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Piwi

January 01, 1970

Piwi (or PIWI) genes were identified as regulatory proteins responsible for stem cell and germ cell differentiation.[4] Piwi is an abbreviation of P-element Induced WImpy testis[a] in Drosophila.[6] Piwi proteins are highly conserved RNA-binding proteins and are present in both plants and animals.[7] Piwi proteins belong to the Argonaute/Piwi family and have been classified as nuclear proteins. Studies on Drosophila have also indicated that Piwi proteins have no slicer activity conferred by the presence of the Piwi domain.[8] In addition, Piwi associates with heterochromatin protein 1, an epigenetic modifier, and piRNA-complementary sequences. These are indications of the role Piwi plays in epigenetic regulation. Piwi proteins are also thought to control the biogenesis of piRNA as many Piwi-like proteins contain slicer activity which would allow Piwi proteins to process precursor piRNA into mature piRNA.

Piwi domain
Structure of the Pyrococcus furiosus Argonaute protein.[1]
Identifiers
SymbolPiwi
PfamPF02171
InterProIPR003165
PROSITEPS50822
CDDcd02826
Available protein structures:
Pfam  structures / ECOD  
PDBRCSB PDB; PDBe; PDBj
PDBsumstructure summary
PDB1u04​, 1w9h​, 1ytu​, 1yvu​, 1z25​, 1z26​, 2bgg​, 2f8s​, 2f8t​, 2nub​, 2w42​
The piwi domain of an argonaute protein with bound siRNA, components of the RNA-induced silencing complex that mediates gene silencing by RNA interference.
All human Piwi proteins and argonaute proteins have the same RNA binding domains, PAZ and Piwi.[2]
Piwi-piRNA interactions: Within the nucleus, this pathway is involved in DNA methylation (A), histone methylation of H3K9 through interactions with heterochromatin protein 1 (HP1) and H3K9 histone methyltransferase (B). The Piwi-piRNA pathway also interacts with the elF translational initiator (C).[3]

Protein structure and function edit

The structure of several Piwi and Argonaute proteins (Ago) have been solved. Piwi proteins are RNA-binding proteins with 2 or 3 domains: The N-terminal PAZ domain binds the 3'-end of the guide RNA; the middle MID domain binds the 5'-phosphate of RNA; and the C-terminal PIWI domain acts as an RNase H endonuclease that can cleave RNA.[9][10] The small RNA partners of Ago proteins are microRNAs (miRNAs). Ago proteins utilize miRNAs to silence genes post-transcriptionally or use small-interfering RNAs (siRNAs) in both transcription and post-transcription silencing mechanisms. Piwi proteins interact with piRNAs (28–33 nucleotides) that are longer than miRNAs and siRNAs (~20 nucleotides), suggesting that their functions are distinct from those of Ago proteins.[9]

Human Piwi proteins edit

Presently there are four known human Piwi proteins—PIWI-like protein 1, PIWI-like protein 2, PIWI-like protein 3 and PIWI-like protein 4. Human Piwi proteins all contain two RNA binding domains, PAZ and Piwi. The four PIWI-like proteins have a spacious binding site within the PAZ domain which allows them to bind the bulky 2’-OCH3 at the 3’ end of piwi-interacting RNA.[11]

One of the major human homologues, whose upregulation is implicated in the formation of tumours such as seminomas, is called hiwi (for human piwi).[12]

Homologous proteins in mice have been called miwi (for mouse piwi).[13]

Role in germline cells edit

PIWI proteins play a crucial role in fertility and germline development across animals and ciliates. Recently identified as a polar granule component, PIWI proteins appear to control germ cell formation so much so that in the absence of PIWI proteins there is a significant decrease in germ cell formation. Similar observations were made with the mouse homologs of PIWI, MILI, MIWI and MIWI2. These homologs are known to be present in spermatogenesis. Miwi is expressed in various stages of spermatocyte formation and spermatid elongation where Miwi2 is expressed in Sertoli cells. Mice deficient in either Mili or Miwi-2 have experienced spermatogenic stem cell arrest and those lacking Miwi-2 underwent a degradation of spermatogonia.[14] The effects of piwi proteins in human and mouse germlines seems to stem from their involvement in translation control as Piwi and the small noncoding RNA, piwi-interacting RNA (piRNA), have been known to co-fractionate polysomes. The piwi-piRNA pathway also induces heterochromatin formation at centromeres,[15] thus affecting transcription. The piwi-piRNA pathway also appears to protect the genome. First observed in Drosophila, mutant piwi-piRNA pathways led to a direct increase in dsDNA breaks in ovarian germ cells. The role of the piwi-piRNA pathway in transposon silencing may be responsible for the reduction in dsDNA breaks in germ cells.

Role in RNA interference edit

The piwi domain[16] is a protein domain found in piwi proteins and a large number of related nucleic acid-binding proteins, especially those that bind and cleave RNA. The function of the domain is double stranded-RNA-guided hydrolysis of single stranded-RNA that has been determined in the argonaute family of related proteins.[1] Argonautes, the most well-studied family of nucleic-acid binding proteins, are RNase H-like enzymes that carry out the catalytic functions of the RNA-induced silencing complex (RISC). In the well-known cellular process of RNA interference, the argonaute protein in the RISC complex can bind both small interfering RNA (siRNA) generated from exogenous double-stranded RNA and microRNA (miRNA) generated from endogenous non-coding RNA, both produced by the ribonuclease Dicer, to form an RNA-RISC complex. This complex binds and cleaves complementary base pairing messenger RNA, destroying it and preventing its translation into protein. Crystallised piwi domains have a conserved basic binding site for the 5' end of bound RNA; in the case of argonaute proteins binding siRNA strands, the last unpaired nucleotide base of the siRNA is also stabilised by base stacking-interactions between the base and neighbouring tyrosine residues.[17]

Recent evidence suggests that the functional role of piwi proteins in germ-line determination is due to their capacity to interact with miRNAs. Components of the miRNA pathway appear to be present in pole plasm and to play a key role in early development and morphogenesis of Drosophila melanogaster embryos, in which germ-line maintenance has been extensively studied.[18]

piRNAs and transposon silencing edit

A novel class of longer-than-average miRNAs known as Piwi-interacting RNAs (piRNAs) has been defined in mammalian cells, about 26-31 nucleotides long as compared to the more typical miRNA or siRNA of about 21 nucleotides. These piRNAs are expressed mainly in spermatogenic cells in the testes of mammals.[19] But studies have reported that piRNA expression can be found in the ovarian somatic cells and neuron cells in invertebrates, as well as in many other mammalian somatic cells. piRNAs have been identified in the genomes of mice, rats, and humans, with an unusual "clustered" genomic organization[20] that may originate from repetitive regions of the genome such as retrotransposons or regions normally organized into heterochromatin, and which are normally derived exclusively from the antisense strand of double-stranded RNA.[21] piRNAs have thus been classified as repeat-associated small interfering RNAs (rasiRNAs).[22]

Although their biogenesis is not yet well understood, piRNAs and Piwi proteins are thought to form an endogenous system for silencing the expression of selfish genetic elements such as retrotransposons and thus preventing the gene products of such sequences from interfering with germ cell formation.[21][23]

Footnotes edit

  1. ^ The word Wimpy itself comes from White color of eyes and Impotency [or profound infertility]. Y is added as a suffix to form a more pronounceable and coherent word.[5]

References edit

  1. ^ a b Rivas FV, Tolia NH, Song JJ, et al. (April 2005). "Purified Argonaute2 and an siRNA form recombinant human RISC". Nat. Struct. Mol. Biol. 12 (4): 340–9. doi:10.1038/nsmb918. PMID 15800637. S2CID 2021813.
  2. ^ "Uniprot: The Universal knowledge database". Nucleic Acids Research. 45 (D1): D158–D169. 2017. doi:10.1093/nar/gkw1099. PMC 5210571. PMID 27899622.
  3. ^ Lindse K (2013). "Piwi-RNAs, the Defenders of the Genome". {{cite journal}}: Cite journal requires |journal= (help)
  4. ^ Cox DN, Chao A, Lin H (2000). "piwi encodes a nucleoplasmic factor whose activity modulates the number and division rate of germline stem cells". Development. 127 (3): 503–14. doi:10.1242/dev.127.3.503. PMID 10631171.
  5. ^ Ross RJ, Weiner MM, Lin H (2014). "PIWI proteins and PIWI-interacting RNAs in the soma". Nature. 505 (7483): 353–359. doi:10.1038/nature12987. PMC 4265809. PMID 24429634.
  6. ^ Lin H, Spradling AC (1997). "A novel group of pumilio mutations affects the asymmetric division of germline stem cells in the Drosophila ovary". Development. 124 (12): 2463–2476. doi:10.1242/dev.124.12.2463. PMID 9199372.
  7. ^ Cox DN, Chao A, Baker J, Chang L, Qiao D, Lin H (1998). "A novel class of evolutionarily conserved genes defined by piwi are essential for stem cell self-renewal". Genes Dev. 12 (23): 3715–27. doi:10.1101/gad.12.23.3715. PMC 317255. PMID 9851978.
  8. ^ Darricarrere N, Liu N, Watanabe T, Lin H (2013). "Function of Piwi, a nuclear Piwi/Argonaute protein, is independent of its slicer activity". Proc Natl Acad Sci USA. 110 (6): 1297–1302. Bibcode:2013PNAS..110.1297D. doi:10.1073/pnas.1213283110. PMC 3557079. PMID 23297219.
  9. ^ a b Zeng, Lei; Zhang, Qiang; Yan, Kelley; Zhou, Ming-Ming (2011-06-01). "Structural insights into piRNA recognition by the human PIWI-like 1 PAZ domain". Proteins: Structure, Function, and Bioinformatics. 79 (6): 2004–2009. doi:10.1002/prot.23003. ISSN 1097-0134. PMC 3092821. PMID 21465557.
  10. ^ Wei, Kai-Fa; Wu, Ling-Juan; Chen, Juan; Chen, Yan-feng; Xie, Dao-Xin (August 2012). "Structural evolution and functional diversification analyses of argonaute protein". Journal of Cellular Biochemistry. 113 (8): 2576–2585. doi:10.1002/jcb.24133. ISSN 1097-4644. PMID 22415963. S2CID 25990631.
  11. ^ Tian Y, Simanshu D, Ma J, Patel D (2010). "Structural basis for piRNA 2'-O-methylated 3'-end recognition by Piwi PAZ (Piwi/Argonaute/Zwille) domains". Proc. Natl. Acad. Sci. USA. 108 (3): 903–910. doi:10.1073/pnas.1017762108. PMC 3024652. PMID 21193640.
  12. ^ Qiao D, Zeeman AM, Deng W, Looijenga LH, Lin H (2002). "Molecular characterization of hiwi, a human member of the piwi gene family whose overexpression is correlated to seminomas". Oncogene. 21 (25): 3988–99. doi:10.1038/sj.onc.1205505. PMID 12037681.
  13. ^ Deng W, Lin H (2002). "miwi, a murine homolog of piwi, encodes a cytoplasmic protein essential for spermatogenesis". Dev Cell. 2 (6): 819–30. doi:10.1016/s1534-5807(02)00165-x. PMID 12062093.
  14. ^ Mani S, Juliano C (2013). "Untangling the Web: The Diverse Functions of the PIWI/piRNA Pathway". Mol. Reprod. Dev. 80 (8): 632–664. doi:10.1002/mrd.22195. PMC 4234069. PMID 23712694.
  15. ^ Thomson T, Lin H (2009). "The Biogenesis and Function PIWI Proteins and piRNAs: Progress and Prospect". Annu. Rev. Cell Dev. Biol. 25: 355–376. doi:10.1146/annurev.cellbio.24.110707.175327. PMC 2780330. PMID 19575643.
  16. ^ Cerutti L, Mian N, Bateman A (October 2000). "Domains in gene silencing and cell differentiation proteins: the novel PAZ domain and redefinition of the Piwi domain". Trends Biochem. Sci. 25 (10): 481–2. doi:10.1016/S0968-0004(00)01641-8. PMID 11050429.
  17. ^ Ma J, Yuan Y, Meister G, Pei Y, Tuschl T, Patel D (2005). "Structural basis for 5'-end-specific recognition of guide RNA by the A. fulgidus Piwi protein". Nature. 434 (7033): 666–70. Bibcode:2005Natur.434..666M. doi:10.1038/nature03514. PMC 4694588. PMID 15800629.
  18. ^ Megosh HB, Cox DN, Campbell C, Lin H (2006). "The role of PIWI and the miRNA machinery in Drosophila germline determination". Curr Biol. 16 (19): 1884–94. doi:10.1016/j.cub.2006.08.051. PMID 16949822. S2CID 6397874.
  19. ^ Kim VN (2006). "Small RNAs just got bigger: Piwi-interacting RNAs (piRNAs) in mammalian testes". Genes Dev. 20 (15): 1993–7. doi:10.1101/gad.1456106. PMID 16882976.
  20. ^ Girard A, Sachidanandam R, Hannon GJ, Carmell MA (2006). "A germline-specific class of small RNAs binds mammalian Piwi proteins". Nature. 442 (7099): 199–202. Bibcode:2006Natur.442..199G. doi:10.1038/nature04917. PMID 16751776. S2CID 3185036.
  21. ^ a b Vagin VV, Sigova A, Li C, Seitz H, Gvozdev V, Zamore PD (2006). "A distinct small RNA pathway silences selfish genetic elements in the germline". Science. 313 (5785): 320–4. Bibcode:2006Sci...313..320V. doi:10.1126/science.1129333. PMID 16809489. S2CID 40471466.
  22. ^ Saito K, Nishida KM, Mori T, Kawamura Y, Miyoshi K, Nagami T, Siomi H, Siomi MC (2006). "Specific association of Piwi with rasiRNAs derived from retrotransposon and heterochromatic regions in the Drosophila genome". Genes Dev. 20 (16): 2214–22. doi:10.1101/gad.1454806. PMC 1553205. PMID 16882972.
  23. ^ Ozata DM, Gainetdinov I, Zoch A, Phillip D, Zamore PD (2019). "PIWI-interacting RNAs: small RNAs with big functions" (PDF). Nature Reviews Genetics. 20 (2): 89–108. doi:10.1038/s41576-018-0073-3. PMID 30446728. S2CID 53565676.

External links edit

  • SCOP 110640 – Piwi domain in SCOP
  • PDOC50822 – Piwi domain in PROSITE
  • UNIPROT Piwi - Piwi domains

January 01, 1970
piwi, piwi, genes, were, identified, regulatory, proteins, responsible, stem, cell, germ, cell, differentiation, abbreviation, element, induced, wimpy, testis, drosophila, proteins, highly, conserved, binding, proteins, present, both, plants, animals, proteins. Piwi or PIWI genes were identified as regulatory proteins responsible for stem cell and germ cell differentiation 4 Piwi is an abbreviation of P element Induced WImpy testis a in Drosophila 6 Piwi proteins are highly conserved RNA binding proteins and are present in both plants and animals 7 Piwi proteins belong to the Argonaute Piwi family and have been classified as nuclear proteins Studies on Drosophila have also indicated that Piwi proteins have no slicer activity conferred by the presence of the Piwi domain 8 In addition Piwi associates with heterochromatin protein 1 an epigenetic modifier and piRNA complementary sequences These are indications of the role Piwi plays in epigenetic regulation Piwi proteins are also thought to control the biogenesis of piRNA as many Piwi like proteins contain slicer activity which would allow Piwi proteins to process precursor piRNA into mature piRNA Piwi domainStructure of the Pyrococcus furiosus Argonaute protein 1 IdentifiersSymbolPiwiPfamPF02171InterProIPR003165PROSITEPS50822CDDcd02826Available protein structures Pfam structures ECOD PDBRCSB PDB PDBe PDBjPDBsumstructure summaryPDB1u04 1w9h 1ytu 1yvu 1z25 1z26 2bgg 2f8s 2f8t 2nub 2w42 The piwi domain of an argonaute protein with bound siRNA components of the RNA induced silencing complex that mediates gene silencing by RNA interference All human Piwi proteins and argonaute proteins have the same RNA binding domains PAZ and Piwi 2 Piwi piRNA interactions Within the nucleus this pathway is involved in DNA methylation A histone methylation of H3K9 through interactions with heterochromatin protein 1 HP1 and H3K9 histone methyltransferase B The Piwi piRNA pathway also interacts with the elF translational initiator C 3 Contents 1 Protein structure and function 2 Human Piwi proteins 3 Role in germline cells 4 Role in RNA interference 5 piRNAs and transposon silencing 6 Footnotes 7 References 8 External linksProtein structure and function editThe structure of several Piwi and Argonaute proteins Ago have been solved Piwi proteins are RNA binding proteins with 2 or 3 domains The N terminal PAZ domain binds the 3 end of the guide RNA the middle MID domain binds the 5 phosphate of RNA and the C terminal PIWI domain acts as an RNase H endonuclease that can cleave RNA 9 10 The small RNA partners of Ago proteins are microRNAs miRNAs Ago proteins utilize miRNAs to silence genes post transcriptionally or use small interfering RNAs siRNAs in both transcription and post transcription silencing mechanisms Piwi proteins interact with piRNAs 28 33 nucleotides that are longer than miRNAs and siRNAs 20 nucleotides suggesting that their functions are distinct from those of Ago proteins 9 Human Piwi proteins editPresently there are four known human Piwi proteins PIWI like protein 1 PIWI like protein 2 PIWI like protein 3 and PIWI like protein 4 Human Piwi proteins all contain two RNA binding domains PAZ and Piwi The four PIWI like proteins have a spacious binding site within the PAZ domain which allows them to bind the bulky 2 OCH3 at the 3 end of piwi interacting RNA 11 One of the major human homologues whose upregulation is implicated in the formation of tumours such as seminomas is called hiwi for human piwi 12 Homologous proteins in mice have been called miwi for mouse piwi 13 Role in germline cells editPIWI proteins play a crucial role in fertility and germline development across animals and ciliates Recently identified as a polar granule component PIWI proteins appear to control germ cell formation so much so that in the absence of PIWI proteins there is a significant decrease in germ cell formation Similar observations were made with the mouse homologs of PIWI MILI MIWI and MIWI2 These homologs are known to be present in spermatogenesis Miwi is expressed in various stages of spermatocyte formation and spermatid elongation where Miwi2 is expressed in Sertoli cells Mice deficient in either Mili or Miwi 2 have experienced spermatogenic stem cell arrest and those lacking Miwi 2 underwent a degradation of spermatogonia 14 The effects of piwi proteins in human and mouse germlines seems to stem from their involvement in translation control as Piwi and the small noncoding RNA piwi interacting RNA piRNA have been known to co fractionate polysomes The piwi piRNA pathway also induces heterochromatin formation at centromeres 15 thus affecting transcription The piwi piRNA pathway also appears to protect the genome First observed in Drosophila mutant piwi piRNA pathways led to a direct increase in dsDNA breaks in ovarian germ cells The role of the piwi piRNA pathway in transposon silencing may be responsible for the reduction in dsDNA breaks in germ cells Role in RNA interference editThe piwi domain 16 is a protein domain found in piwi proteins and a large number of related nucleic acid binding proteins especially those that bind and cleave RNA The function of the domain is double stranded RNA guided hydrolysis of single stranded RNA that has been determined in the argonaute family of related proteins 1 Argonautes the most well studied family of nucleic acid binding proteins are RNase H like enzymes that carry out the catalytic functions of the RNA induced silencing complex RISC In the well known cellular process of RNA interference the argonaute protein in the RISC complex can bind both small interfering RNA siRNA generated from exogenous double stranded RNA and microRNA miRNA generated from endogenous non coding RNA both produced by the ribonuclease Dicer to form an RNA RISC complex This complex binds and cleaves complementary base pairing messenger RNA destroying it and preventing its translation into protein Crystallised piwi domains have a conserved basic binding site for the 5 end of bound RNA in the case of argonaute proteins binding siRNA strands the last unpaired nucleotide base of the siRNA is also stabilised by base stacking interactions between the base and neighbouring tyrosine residues 17 Recent evidence suggests that the functional role of piwi proteins in germ line determination is due to their capacity to interact with miRNAs Components of the miRNA pathway appear to be present in pole plasm and to play a key role in early development and morphogenesis of Drosophila melanogaster embryos in which germ line maintenance has been extensively studied 18 piRNAs and transposon silencing editA novel class of longer than average miRNAs known as Piwi interacting RNAs piRNAs has been defined in mammalian cells about 26 31 nucleotides long as compared to the more typical miRNA or siRNA of about 21 nucleotides These piRNAs are expressed mainly in spermatogenic cells in the testes of mammals 19 But studies have reported that piRNA expression can be found in the ovarian somatic cells and neuron cells in invertebrates as well as in many other mammalian somatic cells piRNAs have been identified in the genomes of mice rats and humans with an unusual clustered genomic organization 20 that may originate from repetitive regions of the genome such as retrotransposons or regions normally organized into heterochromatin and which are normally derived exclusively from the antisense strand of double stranded RNA 21 piRNAs have thus been classified as repeat associated small interfering RNAs rasiRNAs 22 Although their biogenesis is not yet well understood piRNAs and Piwi proteins are thought to form an endogenous system for silencing the expression of selfish genetic elements such as retrotransposons and thus preventing the gene products of such sequences from interfering with germ cell formation 21 23 Footnotes edit The word Wimpy itself comes from White color of eyes and Impotency or profound infertility Y is added as a suffix to form a more pronounceable and coherent word 5 References edit a b Rivas FV Tolia NH Song JJ et al April 2005 Purified Argonaute2 and an siRNA form recombinant human RISC Nat Struct Mol Biol 12 4 340 9 doi 10 1038 nsmb918 PMID 15800637 S2CID 2021813 Uniprot The Universal knowledge database Nucleic Acids Research 45 D1 D158 D169 2017 doi 10 1093 nar gkw1099 PMC 5210571 PMID 27899622 Lindse K 2013 Piwi RNAs the Defenders of the Genome a href Template Cite journal html title Template Cite journal cite journal a Cite journal requires journal help Cox DN Chao A Lin H 2000 piwi encodes a nucleoplasmic factor whose activity modulates the number and division rate of germline stem cells Development 127 3 503 14 doi 10 1242 dev 127 3 503 PMID 10631171 Ross RJ Weiner MM Lin H 2014 PIWI proteins and PIWI interacting RNAs in the soma Nature 505 7483 353 359 doi 10 1038 nature12987 PMC 4265809 PMID 24429634 Lin H Spradling AC 1997 A novel group of pumilio mutations affects the asymmetric division of germline stem cells in the Drosophila ovary Development 124 12 2463 2476 doi 10 1242 dev 124 12 2463 PMID 9199372 Cox DN Chao A Baker J Chang L Qiao D Lin H 1998 A novel class of evolutionarily conserved genes defined by piwi are essential for stem cell self renewal Genes Dev 12 23 3715 27 doi 10 1101 gad 12 23 3715 PMC 317255 PMID 9851978 Darricarrere N Liu N Watanabe T Lin H 2013 Function of Piwi a nuclear Piwi Argonaute protein is independent of its slicer activity Proc Natl Acad Sci USA 110 6 1297 1302 Bibcode 2013PNAS 110 1297D doi 10 1073 pnas 1213283110 PMC 3557079 PMID 23297219 a b Zeng Lei Zhang Qiang Yan Kelley Zhou Ming Ming 2011 06 01 Structural insights into piRNA recognition by the human PIWI like 1 PAZ domain Proteins Structure Function and Bioinformatics 79 6 2004 2009 doi 10 1002 prot 23003 ISSN 1097 0134 PMC 3092821 PMID 21465557 Wei Kai Fa Wu Ling Juan Chen Juan Chen Yan feng Xie Dao Xin August 2012 Structural evolution and functional diversification analyses of argonaute protein Journal of Cellular Biochemistry 113 8 2576 2585 doi 10 1002 jcb 24133 ISSN 1097 4644 PMID 22415963 S2CID 25990631 Tian Y Simanshu D Ma J Patel D 2010 Structural basis for piRNA 2 O methylated 3 end recognition by Piwi PAZ Piwi Argonaute Zwille domains Proc Natl Acad Sci USA 108 3 903 910 doi 10 1073 pnas 1017762108 PMC 3024652 PMID 21193640 Qiao D Zeeman AM Deng W Looijenga LH Lin H 2002 Molecular characterization of hiwi a human member of the piwi gene family whose overexpression is correlated to seminomas Oncogene 21 25 3988 99 doi 10 1038 sj onc 1205505 PMID 12037681 Deng W Lin H 2002 miwi a murine homolog of piwi encodes a cytoplasmic protein essential for spermatogenesis Dev Cell 2 6 819 30 doi 10 1016 s1534 5807 02 00165 x PMID 12062093 Mani S Juliano C 2013 Untangling the Web The Diverse Functions of the PIWI piRNA Pathway Mol Reprod Dev 80 8 632 664 doi 10 1002 mrd 22195 PMC 4234069 PMID 23712694 Thomson T Lin H 2009 The Biogenesis and Function PIWI Proteins and piRNAs Progress and Prospect Annu Rev Cell Dev Biol 25 355 376 doi 10 1146 annurev cellbio 24 110707 175327 PMC 2780330 PMID 19575643 Cerutti L Mian N Bateman A October 2000 Domains in gene silencing and cell differentiation proteins the novel PAZ domain and redefinition of the Piwi domain Trends Biochem Sci 25 10 481 2 doi 10 1016 S0968 0004 00 01641 8 PMID 11050429 Ma J Yuan Y Meister G Pei Y Tuschl T Patel D 2005 Structural basis for 5 end specific recognition of guide RNA by the A fulgidus Piwi protein Nature 434 7033 666 70 Bibcode 2005Natur 434 666M doi 10 1038 nature03514 PMC 4694588 PMID 15800629 Megosh HB Cox DN Campbell C Lin H 2006 The role of PIWI and the miRNA machinery in Drosophila germline determination Curr Biol 16 19 1884 94 doi 10 1016 j cub 2006 08 051 PMID 16949822 S2CID 6397874 Kim VN 2006 Small RNAs just got bigger Piwi interacting RNAs piRNAs in mammalian testes Genes Dev 20 15 1993 7 doi 10 1101 gad 1456106 PMID 16882976 Girard A Sachidanandam R Hannon GJ Carmell MA 2006 A germline specific class of small RNAs binds mammalian Piwi proteins Nature 442 7099 199 202 Bibcode 2006Natur 442 199G doi 10 1038 nature04917 PMID 16751776 S2CID 3185036 a b Vagin VV Sigova A Li C Seitz H Gvozdev V Zamore PD 2006 A distinct small RNA pathway silences selfish genetic elements in the germline Science 313 5785 320 4 Bibcode 2006Sci 313 320V doi 10 1126 science 1129333 PMID 16809489 S2CID 40471466 Saito K Nishida KM Mori T Kawamura Y Miyoshi K Nagami T Siomi H Siomi MC 2006 Specific association of Piwi with rasiRNAs derived from retrotransposon and heterochromatic regions in the Drosophila genome Genes Dev 20 16 2214 22 doi 10 1101 gad 1454806 PMC 1553205 PMID 16882972 Ozata DM Gainetdinov I Zoch A Phillip D Zamore PD 2019 PIWI interacting RNAs small RNAs with big functions PDF Nature Reviews Genetics 20 2 89 108 doi 10 1038 s41576 018 0073 3 PMID 30446728 S2CID 53565676 External links editSCOP 110640 Piwi domain in SCOP PDOC50822 Piwi domain in PROSITE UNIPROT Piwi Piwi domains Retrieved from https en wikipedia org w index php title Piwi amp oldid 1220610644, wikipedia, wiki, book, books, library,

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