in source corpus 22 papers cited in narrative 21 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 6/6 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

DDX4/VASA is a conserved ATP-dependent DEAD-box RNA helicase that organizes germline ribonucleoprotein granules and drives small-RNA-based transposon silencing essential for fertility (PMID:8026330, PMID:24910301). In Drosophila it nucleates a transient secondary-piRNA "Amplifier" complex containing two Piwi proteins, the Tudor protein Qin/Kumo, and antisense piRNA guides, with its helicase domain acting as an RNA clamp that anchors the complex to transposon transcripts and whose ATP-dependent RNP remodeling transfers sliced piRNA precursors between ping-pong partners; loss of this activity causes retrotransposon de-repression and sterility (PMID:17194939, PMID:24910301). Its localization and unwinding activities are genetically separable: posterior/nuage targeting depends on direct protein-protein interactions with Oskar and with the SPRY/SOCS-box protein Gustavus rather than RNA binding, whereas RNA-binding/helicase activity is required after localization for germ cell formation (PMID:8026330, PMID:8804312, PMID:12479811). Helicase activity also governs retention within phase-separated germ granules, and in C. elegans the Vasa homologs (RDE-12, GLH-1) directly bind Argonaute target mRNAs and promote secondary siRNA/piRNA amplification, transgenerational inheritance, and self/non-self discrimination during transcriptome surveillance (PMID:24684931, PMID:31506335, PMID:36085149, PMID:36070689). DDX4 carries conserved symmetrical dimethylarginine modifications installed by PRMT5/dPRMT5 that enable interactions with Tudor-domain proteins (Tdrd1, Tdrd6) and Piwi proteins (Mili, Miwi), a writer-substrate module essential for spermatogenesis (PMID:20080973, PMID:36634107). Independent of RNA silencing, Vasa has translation-independent and translation-dependent cell-cycle roles: it associates with condensin I components Barren and CAP-D2 to promote mitotic chromosome condensation, localizes to the mitotic spindle, and supports translation of cyclinB mRNA and cell-cycle/oncogene factors (PMID:21185189, PMID:21525076, PMID:28612512).

Mechanistic history

Synthesis pass · year-by-year structured walk · 19 steps
  1. 1994 High

    Established that Vasa's localization to germ plasm and its RNA helicase activity are genetically separable functions, defining a protein with distinct targeting and catalytic modules.

    Evidence In vitro RNA-binding/unwinding assays on mutant proteins plus in vivo localization of vas point mutants in Drosophila oocytes

    PMID:8026330

    Open questions at the time
    • Did not identify the RNA substrates unwound in vivo
    • Localization-mediating partners not yet identified
  2. 1996 High

    Identified Oskar as a direct Vasa partner, defining the initiating protein-protein interaction of polar granule assembly.

    Evidence Yeast two-hybrid and in vitro binding, immunoelectron microscopy, and oskar allele epistasis in Drosophila

    PMID:8804312

    Open questions at the time
    • Interaction interface not mapped at residue level
    • Does not explain how the complex recruits downstream granule components
  3. 2002 High

    Defined Gustavus as the dedicated localization factor anchoring Vasa posteriorly, separating Vasa transport from oskar RNA localization.

    Evidence Genetic screen, yeast two-hybrid/pulldown, Vasa deletion mapping, and immunolocalization in gus mutants

    PMID:12479811

    Open questions at the time
    • Mechanism by which GUS couples Vasa to the localization machinery unknown
  4. 2002 High

    Mapped a minimal germline-specific vasa promoter element and placed zygotic vasa expression downstream of maternal Nanos/Pumilio, defining transcriptional control of the gene.

    Evidence Reporter transgene deletion mapping, EMSA with ovarian extracts, and pole cell transplantation in nos/pum mutants

    PMID:11576171 PMID:11850184

    Open questions at the time
    • The ovarian factor binding the element not molecularly identified
  5. 2004 Medium

    Placed Vasa genetically in the germline retrotransposon-silencing pathway alongside Aubergine and Spindle-E, linking the helicase to RNA-based defense.

    Evidence Genetic epistasis with vasa/aub/spn-E alleles and transposon RNA quantification in Drosophila ovary

    PMID:17194939

    Open questions at the time
    • Direct biochemical role in silencing not resolved
    • Single lab; mechanism of Vasa action on transposon RNA undefined
  6. 2010 High

    Identified arginine dimethylation of Vasa by PRMT5 as a conserved PTM that licenses Tudor-domain and Piwi interactions, defining a reader/writer module for germ-granule assembly.

    Evidence Mass spectrometry of dimethylarginine, dPRMT5 mutant analysis, and Co-IP of MVH with Tdrd1/Tdrd6/Mili/Miwi from mouse testes

    PMID:20080973

    Open questions at the time
    • Functional consequence of each individual methyl-arginine not dissected
    • How methylation alters helicase activity unknown
  7. 2010 High

    Revealed a translation-independent role for Vasa in mitotic chromosome condensation via recruitment of condensin I subunits, expanding its function beyond RNA silencing.

    Evidence Co-IP of Vasa with Barr and CAP-D2, condensin localization in vas mutants, and epistasis with aub/spn-E in Drosophila

    PMID:21185189

    Open questions at the time
    • Whether helicase ATPase activity is needed for condensin recruitment unresolved
    • Direct vs. indirect Vasa-condensin contact not distinguished
  8. 2011 High

    Showed Vasa couples cell-cycle progression to translation by associating with the mitotic spindle and promoting cyclinB mRNA translation in sea urchin embryos.

    Evidence Immunofluorescence/live imaging, morpholino knockdown, polysome fractionation for cyclinB, and Cdk inhibitor experiments

    PMID:21525076

    Open questions at the time
    • Direct cyclinB mRNA binding by Vasa not shown
    • Mechanism of spindle association unknown
  9. 2012 Medium

    Demonstrated VASA can drive primordial germ cell differentiation and meiotic progression in human pluripotent stem cells, indicating a post-transcriptional role in human germline development.

    Evidence VASA/DAZL overexpression in hESCs/iPSCs with germ-cell and meiotic marker readouts

    PMID:22162380

    Open questions at the time
    • No direct molecular target of VASA identified in human cells
    • Gain-of-function only; endogenous requirement not tested
  10. 2014 High

    Defined the biochemical core of secondary piRNA biogenesis by reconstituting the Vasa-nucleated Amplifier complex and showing the helicase domain clamps RNA to enable ping-pong precursor transfer.

    Evidence Co-IP/MS, biochemical reconstitution, helicase-domain structure-function and ATPase mutants, and Drosophila fertility assays

    PMID:24910301

    Open questions at the time
    • High-resolution structure of the full Amplifier complex not determined
    • Timing/regulation of complex assembly in vivo unclear
  11. 2014 High

    Showed the C. elegans Vasa homolog RDE-12 bridges upstream and downstream Argonautes, ordering it within siRNA amplification.

    Evidence Co-IP of RDE-12 with WAGO-1, rde-12 mutant RNAi/siRNA analysis, and epistasis in C. elegans

    PMID:24684931

    Open questions at the time
    • Direct mRNA-binding step not biochemically isolated here
    • Helicase ATPase requirement not tested in this study
  12. 2019 High

    Established that helicase activity and RGG repeats govern Vasa/GLH-1 retention in phase-separated P granules, linking enzymatic activity to condensate physics and cytoplasmic compartmentalization.

    Evidence 28 endogenous CRISPR GLH-1 alleles (helicase-dead, RGG deletions), P-granule imaging, and interactome mass spectrometry in C. elegans

    PMID:31506335

    Open questions at the time
    • Mechanistic basis of ribosome/proteasome aversion unresolved
    • Quantitative model of helicase-driven retention lacking
  13. 2019 Medium

    Showed Vasa is required for germline stem cell maintenance and that its loss triggers irreversible Chk2-dependent oogenesis arrest, defining a checkpoint-coupled developmental requirement.

    Evidence Germarium-restricted Vasa loss, chk2 epistasis, and Vasa-restoration rescue in Drosophila

    PMID:31484689

    Open questions at the time
    • Molecular signal activating Chk2 upon Vasa loss unknown
    • Single model organism
  14. 2019 Medium

    Linked DDX4 knockdown to reduced primordial germ cell numbers and decreased aromatase (CYP19A1) expression in chicken gonads, implicating it in female germline gene regulation.

    Evidence Retroviral microRNA knockdown, PGC counts, and RT-qPCR of gonadal markers in chicken embryos

    PMID:30554591

    Open questions at the time
    • Direct vs. indirect link between DDX4 and aromatase unresolved
    • Knockdown specificity limited
  15. 2019 Low

    Reported surface exposure of the DDX4 C-terminus in an overexpression system, raising a non-canonical localization possibility.

    Evidence DDX4-DsRed2 epitope-tag immunocytochemistry and FACS of non-permeabilized HEK 293T cells

    PMID:31212843

    Open questions at the time
    • Single overexpression system in non-native cells; not confirmed endogenously
    • No mechanism for how the C-terminus reaches the surface
  16. 2021 Medium

    Identified LOTUS-domain proteins MIP-1/MIP-2 as direct GLH-1 anchors required for germ-granule coalescence and fertility, defining the scaffold that positions Vasa in P granules.

    Evidence Co-IP of MIP-1/MIP-2 with GLH-1, CRISPR double knockouts, and P-granule imaging in C. elegans

    PMID:34223818

    Open questions at the time
    • Binding interface and stoichiometry not resolved
    • Single lab
  17. 2022 High

    Showed GLH/Vasa helicase activity is required for piRNA silencing fidelity and self/non-self discrimination rather than piRNA production, distinguishing granule formation from small-RNA biogenesis.

    Evidence GLH helicase mutant analysis with genome-wide small RNA sequencing and germ-granule imaging in C. elegans

    PMID:36085149

    Open questions at the time
    • How condensate integrity enforces target discrimination mechanistically unclear
  18. 2022 High

    Demonstrated GLH family members compete to set Argonaute pathway specificity, with GLH-1 directly binding target mRNAs and its ATPase cycle regulating WAGO-1 binding for transgenerational inheritance.

    Evidence Genetic competition, RIP for direct mRNA binding, Co-IP with WAGO-1, ATPase-dead mutants, and small RNA sequencing in C. elegans

    PMID:36070689

    Open questions at the time
    • Structural basis of ATPase-coupled Argonaute handoff unresolved
    • Mammalian conservation of competition model untested
  19. 2023 High

    Confirmed the PRMT5-Vasa methylation module as essential for spermatogenesis in a second insect, mapping specific symmetrically dimethylated arginines required for sperm formation.

    Evidence CRISPR knockout of BmPrmt5 and BmVasa, mass spectrometry of methylation sites, and sperm morphology/RNA-seq in Bombyx mori

    PMID:36634107

    Open questions at the time
    • Downstream effectors of methylated Vasa in sperm formation not defined
    • Effect of methylation on helicase activity not tested

Open questions

Synthesis pass · forward-looking unresolved questions
  • How DDX4 helicase activity, arginine methylation, and condensate scaffolding are integrated to switch between piRNA silencing, mitotic condensation, and translational control in mammalian germ cells remains unresolved.
  • No mammalian structural model of the Amplifier or condensin-associated complexes
  • Direct mammalian mRNA targets of DDX4 not defined
  • Regulatory cross-talk between PTM and helicase cycle unmapped

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140657 ATP-dependent activity 4 GO:0003723 RNA binding 3 GO:0140098 catalytic activity, acting on RNA 3 GO:0016787 hydrolase activity 2 GO:0045182 translation regulator activity 2 GO:0060090 molecular adaptor activity 2
Localization
GO:0005694 chromosome 2 GO:0005815 microtubule organizing center 2 GO:0005829 cytosol 2 GO:0005634 nucleus 1
Pathway
R-HSA-8953854 Metabolism of RNA 4 R-HSA-1266738 Developmental Biology 3 R-HSA-1474165 Reproduction 3 R-HSA-1640170 Cell Cycle 3
Partners
BARRCAP-D2GUSMIP-1OSKPRMT5TDRD1WAGO-1
Complex memberships
condensin Igerm/P granule (nuage/polar granule)piRNA Amplifier complex

Evidence

Reading pass · 21 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1994 Drosophila Vasa protein localizes to perinuclear nuage and pole plasm; pole plasm localization requires protein-protein interactions (not RNA-binding/helicase activity), while RNA-binding activity is required post-localization for germ cell formation. Mutations in RNA-binding and unwinding domains abolish helicase activity but still permit pole plasm localization, demonstrating separable functions. In vitro RNA-binding and unwinding assays on purified mutant Vasa proteins; genetic analysis of vas point mutations; immunolocalization in Drosophila oocytes Development High 8026330
1996 Oskar protein interacts directly with Vasa in yeast two-hybrid and in vitro binding assays; mutations in Oskar that abolish pole plasm formation in vivo also disrupt the Oskar-Vasa interaction. Oskar and Vasa are both components of polar granules, and the Oskar-Vasa interaction represents an essential initial step in polar granule assembly. Yeast two-hybrid assay, in vitro binding assay, immunoelectron microscopy of polar granules, genetic epistasis with oskar alleles Genes & development High 8804312
2002 Gustavus (GUS), a SPRY-domain/SOCS-box protein, directly interacts with Vasa and is required for posterior localization of Vasa in Drosophila oocytes. Deletion of the GUS-binding segment of Vasa blocks posterior localization. GUS is not required for oskar RNA localization, placing GUS specifically in the Vasa localization pathway. Genetic screen, yeast two-hybrid/pulldown interaction assay, immunolocalization of Vasa in gus mutants, deletion mapping of Vasa Developmental cell High 12479811
2002 A 40-bp regulatory element within the vasa promoter is necessary and sufficient for germline-specific expression during both oogenesis and embryogenesis in Drosophila; this region interacts specifically with ovarian protein(s). Maternal Nanos and Pumilio are required autonomously in pole cells for normal zygotic vasa expression during embryogenesis. EGFP-Vasa reporter transgene analysis; promoter deletion mapping; electrophoretic mobility shift assay with ovarian extracts; pole cell transplantation; analysis of nos and pum mutants Mechanisms of development High 11576171 11850184
2004 Vasa RNA helicase, together with Aubergine (Piwi family protein), is involved in retrotransposon silencing (I element, Het-A, copia) in the Drosophila female germline. Mutations in vasa cause accumulation of retrotransposon transcripts similar to spn-E and aub mutants, placing Vasa in the same silencing pathway as these RNAi factors. Genetic epistasis using vasa, aub, and spn-E loss-of-function mutations; quantitative RT-PCR/Northern blot for retrotransposon RNA levels; immunostaining of perinuclear ribonucleoprotein particles RNA biology Medium 17194939
2010 Vasa protein contains symmetrical and asymmetrical dimethylarginine modifications; dPRMT5 (arginine methyltransferase 5) is required in vivo for symmetrical dimethylation of Vasa in Drosophila. Methylated mouse Vasa homolog (MVH) associates with Tudor domain-containing proteins Tdrd1 and Tdrd6, and with Piwi proteins Mili and Miwi, establishing arginine methylation as a conserved PTM enabling Tudor-domain protein interactions. Mass spectrometry identification of dimethylarginine residues; genetic analysis of dPRMT5 mutants; co-immunoprecipitation of MVH with Tdrd1, Tdrd6, Mili, Miwi from mouse testes The Journal of biological chemistry High 20080973
2010 Vasa has a translation-independent function in mitotic chromosome condensation in Drosophila germline cells. During mitosis, Vasa facilitates chromosomal localization of condensin I components Barren (Barr) and CAP-D2 (but not CAP-D3/condensin II). Vasa physically associates with Barr and CAP-D2. Formation of perichromosomal Vasa bodies during mitosis requires piRNA pathway components Aubergine and Spindle-E. Co-immunoprecipitation of Vasa with Barr and CAP-D2; immunofluorescence of condensin localization in vas mutants; genetic epistasis with aub and spn-E mutants Current biology High 21185189
2011 In sea urchin embryos, Vasa is an ATP-dependent RNA helicase present in all blastomeres whose abundance oscillates with the cell cycle. Vasa associates with the mitotic spindle and separating chromatids at metaphase. Inhibition of Vasa protein synthesis arrests cells at M-phase and delays cell cycle progression. Cdk activity is required for proper Vasa localization, and Vasa is required for efficient translation of cyclinB mRNA. Immunofluorescence and live imaging; morpholino knockdown; cell cycle synchronization; polysome fractionation/translation assay for cyclinB mRNA; Cdk inhibitor experiments Development High 21525076
2012 Overexpression of VASA (DDX4) and/or DAZL in human embryonic stem cells (hESCs) and iPSCs promotes differentiation to primordial germ cells and enhances progression through meiosis in vitro, demonstrating that VASA can function as a translational/post-transcriptional factor driving meiotic progression in human germ cells. Overexpression in hESCs and iPSCs; immunofluorescence and flow cytometry for germ cell and meiotic markers; comparison of VASA vs. DAZL vs. combined overexpression Stem cells Medium 22162380
2014 Vasa nucleates a transient 'Amplifier complex' for secondary piRNA biogenesis in insect cells. The complex contains Vasa, two Piwi proteins (participating in the ping-pong cycle), Tudor protein Qin/Kumo, and antisense piRNA guides. Vasa's helicase domain acts as an RNA clamp anchoring the complex to transposon transcripts. ATP-dependent RNP remodeling by Vasa facilitates transfer of 5'-sliced piRNA precursors between ping-pong partners; loss of this activity causes sterility in Drosophila. Co-immunoprecipitation and mass spectrometry identification of Amplifier complex; biochemical reconstitution; structure-function analysis of Vasa helicase domain; ATPase mutant analysis; in vivo fertility assay in Drosophila Cell High 24910301
2014 In C. elegans, the Vasa homolog RDE-12 physically associates with the Argonaute WAGO-1, colocalizes with WAGO-1 in germline P granules and somatic foci, and is required for small RNA amplification (secondary siRNA production) and RNAi. RDE-12 is first recruited to target mRNA by upstream Argonautes (RDE-1, ERGO-1), then promotes WAGO-1 loading. Co-immunoprecipitation of RDE-12 with WAGO-1; genetic analysis of rde-12 mutants for RNAi deficiency and siRNA levels; immunolocalization; genetic epistasis ordering RDE-12 in the RNAi pathway Current biology High 24684931
2019 In C. elegans, GLH-1/Vasa helicase activity is required for its retention in P granules; without helicase activity GLH-1 dissociates from P granules. Glycine-rich (RGG) repeats are required for P-granule wetting-like interactions at the nuclear periphery. Mass spectrometry identifies GLH-1 as part of a piRNA-amplifying complex and reveals association with PCI complexes (proteasome lid, COP9, eIF3), while GLH-1 shows an aversion to assembled ribosomes and the 26S proteasome, suggesting P granules compartmentalize the cytoplasm to shield mRNAs from translation and proteins from degradation. CRISPR/Cas9 generation of 28 endogenous GLH-1 alleles including helicase-dead and RGG deletions; immunofluorescence of P granule localization; mass spectrometry of GLH-1-associated proteins Genetics High 31506335
2019 In Drosophila, absence of Vasa during the germarial stage causes Chk2-dependent oogenesis arrest. Once Chk2 is activated in the germarium by loss of Vasa, the arrest cannot be rescued by subsequent restoration of Vasa expression, and Vasa is required for germline stem cell homeostasis/maintenance. Conditional/germarium-restricted loss of Vasa expression; epistasis with chk2 mutants; rescue experiments with Vasa restoration; immunostaining of germline markers Genetics Medium 31484689
2022 In C. elegans, GLH/VASA helicase mutants form defective perinuclear condensates containing PIWI and small RNA cofactors. These mutants produce largely normal piRNA levels but are defective in triggering piRNA silencing, and hundreds of endogenous genes are aberrantly silenced by piRNAs, demonstrating that perinuclear germ granule formation by GLH/Vasa is required for the fidelity (self vs. non-self discrimination) of piRNA-based transcriptome surveillance. GLH helicase mutant analysis; small RNA sequencing (piRNA levels and targets); immunofluorescence of germ granule components; genetic comparison with other perinuclear germ granule mutants Nature communications High 36085149
2022 In C. elegans, GLH proteins (Vasa homologs) compete with each other to control Argonaute pathway specificity; GLH-1 directly binds Argonaute target mRNAs and promotes amplification of small RNAs required for transgenerational inheritance. The ATPase cycle of GLH-1 regulates its direct binding to the Argonaute WAGO-1. Genetic competition analysis of GLH family members; RIP (RNA immunoprecipitation) to show direct mRNA binding; small RNA sequencing; Co-IP of GLH-1 with WAGO-1; ATPase-dead GLH-1 mutants Cell reports High 36070689
2008 Overexpression of VASA in human ovarian cancer cells (SKOV-3) causes significant downregulation of 14-3-3sigma (identified by 2D proteomics/mass spectrometry), leading to abrogation of the DNA damage-induced G2 checkpoint. Stable VASA overexpression in SKOV-3 cells; 2D gel proteomics and mass spectrometry for protein expression profiling; G2 checkpoint assay after DNA damage Gynecologic oncology Medium 18805576
2017 In human blood-derived cancer cell lines (myeloma IM-9, leukemia THP-1), DDX4 protein localizes to the mitotic spindle. Knockout of DDX4 in IM-9 cells compromises cell proliferation and migration, and downregulates cell cycle/oncogene factors CyclinB and transcription factor E2F1. Immunofluorescence of DDX4 at mitotic spindle; CRISPR/siRNA knockdown/knockout; proliferation and migration assays; Western blot for CyclinB and E2F1 Cancer science Medium 28612512
2023 In Bombyx mori, BmPrmt5 (type II arginine methyltransferase) catalyzes symmetrical dimethylation of BmVasa at R35, R54, and R56 (identified by mass spectrometry). Loss of BmPrmt5 abolishes Vasa arginine methylation and causes the same male and female sterility phenotype as BmVasa loss, demonstrating that the Prmt5-Vasa methylation module is essential for spermatogenesis and apyrene/eupyrene sperm formation in this species. CRISPR/Cas9 knockout of BmPrmt5 and BmVasa; mass spectrometry identification of dimethylarginine residues in Vasa; immunofluorescence of sperm morphology; RNA-seq for downstream gene expression PLoS genetics High 36634107
2019 In chicken embryos, DDX4 (Vasa) knockdown via retroviral microRNA vectors decreases the number of primordial germ cells in both male and female gonads. In female DDX4-knockdown gonads, expression of aromatase (CYP19A1), essential for ovary development, is significantly decreased, linking DDX4 to regulation of aromatase expression in the female germline. Retroviral microRNA-mediated knockdown; immunofluorescence counting of PGCs; RT-qPCR for DMRT1, SOX9, CYP19A1, and FOXL2 Reproduction, fertility, and development Medium 30554591
2021 In C. elegans, novel LOTUS-domain proteins MIP-1 and MIP-2 directly bind and anchor the Vasa homolog GLH-1 within P granules; double loss of MIP-1 and MIP-2 prevents coalescence of GLH-1, MEG-3, and PGL proteins. MIP proteins are required for germline stem cell self-renewal, meiotic progression, and gamete differentiation. Co-immunoprecipitation of MIP-1/MIP-2 with GLH-1; CRISPR/Cas9 double knockouts; immunofluorescence of P granule components; phenotypic analysis of fertility and meiosis eLife Medium 34223818
2019 The C-terminus of DDX4 can be expressed on the cell surface of HEK 293T cells when the full-length protein is expressed, as demonstrated by immunocytochemistry and FACS of non-permeabilized cells using C-terminal epitope tags, despite DDX4 lacking a conventional membrane-targeting or secretory sequence. Expression of DDX4-DsRed2 fusion with C- and N-terminal epitope tags; immunocytochemistry and FACS of non-permeabilized HEK 293T cells; RT-PCR confirmation of DDX4 expression in sorted cells Cells Low 31212843

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2000 The human VASA gene is specifically expressed in the germ cell lineage. Proceedings of the National Academy of Sciences of the United States of America 467 10920202
2000 Expression and intracellular localization of mouse Vasa-homologue protein during germ cell development. Mechanisms of development 454 10781947
2000 Zebrafish vasa RNA but not its protein is a component of the germ plasm and segregates asymmetrically before germline specification. The Journal of cell biology 316 10811828
2000 Isolation of chicken vasa homolog gene and tracing the origin of primordial germ cells. Development (Cambridge, England) 309 10821771
1994 Localization of vasa protein to the Drosophila pole plasm is independent of its RNA-binding and helicase activities. Development (Cambridge, England) 295 8026330
2000 The function and regulation of vasa-like genes in germ-cell development. Genome biology 265 11178242
1999 Expression of vasa(vas)-related genes in germline cells and totipotent somatic stem cells of planarians. Developmental biology 225 9918696
2014 RNA clamping by Vasa assembles a piRNA amplifier complex on transposon transcripts. Cell 207 24910301
1997 A vasa-like gene in zebrafish identifies putative primordial germ cells. Mechanisms of development 205 9376327
2007 The dynamic vasa vasorum. Cardiovascular research 184 17631284
1996 Oskar protein interaction with Vasa represents an essential step in polar granule assembly. Genes & development 179 8804312
2022 High-throughput total RNA sequencing in single cells using VASA-seq. Nature biotechnology 168 35760914
2001 Universal occurrence of the vasa-related genes among metazoans and their germline expression in Hydra. Development genes and evolution 163 11466525
2001 Vasa homolog genes in mammalian germ cell development. Cell structure and function 154 11565805
2000 Differential expression of vasa homologue gene in the germ cells during oogenesis and spermatogenesis in a teleost fish, tilapia, Oreochromis niloticus. Mechanisms of development 144 11091081
1999 Characterization of zebrafish primordial germ cells: morphology and early distribution of vasa RNA. Developmental dynamics : an official publication of the American Association of Anatomists 137 10536055
2012 Divergent RNA-binding proteins, DAZL and VASA, induce meiotic progression in human germ cells derived in vitro. Stem cells (Dayton, Ohio) 132 22162380
2005 Differential expression of vasa RNA and protein during spermatogenesis and oogenesis in the gibel carp (Carassius auratus gibelio), a bisexually and gynogenetically reproducing vertebrate. Developmental dynamics : an official publication of the American Association of Anatomists 119 15880437
2014 Zebrafish vasa is required for germ-cell differentiation and maintenance. Molecular reproduction and development 114 25257909
2005 vasa and nanos expression patterns in a sea anemone and the evolution of bilaterian germ cell specification mechanisms. Evolution & development 109 15876193
2013 The DEAD-box helicase Vasa: evidence for a multiplicity of functions in RNA processes and developmental biology. Biochimica et biophysica acta 105 23587717
2015 Vasa vasorum in atherosclerosis and clinical significance. International journal of molecular sciences 103 26006236
2004 The RNA interference proteins and vasa locus are involved in the silencing of retrotransposons in the female germline of Drosophila melanogaster. RNA biology 102 17194939
2000 Cloning and characterization of a vasa-like gene in rainbow trout and its expression in the germ cell lineage. Molecular reproduction and development 95 10694742
2009 In vivo RNA interference in oyster--vasa silencing inhibits germ cell development. The FEBS journal 83 19476495
1999 A mutation in the zebrafish maternal-effect gene nebel affects furrow formation and vasa RNA localization. Current biology : CB 83 10607587
2006 Dynamic redistribution of vasa homolog and exclusion of somatic cell determinants during germ cell specification in Ciona intestinalis. Development (Cambridge, England) 82 16794033
2002 Identification of a transcriptional regulatory region for germline-specific expression of vasa gene in Drosophila melanogaster. Mechanisms of development 79 11850184
2010 Arginine methylation of vasa protein is conserved across phyla. The Journal of biological chemistry 71 20080973
2002 VASA localization requires the SPRY-domain and SOCS-box containing protein, GUSTAVUS. Developmental cell 67 12479811
2006 Germ cell development in the Honeybee (Apis mellifera); vasa and nanos expression. BMC developmental biology 64 16503992
2017 Ascending Vasa Recta Are Angiopoietin/Tie2-Dependent Lymphatic-Like Vessels. Journal of the American Society of Nephrology : JASN 63 29237738
2022 On vasa vasorum: A history of advances in understanding the vessels of vessels. Science advances 61 35442731
2010 A role for vasa in regulating mitotic chromosome condensation in Drosophila. Current biology : CB 60 21185189
2009 Vasa and the germ line lineage in a colonial urochordate. Developmental biology 58 19406116
2018 Medial Hypoxia and Adventitial Vasa Vasorum Remodeling in Human Ascending Aortic Aneurysm. Frontiers in cardiovascular medicine 55 30276199
1999 Isolation and characterization of a Bombyx vasa-like gene. Development genes and evolution 55 11252184
2011 DDX4 (VASA) is conserved in germ cell development in marsupials and monotremes. Biology of reproduction 54 21653890
2018 Initial characterisation of adult human ovarian cell populations isolated by DDX4 expression and aldehyde dehydrogenase activity. Scientific reports 53 29725036
2011 The DEAD-box RNA helicase Vasa functions in embryonic mitotic progression in the sea urchin. Development (Cambridge, England) 50 21525076
2014 Use of DEAD-box polypeptide-4 (Ddx4) gene promoter-driven fluorescent reporter mice to identify mitotically active germ cells in post-natal mouse ovaries. Molecular human reproduction 47 25147160
2009 Review and Hypothesis: Vulnerable plaque formation from obstruction of Vasa vasorum by homocysteinylated and oxidized lipoprotein aggregates complexed with microbial remnants and LDL autoantibodies. Annals of clinical and laboratory science 47 19201735
2014 Role of the vasa vasorum and vascular resident stem cells in atherosclerosis. BioMed research international 46 24724094
2011 The multiple hats of Vasa: its functions in the germline and in cell cycle progression. Molecular reproduction and development 46 21823188
2019 Germline Maintenance Through the Multifaceted Activities of GLH/Vasa in Caenorhabditis elegans P Granules. Genetics 42 31506335
2016 FACS-sorted putative oogonial stem cells from the ovary are neither DDX4-positive nor germ cells. Scientific reports 42 27301892
2009 Expression of Dazl and Vasa in turtle embryos and ovaries: evidence for inductive specification of germ cells. Evolution & development 42 19754709
2020 Adventitial fibroblast-derived vascular endothelial growth factor promotes vasa vasorum-associated neointima formation and macrophage recruitment. Cardiovascular research 39 31241138
2014 vasa and piwi are required for mitotic integrity in early embryogenesis in the spider Parasteatoda tepidariorum. Developmental biology 39 25257304
2015 VASA (DDX4) is a Putative Marker for Spermatogonia, Spermatocytes and Round Spermatids in Stallions. Reproduction in domestic animals = Zuchthygiene 37 26482643
2017 Classification and Functional Characterization of Vasa Vasorum-Associated Perivascular Progenitor Cells in Human Aorta. Stem cell reports 35 28552602
2008 Germ cell specific protein VASA is over-expressed in epithelial ovarian cancer and disrupts DNA damage-induced G2 checkpoint. Gynecologic oncology 33 18805576
2007 Vasa expression in a colonial ascidian, Botrylloides violaceus. Evolution & development 33 17371399
2010 Cloning and differential expression pattern of vasa in the developing and recrudescing gonads of catfish, Clarias gariepinus. Comparative biochemistry and physiology. Part A, Molecular & integrative physiology 32 20462517
2007 Differential expression of VASA gene in ejaculated spermatozoa from normozoospermic men and patients with oligozoospermia. Asian journal of andrology 32 17486274
2012 Expression of vasa and nanos3 during primordial germ cell formation and migration in Atlantic cod (Gadus morhua L.). Theriogenology 31 22898013
2005 Expression pattern of Bombyx vasa-like (BmVLG) protein and its implications in germ cell development. Development genes and evolution 30 16261344
2021 Building RNA-protein germ granules: insights from the multifaceted functions of DEAD-box helicase Vasa/Ddx4 in germline development. Cellular and molecular life sciences : CMLS 29 34921622
2014 The Vasa Homolog RDE-12 engages target mRNA and multiple argonaute proteins to promote RNAi in C. elegans. Current biology : CB 29 24684931
2012 Germ line specific expression of a vasa homologue gene in turbot (Scophthalmus maximus): evidence for vasa localization at cleavage furrows in euteleostei. Molecular reproduction and development 29 23124920
2018 Local adventitial anti-angiogenic gene therapy reduces growth of vasa-vasorum and in-stent restenosis in WHHL rabbits. Journal of molecular and cellular cardiology 28 30003882
2017 Stage-specific expression of DDX4 and c-kit at different developmental stages of the porcine testis. Animal reproduction science 27 29338902
2016 An unregulated regulator: Vasa expression in the development of somatic cells and in tumorigenesis. Developmental biology 27 27179696
2014 DDX4 (DEAD box polypeptide 4) colocalizes with cancer stem cell marker CD133 in ovarian cancers. Biochemical and biophysical research communications 27 24727449
2005 Characterization of pig vasa homolog gene and specific expression in germ cell lineage. Molecular reproduction and development 26 16094672
2012 Vasa-Like DEAD-Box RNA Helicases of Schistosoma mansoni. PLoS neglected tropical diseases 25 22720105
1999 On the regulation of tone in vasa vasorum. Cardiovascular research 25 10325971
2023 The Prmt5-Vasa module is essential for spermatogenesis in Bombyx mori. PLoS genetics 24 36634107
2013 Expression of MAEL in nuage and non-nuage compartments of rat spermatogenic cells and colocalization with DDX4, DDX25 and MIWI. Histochemistry and cell biology 24 23412502
2021 Novel LOTUS-domain proteins are organizational hubs that recruit C. elegans Vasa to germ granules. eLife 23 34223818
2019 Similar Population of CD133+ and DDX4+ VSEL-Like Stem Cells Sorted from Human Embryonic Stem Cell, Ovarian, and Ovarian Cancer Ascites Cell Cultures: The Real Embryonic Stem Cells? Cells 23 31336813
1996 [The vasa vasorum of the arteries]. Journal des maladies vasculaires 23 8984146
2019 Germ Cell Lineage Homeostasis in Drosophila Requires the Vasa RNA Helicase. Genetics 22 31484689
2013 Cloning, expression promoter analysis of vasa gene in Japanese flounder (Paralichthys olivaceus). Comparative biochemistry and physiology. Part B, Biochemistry & molecular biology 22 23796850
2012 Germ cell specific expression of Vasa in rare minnow, Gobiocypris rarus. Comparative biochemistry and physiology. Part A, Molecular & integrative physiology 22 22357168
2018 Expression of vasa, piwi, and nanos during gametogenesis in Typosyllis antoni (Annelida, Syllidae). Evolution & development 21 30094969
2014 Atherosclerosis and atheroma plaque rupture: normal anatomy of vasa vasorum and their role associated with atherosclerosis. TheScientificWorldJournal 21 24790560
2022 A family of C. elegans VASA homologs control Argonaute pathway specificity and promote transgenerational silencing. Cell reports 20 36070689
2022 GLH/VASA helicases promote germ granule formation to ensure the fidelity of piRNA-mediated transcriptome surveillance. Nature communications 20 36085149
2014 Piwi regulates Vasa accumulation during embryogenesis in the sea urchin. Developmental dynamics : an official publication of the American Association of Anatomists 20 24218044
2014 Light and electron microscopic analyses of Vasa expression in adult germ cells of the fish medaka. Gene 20 24814190
2012 Nestin and WT1 expression in small-sized vasa vasorum from human normal arteries. Histology and histopathology 20 22806906
2009 An evolutionary transition of Vasa regulation in echinoderms. Evolution & development 20 19754712
2007 A vasa-like gene in the giant freshwater prawn, Macrobrachium rosenbergii. Molecular reproduction and development 20 17186538
2017 Multiple Functions of the DEAD-Box Helicase Vasa in Drosophila Oogenesis. Results and problems in cell differentiation 19 28779316
2011 Circular DNA intermediate in the duplication of Nile tilapia vasa genes. PloS one 19 22216289
2003 Hormonal regulation of vasa-like messenger RNA expression in the ovary of the marine teleost Sparus aurata. Biology of reproduction 19 14613903
2006 Identification, localization, and sequencing of fetal bovine VASA homolog. Animal reproduction science 18 17150314
2002 Vasa expression and germ-cell specification in the spider mite Tetranychus urticae. Development genes and evolution 18 12536324
2019 Extracellular Localisation of the C-Terminus of DDX4 Confirmed by Immunocytochemistry and Fluorescence-Activated Cell Sorting. Cells 17 31212843
2017 Germline factor DDX4 functions in blood-derived cancer cell phenotypes. Cancer science 17 28612512
2006 Cloning and pattern of expression of the shiro-uo vasa gene during embryogenesis and its roles in PGC development. The International journal of developmental biology 17 16892175
2021 Ginsenoside Rb1 Enhances Plaque Stability and Inhibits Adventitial Vasa Vasorum via the Modulation of miR-33 and PEDF. Frontiers in cardiovascular medicine 16 34124194
2018 Expression pattern of nanos, piwil, dnd, vasa and pum genes during ontogenic development in Nile tilapia Oreochromis niloticus. Gene 16 30503393
2017 VASA expression suggests shared germ line dynamics in bivalve molluscs. Histochemistry and cell biology 16 28386635
2019 Ddx4+ Oogonial Stem Cells in Postmenopausal Women's Ovaries: A Controversial, Undefined Role. Cells 15 31261822
2016 Discrimination and characterization of Sertoli cell-only syndrome in non-obstructive azoospermia using cell-free seminal DDX4. Reproductive biomedicine online 15 27211570
2019 Knockdown of DEAD-box helicase 4 (DDX4) decreases the number of germ cells in male and female chicken embryonic gonads. Reproduction, fertility, and development 14 30554591
2019 Characterization and expression of a vasa homolog in the gonads and primordial germ cells of the striped catfish (Pangasianodon hypophthalmus). Theriogenology 14 30947076
2001 Maternal Nanos and Pumilio regulate zygotic vasa expression autonomously in the germ-line progenitors of Drosophila melanogaster embryos. Development, growth & differentiation 14 11576171

Missed literature

Know a paper Affinage missed for DDX4? Flag it for the maintainers and the community.

No submissions yet.