Affinage

PIWIL1

Piwi-like protein 1 · UniProt Q96J94

Length
861 aa
Mass
98.6 kDa
Annotated
2026-06-10
100 papers in source corpus 21 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

PIWIL1 (HIWI) is a PIWI-clade Argonaute that, in the germline, loads piRNAs to form piRNA-induced silencing complexes (piRISCs) that direct transposon silencing during spermatogenesis and oogenesis (PMID:28676534, PMID:37644029, PMID:25818294). As a piRNA-guided endoribonuclease, its weak intrinsic slicing activity is potentiated into efficient target cleavage by the accessory factor GTSF1, and this catalytic activity is essential for functional sperm production (PMID:35772669). Unlike AGO-clade Argonautes, PIWIL1 cleaves targets that are only partially base-paired to its piRNA guide, tolerating mismatches at any position and dispensing with seed pairing (PMID:37344600); cryo-EM of mammalian PIWI piRISCs attributes this mismatch tolerance to a wider nucleic-acid channel and a vertebrate-specific lysine that distorts the piRNA seed and relaxes the seed gate upon target engagement (PMID:38658622). PIWIL1 carries symmetrical dimethyl-arginine marks installed by PRMT5 that mediate methylation-dependent Tudor-domain interactions (PMID:20360382), whereas TDRD2 recognizes an unmethylated arginine-rich PIWIL1 sequence in a methylation-independent manner required for piRNA 3'-end trimming (PMID:29118143). In somatic cancers PIWIL1 acts largely independently of piRNAs: in gastric cancer its oncogenic function persists when piRNA binding is abolished and operates through interaction with the UPF1-dependent nonsense-mediated mRNA decay machinery (PMID:32848063); in glioma stem cells it sustains self-renewal by regulating the mRNA stability of tumor suppressors BTG2, FBXW7, and CDKN1B (PMID:33406417); it binds and stabilizes Stathmin1 by blocking its RLIM-mediated ubiquitination and CaMKII-mediated phosphorylation, thereby destabilizing microtubules and promoting migration (PMID:26317901); and it drives tumorigenesis by inducing global DNA hypermethylation that silences cyclin-dependent kinase inhibitors (PMID:22438986). PIWIL1 expression confers stem-like and oncogenic phenotypes across multiple tumor types, including sarcoma, endometrial, and hepatocellular carcinoma, where it also reshapes tumor metabolism and the immune microenvironment (PMID:22438986, PMID:33633112, PMID:26506848).

Mechanistic history

Synthesis pass · year-by-year structured walk · 14 steps
  1. 2002 Medium

    Established the first cellular context for human PIWIL1 — germline-restricted expression and a growth-suppressive/pro-apoptotic activity when ectopically expressed — framing it as a germline developmental regulator.

    Evidence RT-PCR, immunohistochemistry, and overexpression with apoptosis assays in testis and KG1 leukemia cells

    PMID:11154219 PMID:12037681

    Open questions at the time
    • No molecular mechanism for the apoptotic phenotype
    • piRNA/effector activity not yet linked
  2. 2010 Medium

    Defined a post-translational regulatory layer: PRMT5-installed symmetrical dimethyl-arginines on PIWI proteins create docking sites for Tudor-domain partners.

    Evidence Biochemical methyltransferase assays and co-immunoprecipitation synthesized in review

    PMID:20360382

    Open questions at the time
    • Specific Tudor partners of human PIWIL1 not individually mapped here
    • Functional consequence of each interaction unresolved
  3. 2017 High

    Showed an exception to the methylation paradigm — TDRD2 recognizes unmethylated PIWIL1 — and linked this interaction to piRNA biogenesis via 3'-end trimming.

    Evidence Crystal structure, in vitro binding/mutagenesis, and in vitro trimming assay

    PMID:29118143

    Open questions at the time
    • In vivo requirement for trimming in mammals not tested here
    • Trimming nuclease identity not addressed
  4. 2022 High

    Resolved how PIWIL1 becomes catalytically competent — GTSF1 converts its weak intrinsic slicer activity into efficient cleavage — and tied this catalysis to fertility.

    Evidence In vitro cleavage reconstitution with GTSF1 plus mouse knockout fertility phenotypes

    PMID:35772669

    Open questions at the time
    • Structural basis of GTSF1 potentiation not defined here
    • Somatic relevance of catalysis untested
  5. 2023 High

    Explained why PIWI silencing is broad — PIWIL1 cleaves partially paired targets without requiring seed pairing, distinguishing it mechanistically from AGO Argonautes.

    Evidence In vitro binding and cleavage with recombinant mouse MIWI and sponge PIWI, mutational pairing analysis

    PMID:37344600

    Open questions at the time
    • Target repertoire in vivo not enumerated
    • Off-target consequences of mismatch tolerance unexamined
  6. 2024 High

    Provided the structural basis for mismatch tolerance — a wider channel, prearranged extended seed, and a vertebrate-specific lysine that relaxes the seed gate on target binding.

    Evidence Cryo-EM of MILI/HILI piRISCs with functional mutagenesis and in vitro assays

    PMID:38658622

    Open questions at the time
    • Direct structures of PIWIL1 itself not solved here
    • Catalytic step structures not captured
  7. 2015 Medium

    Connected human germline PIWIL1 to its presumed ancestral function — transposon (LINE-1) silencing — across testis and oocytes of multiple species.

    Evidence Immunolocalization, laser-microdissection RT-PCR, proteomics, and small RNA sequencing in human and bovine gonads

    PMID:25818294 PMID:28676534

    Open questions at the time
    • Causality between PIWIL1 and LINE-1 silencing correlative in human tissue
    • piRNA target sites not mapped
  8. 2023 High

    Demonstrated an essential, non-redundant reproductive requirement for PIWIL1 in a mammalian model with full PIWI-family coverage, including partial compensation for PIWIL3.

    Evidence CRISPR knockout of all four PIWI genes in golden hamster with fertility, localization, and RNA-seq analysis

    PMID:37644029

    Open questions at the time
    • Human germline phenotype inferred, not directly tested
    • Molecular targets driving sterility not fully defined
  9. 2012 High

    Opened the somatic-oncogenic chapter, linking PIWIL1 to tumorigenesis through global DNA hypermethylation and CDKI silencing reversible with DNMT inhibitors.

    Evidence In vitro differentiation, transgenic mouse sarcoma model, inducible knockdown, and DNA methylation analysis

    PMID:22438986

    Open questions at the time
    • Direct molecular link between PIWIL1 and DNMT machinery not established
    • piRNA dependence not tested here
  10. 2015 Medium

    Identified a piRNA-independent protein-stabilization mechanism — PIWIL1 binds Stathmin1, blocking its ubiquitination and phosphorylation to destabilize microtubules and drive migration.

    Evidence Co-IP, ubiquitination/phosphorylation assays, microtubule polymerization, and migration assays

    PMID:26317901

    Open questions at the time
    • Single lab without structural definition of the interaction
    • Tissue generality of the mechanism unknown
  11. 2020 High

    Formally separated PIWIL1's somatic oncogenic activity from its piRNA-binding function, implicating the UPF1/NMD pathway in gastric cancer.

    Evidence PIWIL1 knockout, RNA deep sequencing, piRNA-binding mutant rescue, and Co-IP with NMD factors

    PMID:32848063

    Open questions at the time
    • How PIWIL1 modulates NMD activity mechanistically unresolved
    • mRNA targets stabilized/destabilized not fully cataloged
  12. 2021 Medium

    Extended the somatic mRNA-regulatory model to glioma stem cells, where PIWIL1 controls stability of BTG2, FBXW7, and CDKN1B to sustain self-renewal.

    Evidence shRNA knockdown, RNA-seq, mRNA stability assays, and xenograft models

    PMID:33406417

    Open questions at the time
    • Direct PIWIL1–mRNA binding not demonstrated for each target
    • Single lab
  13. 2021 Medium

    Broadened PIWIL1's tumor roles to metabolic reprogramming and immune evasion in hepatocellular carcinoma via C3-mediated MDSC recruitment.

    Evidence Overexpression/knockdown, metabolic flux analysis, MDSC co-culture, C3 neutralization, and p38 MAPK inhibition

    PMID:33633112

    Open questions at the time
    • Mechanism by which PIWIL1 induces C3 secretion unknown
    • Single lab
  14. 2019 Medium

    Documented a residual piRNA-loaded, perinuclear nuage-like PIWIL1 complex in somatic cancer cells engaging carcinogenesis-related target mRNAs.

    Evidence Immunofluorescence, RNA immunoprecipitation, and RNA sequencing in colorectal cancer cells

    PMID:31694219 PMID:33389678

    Open questions at the time
    • Functional contribution of these piRNA-mRNA complexes versus piRNA-independent activity unclear
    • Cleavage versus binding not distinguished

Open questions

Synthesis pass · forward-looking unresolved questions
  • How PIWIL1 switches between germline piRNA-guided slicing and its diverse piRNA-independent somatic functions — and what determines target selection in each mode — remains unresolved.
  • No unifying mechanism linking NMD modulation, mRNA stabilization, Stathmin1 binding, and DNA hypermethylation
  • No structure of human PIWIL1 in either germline or somatic complexes
  • Direct molecular bridge to DNA methylation machinery unidentified

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003723 RNA binding 3 GO:0140098 catalytic activity, acting on RNA 3 GO:0016787 hydrolase activity 2
Localization
GO:0005634 nucleus 2 GO:0005829 cytosol 1
Pathway
R-HSA-1474165 Reproduction 3 R-HSA-1643685 Disease 3 R-HSA-8953854 Metabolism of RNA 3
Partners
GTSF1HENMT1PRMT5STMN1TDRD2UPF1
Complex memberships
piRISC

Evidence

Reading pass · 21 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2002 HIWI (PIWIL1) protein is specifically expressed in germline cells of the human testis, detectable in spermatocytes and round spermatids during spermatogenesis, and its transient overexpression in the human leukemia cell line KG1 caused a dramatic reduction in cellular proliferation and induced programmed cell death (Annexin V assay), suggesting a role as a negative developmental regulator. RT-PCR, immunohistochemistry, transient transfection overexpression, Annexin V apoptosis assay Oncogene Medium 12037681
2001 HIWI (PIWIL1) is expressed in human CD34+ hematopoietic progenitor/stem cells but not in more differentiated cell populations; overexpression in KG1 cells reduced proliferation and induced apoptosis, implicating HIWI in stem cell self-renewal regulation. RT-PCR, 5' RACE cloning, transient transfection, cell proliferation assay, Annexin V apoptosis assay Blood Medium 11154219
2010 PIWI proteins (including PIWIL1/HIWI) contain symmetrical dimethyl arginines (sDMAs) installed by the methyltransferase PRMT5 (Dart5/Capsuleen), and this modification mediates interaction with multiple Tudor (Tud) domain family proteins through sDMA-binding Tudor domains. Biochemical analysis, methyltransferase assays, co-immunoprecipitation, review of prior experimental findings Genes & development Medium 20360382
2017 The extended Tudor domain of mammalian TDRD2 preferentially recognizes an unmethylated arginine-rich sequence from PIWIL1, in contrast to most Tudor domains that bind methylated PIWI proteins. Structural studies revealed that the interface of Tudor and staphylococcal nuclease domains mediates PIWIL1 peptide recognition. Mutations disrupting the TDRD2-PIWIL1 interaction compromised piRNA maturation via 3'-end trimming in vitro. Crystal structure determination, in vitro binding assays, mutagenesis, in vitro trimming assay Proceedings of the National Academy of Sciences of the United States of America High 29118143
2022 GTSF1 potentiates the weak intrinsic piRNA-directed RNA cleavage activity of PIWI proteins including MIWI (PIWIL1), transforming them into efficient endoribonucleases. The piRNA-guided endoribonuclease activities of MIWI (PIWIL1) and MILI are essential for the production of functional sperm. In vitro cleavage assay, biochemical reconstitution, mouse knockout phenotype Nature High 35772669
2023 Unlike AGO-clade Argonaute proteins, PIWI proteins (tested with mouse MIWI/PIWIL1 and sponge PIWI) efficiently cleave target transcripts that are only partially base-paired to their piRNA guides. PIWI slicing tolerates mismatches to any target nucleotide including those flanking the scissile phosphate, and seed pairing is dispensable for PIWI binding or cleavage, in contrast to AGO proteins. In vitro target binding and cleavage assays with recombinant mouse and sponge PIWI proteins, mutational analysis of target:piRNA pairing Nature High 37344600
2024 Cryo-EM structural studies of mammalian MILI (PIWIL2) and human HILI piRNA-induced silencing complexes (piRISCs) revealed that they bind and cleave targets more efficiently than invertebrate counterparts, adopt a wider nucleic-acid-binding channel, and display an extended prearranged piRNA seed in the absence of target. A vertebrate-specific lysine distorts the piRNA seed, shifting the trajectory of the piRNA-target duplex, and promotes target binding and cleavage. In the presence of target, the seed gate adopts a relaxed state, enabling PIWI tolerance of seed-target mismatches. Cryo-EM structure determination, in vitro binding and cleavage assays, mutagenesis of vertebrate-specific lysine Nature structural & molecular biology High 38658622
2020 PIWIL1 is highly expressed in gastric cancer tissues and cell lines. PIWIL1 knockout drastically reduces gastric cancer cell proliferation, migration, metastasis, and tumorigenesis. Surprisingly, few bona fide piRNAs exist in gastric cancer cells, and abolishing the piRNA-binding activity of PIWIL1 does not affect its oncogenic function. PIWIL1 function in gastric cancer involves piRNA-independent interaction with the UPF1-mediated nonsense-mediated mRNA decay (NMD) mechanism. PIWIL1 knockout (PIWIL1-KO), RNA deep sequencing, piRNA-binding mutant functional rescue, co-immunoprecipitation with NMD factors Proceedings of the National Academy of Sciences of the United States of America High 32848063
2015 PIWIL1 directly binds to Stathmin1, upregulates Stathmin1 expression by inhibiting ubiquitin-mediated degradation induced by the E3 ubiquitin ligase RLIM, and reduces phosphorylation of Stathmin1 at Ser-16 by inhibiting the interaction between CaMKII and Stathmin1. Through these mechanisms PIWIL1 suppresses microtubule polymerization and promotes cell proliferation and migration. Co-immunoprecipitation (direct binding), Western blot, ubiquitination assay, phosphorylation assay, microtubule polymerization assay, cell proliferation and migration assays Oncotarget Medium 26317901
2012 Overexpression of HIWI (PIWIL1) in sarcoma precursors inhibits their differentiation in vitro and generates sarcomas in vivo. Transgenic mice expressing mesodermally-restricted Hiwi develop sarcomas. Inducible downregulation of Hiwi in human sarcomas inhibits growth and re-establishes differentiation. Hiwi-associated tumorigenesis is accompanied by increased global DNA methylation and silencing of cyclin-dependent kinase inhibitors (CDKIs), which is reversible with DNA methyltransferase inhibitors. In vitro differentiation assays, transgenic mouse model, inducible knockdown, DNA methylation analysis, DNMT inhibitor treatment PloS one High 22438986
2021 Piwil1 (HIWI) is enriched in glioma stem-like cells (GSCs) and maintains self-renewal. Silencing Piwil1 in GSCs leads to global gene expression changes resulting in cell-cycle arrest, senescence, or apoptosis. Piwil1 knockdown increases expression of BTG2 and FBXW7 (leading to reduced c-Myc and loss of Olig2/Nestin), and Piwil1 regulates mRNA stability of BTG2, FBXW7, and CDKN1B. In animal models, Piwil1 knockdown suppresses tumor growth. shRNA knockdown, RNA-seq, mRNA stability assay, xenograft mouse models, Western blot Cell reports Medium 33406417
2021 PIWIL1 in HCC increases oxygen consumption and energy production via fatty acid metabolism without altering aerobic glycolysis. PIWIL1-overexpressing HCC cells attract myeloid-derived suppressor cells (MDSCs) into the tumor microenvironment. Complement C3, whose secretion is induced by PIWIL1 in HCC cells, mediates the interaction with MDSCs by activating p38 MAPK signaling in MDSCs, which initiates immunosuppressive IL-10 expression. PIWIL1 overexpression/knockdown, metabolic flux analysis, RNA-seq, MDSC co-culture, C3 neutralization, p38 MAPK inhibition Signal transduction and targeted therapy Medium 33633112
2019 PIWIL1 localizes in a nuage-like perinuclear structure in colorectal cancer (COLO 205) cells. RNA immunoprecipitation revealed piRNAs loaded into PIWIL1 that form complexes also comprising target mRNAs encoding key regulatory proteins involved in colorectal carcinogenesis. Immunofluorescence localization, RNA immunoprecipitation (RIP), piRNA methylation analysis, RNA sequencing Cells Medium 31694219
2012 RASSF1C overexpression increases PIWIL1 gene expression in non-small cell lung cancer cells, and RASSF1C silencing decreases PIWIL1 expression. Inhibition of the MEK-ERK1/2 pathway suppresses PIWIL1 expression, suggesting RASSF1C regulates PIWIL1 through activation of the MEK-ERK1/2 pathway. RASSF1C overexpression, siRNA knockdown, Affymetrix microarray, qRT-PCR, Western blot, immunostaining, MEK inhibitor treatment BMC research notes Low 22591718
2021 PIWIL1 interacting with piR-017061 facilitates EFNA5 mRNA degradation in pancreatic cancer cells. Loss of piR-017061 results in EFNA5 accumulation, which promotes pancreatic cancer development. Direct binding between piR-017061 and EFNA5 mRNA is mediated by PIWIL1. RNA immunoprecipitation, qRT-PCR, cell proliferation assay, in vivo xenograft, mRNA stability assay Human cell Medium 33389678
2015 HENMT1 and PIWIL1 are coexpressed in pachytene spermatocytes and spermatids in the human testis, and samples expressing HENMT1, PIWIL1, and PIWIL2 showed significantly lower transposon (LINE-1) expression compared to samples without expression, supporting conservation of the transposon-regulating piRNA pathway function in adult human testis. Immunolocalization, RT-PCR after laser microdissection, RT-qPCR in 222 testis biopsies, HENMT1-overexpressing cell lines Reproduction (Cambridge, England) Medium 28676534
2023 In golden hamsters, knockout of PIWIL1 leads to female sterility (PIWIL1 is highly expressed throughout oogenesis and early embryogenesis), and PIWIL1 can partially compensate for TE silencing in PIWIL3 knockout females. In testes, PIWIL1 is the predominant PIWI in adult hamsters, and loss of PIWIL1 causes sterility with severe spermatogenesis disorders. CRISPR/Cas9 knockout, immunofluorescence subcellular localization, RNA-seq, fertility assay Nature communications High 37644029
2015 In human testes, PIWIL1 is expressed in spermatocytes and spermatids and is associated with transposon silencing. Expression of PIWIL1 in oocytes of adult human and bovine ovaries was demonstrated, with PIWIL1 expressed in adult human and bovine oocytes (bovine oocytes exclusively expressed PIWIL1 among PIWI family members). Immunohistochemistry, Western blot, quantitative proteome analysis, small RNA sequencing Cell reports Medium 25818294
2015 PIWIL1 overexpression in endometrial cancer cells maintained stem-like characteristics including enhanced migration, invasion, and sphere-forming activity, and induced an EMT-like phenotype with increased mesenchymal markers and suppression of E-cadherin, as well as upregulated CD44 and ALDH1 expression. Stable transfection/shRNA knockdown, MTT assay, colony formation, transwell invasion, xenograft, qRT-PCR, Western blot, immunofluorescence BMC cancer Medium 26506848
2019 Viral-mediated knockdown of Piwil1 in the dorsal hippocampus of adult mice leads to enhanced contextual fear memory without affecting generalized anxiety, implicating PIWIL1 in behavioral regulation in the adult mammalian brain through modulation of plasticity-related gene expression. Viral-mediated shRNA knockdown in vivo, contextual fear conditioning behavioral assay Neurobiology of learning and memory Low 30965112
2015 In colorectal cancer cells, Hiwi (PIWIL1) overexpression promoted cell proliferation and increased global DNA methylation levels. Chemical inhibition of DNA methylation significantly restrained this proliferation-promoting effect, suggesting PIWIL1 promotes cancer cell proliferation via increasing global DNA methylation. Adenovirus-mediated overexpression, cell proliferation assay, global DNA methylation measurement, DNMT inhibitor treatment Disease markers Low 26355242

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2019 PIWI-interacting RNAs: small RNAs with big functions. Nature reviews. Genetics 853 30446728
2007 The Piwi-piRNA pathway provides an adaptive defense in the transposon arms race. Science (New York, N.Y.) 808 17975059
2015 PIWI-Interacting RNA: Its Biogenesis and Functions. Annual review of biochemistry 552 25747396
2009 The biogenesis and function of PIWI proteins and piRNAs: progress and prospect. Annual review of cell and developmental biology 413 19575643
2014 PIWI proteins and PIWI-interacting RNAs in the soma. Nature 330 24429634
2011 Uniting germline and stem cells: the function of Piwi proteins and the piRNA pathway in diverse organisms. Annual review of genetics 294 21942366
2019 The emerging role of the piRNA/piwi complex in cancer. Molecular cancer 271 31399034
2013 PIWI-interacting RNAs: from generation to transgenerational epigenetics. Nature reviews. Genetics 256 23797853
2002 Molecular characterization of hiwi, a human member of the piwi gene family whose overexpression is correlated to seminomas. Oncogene 243 12037681
2022 Emerging roles and functional mechanisms of PIWI-interacting RNAs. Nature reviews. Molecular cell biology 217 36104626
2015 Piwi proteins and piRNAs in mammalian oocytes and early embryos. Cell reports 182 25818294
2001 Human CD34(+) stem cells express the hiwi gene, a human homologue of the Drosophila gene piwi. Blood 169 11154219
2010 How does the royal family of Tudor rule the PIWI-interacting RNA pathway? Genes & development 159 20360382
2006 Expression of hiwi gene in human gastric cancer was associated with proliferation of cancer cells. International journal of cancer 152 16287078
2013 PIWI proteins and PIWI-interacting RNAs function in Hydra somatic stem cells. Proceedings of the National Academy of Sciences of the United States of America 127 24367095
2012 Piwis and piwi-interacting RNAs in the epigenetics of cancer. Journal of cellular biochemistry 125 21928326
2012 PIWI Expression and Function in Cancer. Frontiers in genetics 118 23087701
2017 PIWI Proteins and PIWI-Interacting RNA: Emerging Roles in Cancer. Cellular physiology and biochemistry : international journal of experimental cellular physiology, biochemistry, and pharmacology 111 29130960
2013 Beyond transposons: the epigenetic and somatic functions of the Piwi-piRNA mechanism. Current opinion in cell biology 111 23465540
2006 Expression of the stem cell self-renewal gene Hiwi and risk of tumour-related death in patients with soft-tissue sarcoma. Oncogene 108 16953229
2008 The stem cell-associated Hiwi gene in human adenocarcinoma of the pancreas: expression and risk of tumour-related death. British journal of cancer 103 18781170
2017 PIWI-Interacting RNA in Drosophila: Biogenesis, Transposon Regulation, and Beyond. Chemical reviews 93 29281264
2013 Untangling the web: the diverse functions of the PIWI/piRNA pathway. Molecular reproduction and development 86 23712694
2023 Relaxed targeting rules help PIWI proteins silence transposons. Nature 73 37344600
2020 PIWIL1 promotes gastric cancer via a piRNA-independent mechanism. Proceedings of the National Academy of Sciences of the United States of America 70 32848063
2021 The biogenesis and biological function of PIWI-interacting RNA in cancer. Journal of hematology & oncology 65 34118972
2012 Hiwi mediated tumorigenesis is associated with DNA hypermethylation. PloS one 64 22438986
2023 The epigenetic regulatory mechanism of PIWI/piRNAs in human cancers. Molecular cancer 62 36882835
2020 Functions of PIWI Proteins in Gene Regulation: New Arrows Added to the piRNA Quiver. Trends in genetics : TIG 61 32951946
2020 Crystal structure of Drosophila Piwi. Nature communications 57 32051406
2016 Abundant and Altered Expression of PIWI-Interacting RNAs during Cardiac Hypertrophy. Heart, lung & circulation 57 27067666
2019 PIWI Proteins and piRNAs in the Nervous System. Molecules and cells 56 31838836
2021 Terminal modification, sequence, length, and PIWI-protein identity determine piRNA stability. Molecular cell 55 34626567
2021 PIWIL1 governs the crosstalk of cancer cell metabolism and immunosuppressive microenvironment in hepatocellular carcinoma. Signal transduction and targeted therapy 54 33633112
2019 The emerging roles of PIWI-interacting RNA in human cancers. Cancer management and research 51 31303794
2022 GTSF1 accelerates target RNA cleavage by PIWI-clade Argonaute proteins. Nature 50 35772669
2012 Concise review: The Piwi-piRNA axis: pivotal beyond transposon silencing. Stem cells (Dayton, Ohio) 50 22996918
2021 Piwil1 Regulates Glioma Stem Cell Maintenance and Glioblastoma Progression. Cell reports 49 33406417
2019 PIWI pathway against viruses in insects. Wiley interdisciplinary reviews. RNA 48 31183996
2014 Hiwi facilitates chemoresistance as a cancer stem cell marker in cervical cancer. Oncology reports 48 25119492
2014 Silencing HIWI suppresses the growth, invasion and migration of glioma cells. International journal of oncology 48 25269862
2014 Piwi and potency: PIWI proteins in animal stem cells and regeneration. Integrative and comparative biology 47 24948137
2017 Cancer-testis gene PIWIL1 promotes cell proliferation, migration, and invasion in lung adenocarcinoma. Cancer medicine 44 29168346
2015 Stem cell protein Piwil1 endowed endometrial cancer cells with stem-like properties via inducing epithelial-mesenchymal transition. BMC cancer 39 26506848
2020 Role of PIWI-interacting RNAs on cell survival: Proliferation, apoptosis, and cycle. IUBMB life 38 32697419
2017 The Piwi-piRNA pathway: road to immortality. Aging cell 37 28653810
2017 Expression patterns of HENMT1 and PIWIL1 in human testis: implications for transposon expression. Reproduction (Cambridge, England) 37 28676534
2014 Hiwi downregulation, mediated by shRNA, reduces the proliferation and migration of human hepatocellular carcinoma cells. Molecular medicine reports 37 25370791
2010 Expression of PSCA, PIWIL1, and TBX2 in endometrial adenocarcinoma. Onkologie 37 20502058
2023 The burgeoning importance of PIWI-interacting RNAs in cancer progression. Science China. Life sciences 36 38198029
2017 MiRNA-154-5p inhibits cell proliferation and metastasis by targeting PIWIL1 in glioblastoma. Brain research 35 28842123
2014 Overexpression of hiwi promotes growth of human breast cancer cells. Asian Pacific journal of cancer prevention : APJCP 34 25292027
2018 PIWI-piRNA pathway: Setting the pace of aging by reducing DNA damage. Mechanisms of ageing and development 32 29580825
2024 Critical appraisal of the piRNA-PIWI axis in cancer and cancer stem cells. Biomarker research 31 38303021
2021 PIWIL1 interacting RNA piR-017061 inhibits pancreatic cancer growth via regulating EFNA5. Human cell 31 33389678
2021 Piwi-Interacting RNAs: A New Class of Regulator in Human Breast Cancer. Frontiers in oncology 31 34295823
2020 PIWI-interacting RNAs in human cancer. Seminars in cancer biology 31 32877760
2019 PIWI-interacting RNA 39980 promotes tumor progression and reduces drug sensitivity in neuroblastoma cells. Journal of cellular physiology 31 31478570
2018 Epigenetic roles of PIWI‑interacting RNAs (piRNAs) in cancer metastasis (Review). Oncology reports 31 30226604
2015 Piwi-pathway alteration induces LINE-1 transposon derepression and infertility development in cryptorchidism. Sexual development : genetics, molecular biology, evolution, endocrinology, embryology, and pathology of sex determination and differentiation 31 25791297
2021 A Review of Discovery Profiling of PIWI-Interacting RNAs and Their Diverse Functions in Metazoans. International journal of molecular sciences 30 34681826
2017 Structural basis for arginine methylation-independent recognition of PIWIL1 by TDRD2. Proceedings of the National Academy of Sciences of the United States of America 30 29118143
2010 The universal overexpression of a cancer testis antigen hiwi is associated with cancer angiogenesis. Oncology reports 30 20204292
2023 Novel roles of PIWI proteins and PIWI-interacting RNAs in human health and diseases. Cell communication and signaling : CCS 28 38031146
2023 What Are the Functional Roles of Piwi Proteins and piRNAs in Insects? Insects 27 36835756
2023 PIWI-interacting RNAs: Critical roles and therapeutic targets in cancer. Cancer letters 27 37076042
2024 AVCAPIR: A Novel Procalcific PIWI-Interacting RNA in Calcific Aortic Valve Disease. Circulation 25 38258575
2024 Mammalian PIWI-piRNA-target complexes reveal features for broad and efficient target silencing. Nature structural & molecular biology 25 38658622
2019 Disrupting the hippocampal Piwi pathway enhances contextual fear memory in mice. Neurobiology of learning and memory 25 30965112
2016 Expression and Regulation of PIWIL-Proteins and PIWI-Interacting RNAs in Rheumatoid Arthritis. PloS one 25 27893851
2020 PIWI-piRNA pathway-mediated transposable element repression in Hydra somatic stem cells. RNA (New York, N.Y.) 24 32075940
2018 Transposons and the PIWI pathway: genome defense in gametes and embryos. Reproduction (Cambridge, England) 24 30304932
2015 Correlation of HIWI and HILI Expression with Cancer Stem Cell Markers in Colorectal Cancer. Anticancer research 24 26026091
2013 Hiwi knockdown inhibits the growth of lung cancer in nude mice. Asian Pacific journal of cancer prevention : APJCP 24 23621188
2012 RASSF1C modulates the expression of a stem cell renewal gene, PIWIL1. BMC research notes 24 22591718
2020 PIWI-interacting RNAs: Mitochondria-based biogenesis and functions in cancer. Genes & diseases 23 34291132
2018 Investigation of piwi-interacting RNA pathway genes role in idiopathic non-obstructive azoospermia. Scientific reports 23 29317647
2019 Molecular and Functional Characterization of the Somatic PIWIL1/piRNA Pathway in Colorectal Cancer Cells. Cells 22 31694219
2012 Altered expression of porcine Piwi genes and piRNA during development. PloS one 22 22952772
2022 piRNA/PIWI Protein Complex as a Potential Biomarker in Sporadic Amyotrophic Lateral Sclerosis. Molecular neurobiology 21 35015250
2022 PIWI-Interacting RNAs (piRNAs): Promising Applications as Emerging Biomarkers for Digestive System Cancer. Frontiers in molecular biosciences 21 35155584
2018 Investigating piwi-interacting RNA regulome in human neuroblastoma. Genes, chromosomes & cancer 21 29516567
2018 Aberrant Expression of PIWIL1 and PIWIL2 and Their Clinical Significance in Ductal Breast Carcinoma. Anticancer research 21 29599319
2017 Drosophila PAF1 Modulates PIWI/piRNA Silencing Capacity. Current biology : CB 21 28844648
2015 Hiwi Promotes the Proliferation of Colorectal Cancer Cells via Upregulating Global DNA Methylation. Disease markers 21 26355242
2021 Critical Roles of PIWIL1 in Human Tumors: Expression, Functions, Mechanisms, and Potential Clinical Implications. Frontiers in cell and developmental biology 20 33718392
2019 Epigenetic roles of PIWI proteins and piRNAs in lung cancer. Cell & bioscience 20 31890151
2017 The PIWI-Interacting RNA Molecular Pathway: Insights From Cultured Silkworm Germline Cells. BioEssays : news and reviews in molecular, cellular and developmental biology 20 29164638
2023 PIWIL1 interacting RNA piR-017724 inhibits proliferation, invasion, and migration, and inhibits the development of HCC by silencing PLIN3. Frontiers in oncology 19 37503320
2020 PIWI-interacting RNAs and PIWI proteins in glioma: molecular pathogenesis and role as biomarkers. Cell communication and signaling : CCS 19 33109195
2011 Expression of HIWI in human hepatocellular carcinoma. Cell biochemistry and biophysics 19 21327579
2023 PIWI-interacting RNA expression regulates pathogenesis in a Caenorhabditis elegans model of Lewy body disease. Nature communications 18 37783675
2021 Knockout Gene-Based Evidence for PIWI-Interacting RNA Pathway in Mammals. Frontiers in cell and developmental biology 18 34336834
2020 DEPS-1 is required for piRNA-dependent silencing and PIWI condensate organisation in Caenorhabditis elegans. Nature communications 18 32843637
2020 Diverse Defenses: A Perspective Comparing Dipteran Piwi-piRNA Pathways. Cells 18 32992598
2023 The non-redundant functions of PIWI family proteins in gametogenesis in golden hamsters. Nature communications 17 37644029
2022 PiWi RNA in Neurodevelopment and Neurodegenerative Disorders. Current molecular pharmacology 17 34212832
2022 Emerging roles of PIWI-interacting RNAs (piRNAs) and PIWI proteins in head and neck cancer and their potential clinical implications. Biochimica et biophysica acta. Reviews on cancer 17 35931391
2020 Effects of fluoride on PIWI-interacting RNA expression profiling in testis of mice. Chemosphere 17 33213873
2015 PIWIL1 destabilizes microtubule by suppressing phosphorylation at Ser16 and RLIM-mediated degradation of Stathmin1. Oncotarget 17 26317901

Missed literature

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