226:
different transposons, including multiple
Gypsies), all facing the same direction. Indeed, piRNAs are all found in clusters throughout animal genomes; these clusters may contain as few as ten or many thousands of piRNAs matching different, phased transposon fragments. This led to the idea in 2007 that in germlines a pool of primary piRNAs is processed from long single-stranded transcripts encoded by piRNA clusters in the opposite orientation of the transposons, so that the piRNAs can anneal to and complement the transposon-encoded transcripts, thereby triggering their degradation. Any transposon landing in the correct orientation in such a cluster will make the individual more or less immune to that transposon, and such an advantageous mutation will spread quickly through the population. The original mutations in the flamenco locus inhibited the transcription of the master transcript, thereby deactivating this defense system.
459:
Aub and Ago3 target cleavage triggers the 'phased' loading of piRNA into Piwi. Phasing begins with the targeting and cleavage of a complementary target by either Aub or Ago3 associated with a 'responder' piRNA. Once cleaved, the targeted transcript is then processed further by a mechanism believed to require the mitochondrial-associated endonuclease, Zucchini, which leads to the loading of Piwi protein with sequential fragments of the targeted transcript. In this way, the Aub or Ago3 'responder' piRNA sequence cleaves a complementary target that is then sliced at periodic intervals of approximately 27 nucleotides that are sequentially loaded into Piwi protein. Once loaded with piRNA, Piwi then enters the germ cell nucleus to co-transcriptionally silence nascent transcripts with complementarity to its piRNA guide. It is currently unknown whether phasing occurs in other organisms.
414:
the main factor in targeting deleterious transcripts through complementarity. Conversely, Ago3 piRNA sequences are predominantly of sense orientation to transposable element transcripts and are derived from the product of Aub cleavage of transposon mRNA. As such, Ago3 piRNA lack the ability to target transposable element transcripts directly. Therefore, it was proposed that Ago3 piRNA guide the production of piRNA that are loaded into Aub by targeting newly exported piRNA cluster transcripts. Several lines of evidence support the effect of Ago3 on the production of Aub piRNA, in particular from examining the piRNA repertoire in
4702:
315:
438:) was also shown to interact with both Aub and Ago3 through its Tudor domains while also binding itself through its N-terminal Krimper domain. Specifically, Krimper interacts with Ago3 in its piRNA-unloaded state, while its interaction with Aub is dependent on the symmetrical dimethylation of arginine residues in the N-terminal region of Aub. In Silkmoth germ cells, it was proposed that
306:, a species of yeast, as well in some plants, neither of which have been observed to contain the Piwi subfamily of Argonaute proteins. It has been observed that both rasiRNA and piRNA are maternally linked, but more specifically it is the Piwi protein subfamily that is maternally linked and therefore leads to the observation that rasiRNA and piRNA are maternally linked.
734:-based methods have been developed in response to this difficulty. However, research has also revealed that a number of annotated piRNAs may be false positives; for instance, a majority of piRNAs that were expressed in somatic non-gonadal tissues were considered to derive from non-coding RNA fragments.
458:
piRNA pathway can be separated into two branches: the cytoplasmic branch consisting of Aub and Ago3 operating the Ping-Pong mechanism, and the nuclear branch, pertaining to the co-transcriptional silencing of genomic loci by Piwi in the nucleus. Through complementary strategies, two studies show that
413:
or in mouse remains to be understood, but a leading hypothesis is that the interaction between Aub and Ago3 allows for a cyclic refinement of piRNA that are best suited to target active transposon sequences. Aub piRNA are primarily antisense to transposable element transcripts and are believed to be
589:. However, in mosquitoes the PIWI family of proteins has expanded and some PIWI proteins have been identified as antiviral such as Piwi4. As such virus infections in mosquitoes commonly produce virus-derived piRNAs in diverse positive-sense RNA, negative-sense RNA and single-stranded DNA viruses.
408:
where the piRNA associated with the two cytoplasmic Piwi proteins, Aubergine (Aub) and
Argonaute-3 (Ago3) exhibited a high frequency of sequence complementarity over exactly 10 nucleotides at their 5′ ends. This relationship is known as the "ping-pong signature" and is also observed in associated
299:
and some unicellular eukaryotes but its presence in mammals has not been determined, unlike piRNA which has been observed in many species of invertebrates and vertebrates including mammals; however, since proteins which associate with rasiRNA are found in both vertebrates and invertebrates, it is
225:
in the germline and by 2003 the idea had emerged that vestiges of transposons might produce dsRNAs required for the silencing of "live" transposons. Sequencing of the 200,000-bp flamenco locus was difficult, as it turned out to be packed with transposable element fragments (104 insertions of 42
237:
genome in the mid-20th century, and, through interbreeding, within decades all wild fruit flies worldwide (though not the reproductively isolated lab strains) contained the same P-element. Repression of further P-element activity, spreading near-simultaneously, appears to have occurred by the
1924:
Aravin A, Gaidatzis D, Pfeffer S, Lagos-Quintana M, Landgraf P, Iovino N, Morris P, Brownstein MJ, Kuramochi-Miyagawa S, Nakano T, Chien M, Russo JJ, Ju J, Sheridan R, Sander C, Zavolan M, Tuschl T (July 2006). "A novel class of small RNAs bind to MILI protein in mouse testes".
632:
that are known to bind symmetrically dimethylated arginine residues (sDMA) present in methylation motifs of Piwi proteins. Piwi proteins are symmetrically dimethylated by the PRMT5 methylosome complex, consisting of Valois (MEP50) and Capsulèen (dart5; PRMT5).
729:
platform sequencing. These techniques allow analysis of highly complex and heterogeneous RNA populations like piRNAs. Due to their small size, expression and amplification of small RNAs can be challenging, so specialised
538:. Three piwi subfamily proteins – MIWI, MIWI2, and MILI – have been found to be essential for spermatogenesis in mice. piRNAs direct the piwi proteins to their transposon targets. A decrease or absence of PIWI
395:
across species. Ping-pong signatures have been identified in very primitive animals such as sponges and cnidarians, pointing to the existence of the ping-pong cycle already in the early branches of metazoans.
542:
is correlated with an increased expression of transposons. Transposons have a high potential to cause deleterious effects on their hosts and, in fact, mutations in piRNA pathways have been found to reduce
487:
to transposon sequences, suggesting that transposons are targets of the piRNAs. In mammals, it appears that the activity of piRNAs in transposon silencing is most important during the development of the
445:
It is likely that the mechanism of Ping-Pong is primarily coordinated by
Krimper but factors such as Kumo/Qin and Vasa, in addition to other factors have necessary functions in the Ping-Pong mechanism.
1420:
Houwing S, Kamminga LM, Berezikov E, Cronembold D, Girard A, van den Elst H, et al. (April 2007). "A role for Piwi and piRNAs in germ cell maintenance and transposon silencing in
Zebrafish".
1289:
Carmen L, Michela B, Rosaria V, Gabriella M (2009). "Existence of snoRNA, microRNA, piRNA characteristics in a novel non-coding RNA: x-ncRNA and its biological implication in Homo sapiens".
1603:
Das PP, Bagijn MP, Goldstein LD, Woolford JR, Lehrbach NJ, Sapetschnig A, Buhecha HR, Gilchrist MJ, Howe KL, Stark R, Matthews N, Berezikov E, Ketting RF, Tavaré S, Miska EA (July 2008).
3463:
Anne J, Mechler BM (May 2005). "Valois, a component of the nuage and pole plasm, is involved in assembly of these structures, and binds to Tudor and the methyltransferase Capsuléen".
330:
only, and this may indicate that they are the product of long single stranded precursor molecules. A primary processing pathway is suggested to be the only pathway used to produce
2462:
Li C, Vagin VV, Lee S, Xu J, Ma S, Xi H, Seitz H, Horwich MD, Syrzycka M, Honda BM, Kittler EL, Zapp ML, Klattenhoff C, Schulz N, Theurkauf WE, Weng Z, Zamore PD (May 2009).
3304:"Aedes Anphevirus: an Insect-Specific Virus Distributed Worldwide in Aedes aegypti Mosquitoes That Has Complex Interplays with Wolbachia and Dengue Virus Infection in Cells"
326:
of piRNAs is not yet fully understood, although possible mechanisms have been proposed. piRNAs show a significant strand bias, that is, they are derived from one strand of
3827:"Identification and characterization of two novel classes of small RNAs in the mouse germline: retrotransposon-derived siRNAs in oocytes and germline small RNAs in testes"
613:
Genetic screens examining fertility defects identified a number of proteins that are not Piwi-clade
Argonautes, yet produce the same sterility phenotypes as Piwi mutants.
115:
98:
in 2001. By 2008, it was still unclear how piRNAs are generated, but potential methods had been suggested, and it was certain their biogenesis pathway is distinct from
295:. Unlike the Ago subfamily which is present in animals, plants, and fission yeast, the Piwi subfamily has only been found in animals. RasiRNA has been observed in
358:
at the tenth position. Since the piRNA involved in the ping pong cycle directs its attacks on transposon transcripts, the ping pong cycle acts only at the level of
3353:"Density-dependent enhanced replication of a densovirus in Wolbachia-infected Aedes cells is associated with production of piRNAs and higher virus-derived siRNAs"
2656:
Xiol J, Spinelli P, Laussmann MA, Homolka D, Yang Z, Cora E, Couté Y, Conn S, Kadlec J, Sachidanandam R, Kaksonen M, Cusack S, Ephrussi A, Pillai RS (June 2014).
2078:, Plasterk RH, Hannon GJ, Draper BW, Ketting RF (April 2007). "A role for Piwi and piRNAs in germ cell maintenance and transposon silencing in Zebrafish".
173:. This 3’ modification is a 2’-O-methylation; the reason for this modification is not clear, but it has been suggested that it increases piRNA stability.
605:
effects. The activity of specific piRNAs in the epigenetic process also requires interactions between piwi proteins and HP1a, as well as other factors.
1360:
Vagin VV, Sigova A, Li C, Seitz H, Gvozdev V, Zamore PD (July 2006). "A distinct small RNA pathway silences selfish genetic elements in the germline".
3657:
Lau NC, Seto AG, Kim J, Kuramochi-Miyagawa S, Nakano T, Bartel DP, Kingston RE (July 2006). "Characterization of the piRNA complex from rat testes".
585:
ovarian somatic sheet (OSS) cells. Subsequent experimental studies have demonstrated that the piRNA pathway is not required for antiviral defence in
3400:
3288:
3223:
3157:
3092:
3015:
354:
from the 5’ end of the primary piRNA, producing the secondary piRNA. These secondary piRNAs are targeted toward sequences that possess an
1605:"Piwi and piRNAs act upstream of an endogenous siRNA pathway to suppress Tc3 transposon mobility in the Caenorhabditis elegans germline"
3737:
Girard A, Sachidanandam R, Hannon GJ, Carmell MA (July 2006). "A germline-specific class of small RNAs binds mammalian Piwi proteins".
2131:
Girard A, Sachidanandam R, Hannon GJ, Carmell MA (July 2006). "A germline-specific class of small RNAs binds mammalian Piwi proteins".
1976:
Tam OH, Aravin AA, Stein P, Girard A, Murchison EP, Cheloufi S, Hodges E, Anger M, Sachidanandam R, Schultz RM, Hannon GJ (May 2008).
844:"Computational Identification of piRNAs Using Features Based on RNA Sequence, Structure, Thermodynamic and Physicochemical Properties"
155:
have a 5’ monophosphate and a 3’ modification that acts to block either the 2’ or 3’ oxygen; this has also been confirmed to exist in
2905:
342:. Also proposed is a ‘Ping Pong’ mechanism wherein primary piRNAs recognise their complementary targets and cause the recruitment of
79:(miRNA) in size (26–31 nucleotides as opposed to 21–24 nt), lack of sequence conservation, increased complexity, and independence of
4297:
991:"Drosophila rasiRNA pathway mutations disrupt embryonic axis specification through activation of an ATR/Chk2 DNA damage response"
20:
4035:
1184:"Double-stranded RNA-mediated silencing of genomic tandem repeats and transposable elements in the D. melanogaster germline"
425:
was reported to coordinate the loading of Ago3 with piRNA, in addition to interacting with both Aub and Ago3. However, the
268:, although they only seem to be required in males. In invertebrates, piRNAs have been detected in both the male and female
3414:
Kirino Y, Kim N, de
Planell-Saguer M, Khandros E, Chiorean S, Klein PS, Rigoutsos I, Jongens TA, Mourelatos Z (May 2009).
942:"Delving into the diversity of silencing pathways. Symposium on MicroRNAs and siRNAs: biological functions and mechanisms"
64:
and other spurious or repeat-derived transcripts, but can also be involved in the regulation of other genetic elements in
471:
function across species contributes to the difficulty in establishing the functionality of piRNAs. However, like other
143:
4383:
57:
2348:
Grimson A, Srivastava M, Fahey B, Woodcroft BJ, Chiang HR, King N, Degnan BM, Rokhsar DS, Bartel DP (October 2008).
205:
4670:
4141:
894:
Siomi MC, Sato K, Pezic D, Aravin AA (April 2011). "PIWI-interacting small RNAs: the vanguard of genome defence".
4675:
4069:
300:
possible that active rasiRNA exist and have yet to be observed in other animals. RasiRNAs have been observed in
3606:"Non-coding RNA fragments account for the majority of annotated piRNAs expressed in somatic non-gonadal tissues"
2795:
Le Thomas A, Rogers AK, Webster A, Marinov GK, Liao SE, Perkins EM, Hur JK, Aravin AA, Tóth KF (February 2013).
2297:
1035:"History of the discovery of a master locus producing piRNAs: the flamenco/COM locus in Drosophila melanogaster"
283:, suggesting that piRNA pathways may function in both of these areas and, therefore, may have multiple effects.
134:
and modes of action do vary somewhat between species, a number of features are conserved. piRNAs have no clear
1128:
Xie Z, Johansen LK, Gustafson AM, Kasschau KD, Lellis AD, Zilberman D, Jacobsen SE, Carrington JC (May 2004).
4572:
4458:
2568:"Aub and Ago3 Are Recruited to Nuage through Two Mechanisms to Form a Ping-Pong Complex Assembled by Krimper"
2242:
Aravin AA, Sachidanandam R, Bourc'his D, Schaefer C, Pezic D, Toth KF, Bestor T, Hannon GJ (September 2008).
2182:
Tomari Y, Du T, Haley B, Schwarz DS, Bennett R, Cook HA, Koppetsch BS, Theurkauf WE, Zamore PD (March 2004).
1571:
Lin H, Yin H, Beyret E, Findley S, Deng W (2008). "The role of the piRNA pathway in stem cell self-renewal".
515:
1473:
Kirino Y, Mourelatos Z (April 2007). "Mouse Piwi-interacting RNAs are 2′-O-methylated at their 3′ termini".
2699:"Noncoding RNA. piRNA-guided slicing specifies transcripts for Zucchini-dependent, phased piRNA biogenesis"
2074:
Houwing S, Kamminga LM, Berezikov E, Cronembold D, Girard A, van den Elst H, Filippov DV, Blaser H, Raz E,
421:
The molecular mechanism that underpins Ping-Pong likely involves several piRNA pathway associated factors.
71:
piRNAs are mostly created from loci that function as transposon traps which provide a kind of RNA-mediated
3996:
4685:
4680:
4665:
4290:
4028:
2797:"Piwi induces piRNA-guided transcriptional silencing and establishment of a repressive chromatin state"
2748:"Noncoding RNA. piRNA-guided transposon cleavage initiates Zucchini-dependent, phased piRNA production"
2514:
Zhang Z, Xu J, Koppetsch BS, Wang J, Tipping C, Ma S, Weng Z, Theurkauf WE, Zamore PD (November 2011).
722:
573:
which are required to recognise and silence transposons, but this relationship is not well understood.
3173:
2405:
Gunawardane LS, Saito K, Nishida KM, Miyoshi K, Kawamura Y, Nagami T, Siomi H, Siomi MC (March 2007).
1313:"Large-scale sequencing reveals 21U-RNAs and additional microRNAs and endogenous siRNAs in C. elegans"
4008:
731:
558:(endo-siRNA) may have comparable and even redundant functionality in transposon control in mammalian
409:
piRNA from Mili and Miwi2 proteins isolated from mouse testes. The proposed function of Ping-Pong in
302:
2615:
Sato K, Iwasaki YW, Shibuya A, Carninci P, Tsuchizawa Y, Ishizu H, Siomi MC, Siomi H (August 2015).
4254:
3868:
Carmell MA, Girard A, van de Kant HJ, Bourc'his D, Bestor TH, de Rooij DG, Hannon GJ (April 2007).
1826:"Reexamining the P-Element Invasion of Drosophila melanogaster Through the Lens of piRNA Silencing"
581:
In
Dipterans viral-derived piRNAs derived from positive-sense RNA viruses were first identified in
359:
335:
3416:"Arginine methylation of Piwi proteins catalysed by dPRMT5 is required for Ago3 and Aub stability"
4731:
3870:"MIWI2 is essential for spermatogenesis and repression of transposons in the mouse male germline"
2913:
989:
Klattenhoff C, Bratu DP, McGinnis-Schultz N, Koppetsch BS, Cook HA, Theurkauf WE (January 2007).
385:
contain adenine at their tenth position, and this has been interpreted as possible evidence of a
381:
193:
157:
4705:
4398:
4283:
4079:
2244:"A piRNA pathway primed by individual transposons is linked to de novo DNA methylation in mice"
494:
368:
151:
2617:"Krimper Enforces an Antisense Bias on piRNA Pools by Binding AGO3 in the Drosophila Germline"
2566:
Webster A, Li S, Hur JK, Wachsmuth M, Bois JS, Perkins EM, Patel DJ, Aravin AA (August 2015).
2298:"Discrete small RNA-generating loci as master regulators of transposon activity in Drosophila"
4582:
4426:
4171:
4021:
3394:
3282:
3217:
3151:
3086:
3028:
3009:
555:
372:, for instance, does have piRNAs, but does not appear to use the ping pong mechanism at all.
176:
More than 50,000 unique piRNA sequences have been discovered in mice and more than 13,000 in
3825:
Watanabe T, Takeda A, Tsukiyama T, Mise K, Okuno T, Sasaki H, Minami N, Imai H (July 2006).
3174:
Varjak M, Maringer K, Watson M, Sreenu VB, Fredericks AC, Pondeville E; et al. (2017).
4617:
4600:
3746:
3666:
3046:
2969:
2860:
2516:"Heterotypic piRNA Ping-Pong requires qin, a protein with both E3 ligase and Tudor domains"
2418:
2361:
2296:
Brennecke J, Aravin AA, Stark A, Dus M, Kellis M, Sachidanandam R, Hannon GJ (March 2007).
2140:
1989:
1934:
1672:
1369:
1195:
61:
2464:"Collapse of germline piRNAs in the absence of Argonaute3 reveals somatic piRNAs in flies"
2100:
1875:"proTRAC—a software for probabilistic piRNA cluster detection, visualization and analysis"
1442:
221:
proposed that double-stranded (ds) RNA-mediated silencing is implicated in the control of
8:
4612:
4595:
4453:
4336:
4151:
4100:
392:
200:
138:
motifs, due to the fact that the length of a piRNA varies between species (from 21 to 31
135:
3750:
3670:
3050:
2973:
2864:
2422:
2407:"A slicer-mediated mechanism for repeat-associated siRNA 5′ end formation in Drosophila"
2365:
2144:
1993:
1938:
1676:
1659:
Brennecke J, Malone CD, Aravin AA, Sachidanandam R, Stark A, Hannon GJ (November 2008).
1373:
1311:
Ruby JG, Jan C, Player C, Axtell MJ, Lee W, Nusbaum C, Ge H, Bartel DP (December 2006).
1199:
180:. It is thought that there are many hundreds of thousands of different piRNA species in
4506:
4501:
4376:
4181:
4176:
3931:
3904:
3851:
3826:
3808:
3783:
3770:
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3382:
3328:
3303:
3265:
3238:
3200:
3175:
3134:
3107:
3069:
3030:
2992:
2958:"Virus discovery by deep sequencing and assembly of virus-derived small silencing RNAs"
2957:
2938:
2881:
2848:
2821:
2796:
2772:
2747:
2723:
2698:
2592:
2567:
2540:
2515:
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2349:
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2213:
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2113:
2010:
1977:
1958:
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1801:
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1629:
1604:
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1498:
1455:
1393:
1342:
1221:
1061:
1034:
966:
941:
919:
868:
843:
726:
386:
3108:"Comparative genomics of small RNA regulatory pathway components in vector mosquitoes"
2200:
2183:
1208:
1183:
1156:
1129:
859:
4590:
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3333:
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2545:
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2436:
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2322:
2273:
2205:
2156:
2105:
2056:
2015:
1978:"Pseudogene-derived small interfering RNAs regulate gene expression in mouse oocytes"
1950:
1906:
1855:
1806:
1750:
1698:
1634:
1550:
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1447:
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1334:
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1012:
971:
911:
873:
821:
566:
72:
3694:
3386:
3176:"Aedes aegypti Piwi4 Is a Noncanonical PIWI Protein Involved in Antiviral Responses"
2942:
2658:"RNA clamping by Vasa assembles a piRNA amplifier complex on transposon transcripts"
2448:
2217:
2117:
1502:
1459:
1397:
1346:
1225:
4416:
4005:– a software for probabilistic piRNA cluster detection, visualization, and analysis
3961:
3926:
3916:
3881:
3846:
3838:
3803:
3795:
3774:
3754:
3715:
3674:
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2005:
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1007:
990:
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539:
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222:
3704:"Small RNAs just got bigger: Piwi-interacting RNAs (piRNAs) in mammalian testes"
3369:
3352:
1841:
1584:
1182:
Aravin AA, Naumova NM, Tulin AV, Vagin VV, Rozovsky YM, Gvozdev VA (July 2001).
213:. The site of the mutations that made these Gypsies "dance" was thus called the
4632:
4535:
4352:
3990:
3572:
3523:
2674:
2657:
2479:
2350:"Early origins and evolution of microRNAs and Piwi-interacting RNAs in animals"
2317:
2091:
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1536:
1433:
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778:
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242:
34:
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4136:
4064:
4059:
3191:
2075:
1089:"Identification and characterization of small RNAs involved in RNA silencing"
1051:
957:
721:
Major advances in the study of piRNA have been achieved thanks to the use of
511:
484:
442:
protein coordinates the Ping-Pong mechanism of
Silkmoth Aub (Siwi) and Ago3.
426:
347:
291:
There are at least three
Argonaute (Ago) subfamilies that have been found in
3678:
3124:
3059:
2982:
2763:
2714:
2431:
2406:
1684:
1661:"An epigenetic role for maternally inherited piRNAs in transposon silencing"
1381:
249:
and vertebrate piRNAs have been located in areas lacking any protein-coding
4726:
4650:
4470:
4443:
4388:
4307:
4264:
4074:
3975:
3940:
3895:
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3541:
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3143:
3078:
3001:
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1702:
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1389:
1338:
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1217:
1165:
1114:
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1016:
975:
915:
877:
825:
629:
535:
389:
276:
127:
3993:– a software for finding ping-pong signatures and ping-pong cycle activity
4562:
4408:
4275:
3319:
988:
570:
434:
90:
Double-stranded RNAs capable of silencing repeat elements, then known as
53:
4363:
3842:
3799:
3758:
3720:
3703:
2373:
2241:
2152:
2001:
1946:
1130:"Genetic and functional diversification of small RNA pathways in plants"
418:
ovaries that are mutant for Ago3 and the Tudor-domain protein Kumo/Qin.
4622:
4331:
4326:
4321:
4228:
4202:
3966:
3949:
2955:
1266:
1249:
602:
552:
480:
351:
323:
139:
131:
123:
3476:
3350:
3105:
3029:
Petit M, Mongelli V, Frangeul L, Blanc H, Jiggins F, Saleh MC (2016).
4486:
4259:
4233:
4115:
4105:
3905:"To be or not to be a piRNA: genomic origin and processing of piRNAs"
3867:
2849:"A C. elegans Piwi, PRG-1, regulates 21U-RNAs during spermatogenesis"
1486:
682:
544:
531:
527:
523:
472:
439:
376:
331:
292:
280:
230:
162:
149:
is common to piRNAs in both vertebrates and invertebrates. piRNAs in
65:
49:
3431:
1923:
907:
318:
The ping-pong mechanism for the biogenesis of the 5′ end of rasiRNA.
192:
In the early 1980s, it was discovered that a single mutation in the
75:
against transposon expansions and invasions. They are distinct from
4640:
4491:
4421:
4156:
3921:
3508:"A PCR-based method for detection and quantification of small RNAs"
2035:"Tiny RNA: Where do we come from? What are we? Where are we going?"
269:
210:
76:
3413:
2073:
1658:
1419:
4645:
4166:
4110:
939:
526:. These are active in the testes of mammals and are required for
363:
355:
339:
314:
265:
229:
A historical example of invasion and Piwi response is known: the
146:
41:
3736:
2130:
4660:
4655:
4605:
4238:
4146:
4013:
3239:"PIWIs Go Viral: Arbovirus-Derived piRNAs in Vector Mosquitoes"
3236:
2347:
940:
Dorner S, Eulalio A, Huntzinger E, Izaurralde E (August 2007).
597:
piRNAs can be transmitted maternally, and based on research in
559:
489:
404:
The piRNA Ping-Pong pathway was first proposed from studies in
261:
196:
181:
2904:
Ozata DM, Gainetdinov I, Zoch A, Phillip D, Zamore PD (2019).
2404:
2184:"RISC assembly defects in the Drosophila RNAi mutant armitage"
672:
275:
At the cellular level, piRNAs have been found within both the
4463:
4120:
2956:
Wu Q, Luo Y, Lu R, Lau N, Lai EC, Li WX; et al. (2010).
2794:
1288:
362:. One or both of these mechanisms may be acting in different
250:
103:
99:
80:
3656:
2655:
1127:
691:
3824:
2903:
1721:"Mighty Piwis defend the germline against genome intruders"
519:
468:
343:
166:
84:
52:
proteins. These piRNA complexes are mostly involved in the
45:
3784:"A novel class of small RNAs in mouse spermatogenic cells"
3555:
Tang F, Hayashi K, Kaneda M, Lao K, Surani MA (May 2008).
2614:
1602:
608:
522:
proteins that are part of a family of proteins called the
241:
piRNA clusters in genomes can now readily be detected via
118:
Proposed piRNA structure, with the 3′ end 2′-O-methylation
16:
Largest class of small non-coding RNA molecules in animals
4525:
4346:
4045:
1032:
327:
170:
37:
3506:
Ro S, Park C, Jin J, Sanders KM, Yan W (December 2006).
3031:"piRNA pathway is not required for antiviral defense in
114:
2295:
1181:
2906:"PIWI-interacting RNAs: small RNAs with big functions"
106:, while rasiRNA is now considered a piRNA subspecies.
19:"piRNA" redirects here. For the software package, see
2565:
40:
molecules expressed in animal cells. piRNAs form RNA-
3554:
2746:
Han BW, Wang W, Li C, Weng Z, Zamore PD (May 2015).
2696:
2513:
2181:
1975:
1247:
893:
3999:– a web resource on classified and clustered piRNAs
3902:
3781:
3561:
3512:
Biochemical and Biophysical Research Communications
1570:
1310:
616:
3557:"A sensitive multiplex assay for piRNA expression"
1872:
1718:
1359:
257:have been identified amidst protein-coding genes.
3603:
3301:
1518:
338:resulting in piRNAs with a tendency to target 5’
4718:
3903:Le Thomas A, Tóth KF, Aravin AA (January 2014).
3782:Grivna ST, Beyret E, Wang Z, Lin H (July 2006).
3505:
2561:
2559:
1472:
1033:Goriaux C, Théron E, Brasset E, Vaury C (2014).
334:piRNAs; in this mechanism, piRNA precursors are
3351:Parry R, Bishop C, De Hayr L, Asgari S (2019).
3169:
3167:
3106:Campbell CL, Black WC, Hess AM, Foy BD (2008).
2509:
2507:
1291:Journal of Bioinformatics and Sequence Analysis
797:
624:Many factors required for the piRNA pathway in
601:, piRNAs may be involved in maternally derived
1243:
1241:
1239:
1237:
1235:
889:
887:
4291:
4029:
2745:
2690:
2556:
2291:
2289:
2287:
1250:"Biogenesis and germline functions of piRNAs"
841:
375:A significant number of piRNAs identified in
3399:: CS1 maint: multiple names: authors list (
3344:
3295:
3287:: CS1 maint: multiple names: authors list (
3230:
3222:: CS1 maint: multiple names: authors list (
3164:
3156:: CS1 maint: multiple names: authors list (
3099:
3091:: CS1 maint: multiple names: authors list (
3022:
3014:: CS1 maint: multiple names: authors list (
2949:
2897:
2504:
2461:
2175:
2124:
2067:
1774:
1654:
1652:
1650:
1648:
1086:
199:could specifically activate all copies of a
83:for biogenesis, at least in animals. (Plant
3604:Tosar JP, Rovira C, Cayota A (2018-01-22).
3462:
2697:Mohn F, Handler D, Brennecke J (May 2015).
2398:
1714:
1712:
1282:
1248:Klattenhoff C, Theurkauf W (January 2008).
1232:
982:
884:
756:
754:
752:
750:
748:
746:
87:may play a role in rasi/piRNA biogenesis.)
4305:
4298:
4284:
4036:
4022:
3947:
2846:
2649:
2284:
2237:
2235:
2233:
2231:
2229:
2227:
1866:
1514:
1512:
1415:
1413:
1411:
1409:
1407:
798:Seto AG, Kingston RE, Lau NC (June 2007).
467:The wide variation in piRNA sequences and
3965:
3930:
3920:
3885:
3850:
3807:
3719:
3629:
3580:
3531:
3439:
3368:
3327:
3264:
3254:
3199:
3133:
3123:
3068:
3058:
2991:
2981:
2880:
2820:
2771:
2739:
2722:
2673:
2632:
2608:
2591:
2539:
2487:
2430:
2381:
2316:
2267:
2199:
2099:
2050:
2009:
1900:
1890:
1873:Rosenkranz D, Zischler H (January 2012).
1849:
1800:
1770:
1768:
1766:
1764:
1744:
1692:
1645:
1628:
1544:
1519:Faehnle CR, Joshua-Tor L (October 2007).
1475:Nature Structural & Molecular Biology
1441:
1328:
1265:
1207:
1177:
1175:
1155:
1145:
1104:
1060:
1050:
1028:
1026:
1006:
965:
867:
815:
777:
3237:Miesen P, Joosten J, van Rij RP (2016).
2788:
2455:
1823:
1709:
935:
933:
743:
551:. Further, it is thought that piRNA and
313:
113:
2842:
2840:
2341:
2224:
1777:"Small RNAs as guardians of the genome"
1509:
1404:
1306:
1304:
837:
835:
609:Accessory proteins of the piRNA pathway
475:, piRNAs are thought to be involved in
92:repeat associated small interfering RNA
21:Partition function for Interacting RNAs
4719:
2032:
2026:
1917:
1775:Malone CD, Hannon GJ (February 2009).
1761:
1598:
1596:
1594:
1566:
1564:
1466:
1172:
1082:
1080:
1023:
793:
791:
789:
4279:
4017:
3950:"piRNAs: from biogenesis to function"
3948:Weick EM, Miska EA (September 2014).
1719:O'Donnell KA, Boeke JD (April 2007).
1353:
930:
896:Nature Reviews Molecular Cell Biology
842:Monga I, Banerjee I (November 2019).
800:"The coming of age for Piwi proteins"
725:techniques, such as Solexa, 454, and
592:
498:and humans, piRNAs are necessary for
260:In mammals, piRNAs are found both in
3548:
2837:
1521:"Argonautes confront new small RNAs"
1301:
832:
576:
122:piRNAs have been identified in both
44:complexes through interactions with
3701:
1969:
1591:
1561:
1525:Current Opinion in Chemical Biology
1087:Aravin A, Tuschl T (October 2005).
1077:
786:
565:piRNAs appear to affect particular
187:
13:
3649:
3499:
109:
14:
4743:
4422:Micro
3984:
860:10.2174/1389202920666191129112705
350:of the transcript at a point ten
286:
4701:
4700:
4142:Cis-natural antisense transcript
4043:
716:
505:
479:, specifically the silencing of
449:
33:) is the largest class of small
4377:precursor, heterogenous nuclear
4070:Signal recognition particle RNA
3597:
3456:
3407:
1817:
4507:Trans-acting small interfering
4471:Enhancer RNAs
4389:Transfer
2847:Wang G, Reinke V (June 2008).
2101:11858/00-001M-0000-0012-E169-6
1443:11858/00-001M-0000-0012-E169-6
1121:
695:nuclear piRNA pathway proteins
346:proteins. This results in the
238:Piwi-interacting RNA pathway.
1:
4394:Ribosomal
4372:Messenger
2201:10.1016/S0092-8674(04)00218-1
2052:10.1016/j.tplants.2008.05.005
1209:10.1016/S0960-9822(01)00299-8
1106:10.1016/j.febslet.2005.08.009
737:
518:(RISC). piRNAs interact with
516:RNA-induced silencing complex
483:. The majority of piRNAs are
309:
3887:10.1016/j.devcel.2007.03.001
3256:10.1371/journal.ppat.1006017
2634:10.1016/j.molcel.2015.06.024
2584:10.1016/j.molcel.2015.07.017
2532:10.1016/j.molcel.2011.10.011
2260:10.1016/j.molcel.2008.09.003
1621:10.1016/j.molcel.2008.06.003
1147:10.1371/journal.pbio.0020104
1008:10.1016/j.devcel.2006.12.001
817:10.1016/j.molcel.2007.05.021
399:
94:(rasiRNA), were proposed in
7:
3370:10.1016/j.virol.2018.12.006
1842:10.1534/genetics.115.184119
1824:Kelleher ES (August 2016).
1585:10.1016/j.ydbio.2008.05.048
462:
10:
4748:
4573:Multicopy single-stranded
4417:Interferential
4229:Reverse transcribing virus
3573:10.1016/j.bbrc.2008.03.035
3524:10.1016/j.bbrc.2006.10.105
3302:Parry R, Asgari S (2018).
2675:10.1016/j.cell.2014.05.018
2480:10.1016/j.cell.2009.04.027
2318:10.1016/j.cell.2007.01.043
2092:10.1016/j.cell.2007.03.026
1793:10.1016/j.cell.2009.01.045
1737:10.1016/j.cell.2007.03.028
1537:10.1016/j.cbpa.2007.08.032
1434:10.1016/j.cell.2007.03.026
1330:10.1016/j.cell.2006.10.040
779:10.1016/j.cell.2006.07.012
762:"Molecular Biology Select"
723:next-generation sequencing
18:
4696:
4631:
4581:
4524:
4487:Guide
4479:
4407:
4362:
4345:
4314:
4247:
4216:
4195:
4129:
4088:
4052:
3622:10.1038/s42003-017-0001-7
2927:10.1038/s41576-018-0073-3
2873:10.1016/j.cub.2008.05.009
772:(2): 223–225. July 2006.
303:Schizosaccharomyces pombe
4449:Small nuclear
4009:piRNA cluster – database
3192:10.1128/mSphere.00144-17
3039:Proc Natl Acad Sci U S A
2962:Proc Natl Acad Sci U S A
1052:10.3389/fgene.2014.00257
958:10.1038/sj.embor.7401015
514:via the formation of an
4563:Genomic
4196:Cis-regulatory elements
4167:Repeat-associated siRNA
3831:Genes & Development
3788:Genes & Development
3708:Genes & Development
3679:10.1126/science.1130164
3125:10.1186/1471-2164-9-425
3060:10.1073/pnas.1607952113
3033:Drosophila melanogaster
2983:10.1073/pnas.0911353107
2914:Nature Reviews Genetics
2801:Genes & Development
2764:10.1126/science.aaa1264
2715:10.1126/science.aaa1039
2432:10.1126/science.1140494
2039:Trends in Plant Science
1685:10.1126/science.1165171
1382:10.1126/science.1129333
587:Drosophila melanogaster
235:Drosophila melanogaster
201:retrovirus-like element
158:Drosophila melanogaster
4666:Artificial chromosomes
4454:Small nucleolar
4080:Transfer-messenger RNA
3702:Kim VN (August 2006).
3610:Communications Biology
2813:10.1101/gad.209841.112
2033:Ruvkun G (July 2008).
1892:10.1186/1471-2105-13-5
677:piRNA pathway proteins
319:
152:Caenorhabditis elegans
142:), and the bias for a
119:
4459:Small Cajal Body RNAs
4172:Small interfering RNA
1573:Developmental Biology
1039:Frontiers in Genetics
620:Tudor domain proteins
556:small interfering RNA
317:
233:transposon invaded a
117:
62:transposable elements
4512:Subgenomic messenger
4427:Small interfering
4399:Transfer-messenger
4162:Piwi-interacting RNA
3320:10.1128/JVI.00224-18
665:Brother of Yb (BoYB)
510:piRNA has a role in
58:post-transcriptional
27:Piwi-interacting RNA
4101:Small nucleolar RNA
3843:10.1101/gad.1425706
3800:10.1101/gad.1434406
3759:10.1038/nature04917
3751:2006Natur.442..199G
3721:10.1101/gad.1456106
3671:2006Sci...313..363L
3051:2016PNAS..113E4218P
2974:2010PNAS..107.1606W
2865:2008CBio...18..861W
2423:2007Sci...315.1587G
2417:(5818): 1587–1590.
2374:10.1038/nature07415
2366:2008Natur.455.1193G
2360:(7217): 1193–1197.
2153:10.1038/nature04917
2145:2006Natur.442..199G
2002:10.1038/nature06904
1994:2008Natur.453..534T
1947:10.1038/nature04916
1939:2006Natur.442..203A
1677:2008Sci...322.1387B
1671:(5906): 1387–1392.
1374:2006Sci...313..320V
1200:2001CBio...11.1017A
668:Sister of Yb (SoYB)
136:secondary structure
4541:Chloroplast
4384:modified Messenger
4347:Ribonucleic acids
4182:Trans-acting siRNA
4177:Small temporal RNA
4152:Long noncoding RNA
3967:10.1242/dev.094037
3874:Developmental Cell
1879:BMC Bioinformatics
1267:10.1242/dev.006486
995:Developmental Cell
593:Epigenetic effects
567:methyltransferases
320:
217:. In 2001, Aravin
120:
4714:
4713:
4591:Xeno
4553:Complementary
4526:Deoxyribonucleic
4520:
4519:
4497:Small hairpin
4273:
4272:
4187:Short hairpin RNA
4096:Small nuclear RNA
4053:Protein synthesis
3960:(18): 3458–3471.
3837:(13): 1732–1743.
3794:(13): 1709–1714.
3745:(7099): 199–202.
3714:(15): 1993–1997.
3665:(5785): 363–367.
3477:10.1242/dev.01809
2758:(6236): 817–821.
2709:(6236): 812–817.
2139:(7099): 199–202.
1988:(7194): 534–538.
1933:(7099): 203–207.
1368:(5785): 320–324.
1194:(13): 1017–1027.
1099:(26): 5830–5840.
577:Antiviral effects
569:that perform the
73:adaptive immunity
4739:
4704:
4703:
4681:Yeast
4502:Small temporal
4432:Piwi-interacting
4360:
4359:
4356:
4337:Deoxynucleotides
4300:
4293:
4286:
4277:
4276:
4038:
4031:
4024:
4015:
4014:
3979:
3969:
3944:
3934:
3924:
3899:
3889:
3864:
3854:
3821:
3811:
3778:
3733:
3723:
3698:
3644:
3643:
3633:
3601:
3595:
3594:
3584:
3567:(4): 1190–1194.
3552:
3546:
3545:
3535:
3503:
3497:
3496:
3460:
3454:
3453:
3443:
3411:
3405:
3404:
3398:
3390:
3372:
3348:
3342:
3341:
3331:
3299:
3293:
3292:
3286:
3278:
3268:
3258:
3249:(12): e1006017.
3234:
3228:
3227:
3221:
3213:
3203:
3171:
3162:
3161:
3155:
3147:
3137:
3127:
3103:
3097:
3096:
3090:
3082:
3072:
3062:
3045:(29): E4218-27.
3026:
3020:
3019:
3013:
3005:
2995:
2985:
2953:
2947:
2946:
2910:
2901:
2895:
2894:
2884:
2844:
2835:
2834:
2824:
2792:
2786:
2785:
2775:
2743:
2737:
2736:
2726:
2694:
2688:
2687:
2677:
2668:(7): 1698–1711.
2653:
2647:
2646:
2636:
2612:
2606:
2605:
2595:
2563:
2554:
2553:
2543:
2511:
2502:
2501:
2491:
2459:
2453:
2452:
2434:
2402:
2396:
2395:
2385:
2345:
2339:
2338:
2320:
2311:(6): 1089–1103.
2302:
2293:
2282:
2281:
2271:
2239:
2222:
2221:
2203:
2179:
2173:
2172:
2128:
2122:
2121:
2103:
2071:
2065:
2064:
2054:
2030:
2024:
2023:
2013:
1973:
1967:
1966:
1921:
1915:
1914:
1904:
1894:
1870:
1864:
1863:
1853:
1836:(4): 1513–1531.
1821:
1815:
1814:
1804:
1772:
1759:
1758:
1748:
1716:
1707:
1706:
1696:
1656:
1643:
1642:
1632:
1600:
1589:
1588:
1568:
1559:
1558:
1548:
1516:
1507:
1506:
1487:10.1038/nsmb1218
1470:
1464:
1463:
1445:
1417:
1402:
1401:
1357:
1351:
1350:
1332:
1323:(6): 1193–1207.
1308:
1299:
1298:
1286:
1280:
1279:
1269:
1245:
1230:
1229:
1211:
1179:
1170:
1169:
1159:
1149:
1125:
1119:
1118:
1108:
1084:
1075:
1074:
1064:
1054:
1030:
1021:
1020:
1010:
986:
980:
979:
969:
937:
928:
927:
891:
882:
881:
871:
848:Current Genomics
839:
830:
829:
819:
795:
784:
783:
781:
758:
709:SetDB1 (Eggless)
687:Maelstrom (Mael)
643:Spindle-E (SpnE)
437:
223:retrotransposons
188:History and loci
4747:
4746:
4742:
4741:
4740:
4738:
4737:
4736:
4717:
4716:
4715:
4710:
4692:
4633:Cloning vectors
4627:
4613:Locked
4577:
4527:
4516:
4475:
4403:
4350:
4349:
4341:
4310:
4304:
4274:
4269:
4243:
4224:Retrotransposon
4212:
4191:
4130:Gene regulation
4125:
4084:
4048:
4042:
3987:
3982:
3652:
3650:Further reading
3647:
3602:
3598:
3553:
3549:
3504:
3500:
3461:
3457:
3432:10.1038/ncb1872
3412:
3408:
3392:
3391:
3349:
3345:
3300:
3296:
3280:
3279:
3235:
3231:
3215:
3214:
3172:
3165:
3149:
3148:
3104:
3100:
3084:
3083:
3027:
3023:
3007:
3006:
2954:
2950:
2908:
2902:
2898:
2859:(12): 861–867.
2853:Current Biology
2845:
2838:
2793:
2789:
2744:
2740:
2695:
2691:
2654:
2650:
2613:
2609:
2564:
2557:
2512:
2505:
2460:
2456:
2403:
2399:
2346:
2342:
2300:
2294:
2285:
2240:
2225:
2180:
2176:
2129:
2125:
2072:
2068:
2031:
2027:
1974:
1970:
1922:
1918:
1871:
1867:
1822:
1818:
1773:
1762:
1717:
1710:
1657:
1646:
1601:
1592:
1569:
1562:
1517:
1510:
1471:
1467:
1418:
1405:
1358:
1354:
1309:
1302:
1287:
1283:
1246:
1233:
1188:Current Biology
1180:
1173:
1126:
1122:
1085:
1078:
1031:
1024:
987:
983:
938:
931:
908:10.1038/nrm3089
892:
885:
840:
833:
796:
787:
760:
759:
744:
740:
719:
697:
679:
622:
611:
599:D. melanogaster
595:
579:
549:D. melanogaster
540:gene expression
534:development in
508:
500:spermatogenesis
465:
452:
433:
402:
382:D. melanogaster
312:
289:
247:D. melanogaster
245:methods. While
190:
178:D. melanogaster
130:, and although
112:
110:Characteristics
24:
17:
12:
11:
5:
4745:
4735:
4734:
4732:Non-coding RNA
4729:
4712:
4711:
4709:
4708:
4697:
4694:
4693:
4691:
4690:
4689:
4688:
4683:
4678:
4673:
4663:
4658:
4653:
4648:
4643:
4637:
4635:
4629:
4628:
4626:
4625:
4620:
4618:Peptide
4615:
4610:
4609:
4608:
4603:
4598:
4596:Glycol
4587:
4585:
4579:
4578:
4576:
4575:
4570:
4565:
4560:
4555:
4550:
4549:
4548:
4543:
4532:
4530:
4522:
4521:
4518:
4517:
4515:
4514:
4509:
4504:
4499:
4494:
4489:
4483:
4481:
4477:
4476:
4474:
4473:
4468:
4467:
4466:
4461:
4456:
4451:
4441:
4436:
4435:
4434:
4429:
4424:
4413:
4411:
4405:
4404:
4402:
4401:
4396:
4391:
4386:
4381:
4380:
4379:
4368:
4366:
4357:
4343:
4342:
4340:
4339:
4334:
4329:
4324:
4318:
4316:
4312:
4311:
4308:nucleic acids
4303:
4302:
4295:
4288:
4280:
4271:
4270:
4268:
4267:
4262:
4257:
4255:Telomerase RNA
4251:
4249:
4245:
4244:
4242:
4241:
4236:
4231:
4226:
4220:
4218:
4214:
4213:
4211:
4210:
4205:
4199:
4197:
4193:
4192:
4190:
4189:
4184:
4179:
4174:
4169:
4164:
4159:
4154:
4149:
4144:
4139:
4133:
4131:
4127:
4126:
4124:
4123:
4118:
4113:
4108:
4103:
4098:
4092:
4090:
4089:RNA processing
4086:
4085:
4083:
4082:
4077:
4072:
4067:
4062:
4056:
4054:
4050:
4049:
4041:
4040:
4033:
4026:
4018:
4012:
4011:
4006:
4000:
3994:
3986:
3985:External links
3983:
3981:
3980:
3945:
3922:10.1186/gb4154
3909:Genome Biology
3900:
3880:(4): 503–514.
3865:
3822:
3779:
3734:
3699:
3653:
3651:
3648:
3646:
3645:
3596:
3547:
3518:(3): 756–763.
3498:
3471:(9): 2167–77.
3455:
3420:Nat. Cell Biol
3406:
3343:
3294:
3229:
3163:
3098:
3021:
2968:(4): 1606–11.
2948:
2896:
2836:
2807:(4): 390–399.
2787:
2738:
2689:
2648:
2627:(4): 553–563.
2621:Molecular Cell
2607:
2578:(4): 564–575.
2572:Molecular Cell
2555:
2526:(4): 572–584.
2520:Molecular Cell
2503:
2474:(3): 509–521.
2454:
2397:
2340:
2283:
2254:(6): 785–799.
2248:Molecular Cell
2223:
2194:(6): 831–841.
2174:
2123:
2066:
2045:(7): 313–316.
2025:
1968:
1916:
1865:
1816:
1787:(4): 656–668.
1760:
1708:
1644:
1609:Molecular Cell
1590:
1560:
1531:(5): 569–577.
1508:
1481:(4): 347–348.
1465:
1403:
1352:
1300:
1281:
1231:
1171:
1120:
1076:
1022:
981:
952:(8): 723–729.
929:
902:(4): 246–258.
883:
854:(7): 508–518.
831:
810:(5): 603–609.
804:Molecular Cell
785:
741:
739:
736:
718:
715:
714:
713:
710:
707:
704:
701:
696:
690:
689:
688:
685:
678:
671:
670:
669:
666:
663:
656:
653:
652:Vreteno (Vret)
650:
647:
644:
641:
638:
621:
615:
610:
607:
594:
591:
578:
575:
507:
504:
492:, and in both
477:gene silencing
464:
461:
451:
448:
401:
398:
311:
308:
288:
287:Classification
285:
243:bioinformatics
215:flamenco locus
209:in the female
189:
186:
111:
108:
15:
9:
6:
4:
3:
2:
4744:
4733:
4730:
4728:
4725:
4724:
4722:
4707:
4699:
4698:
4695:
4687:
4684:
4682:
4679:
4677:
4674:
4672:
4669:
4668:
4667:
4664:
4662:
4659:
4657:
4654:
4652:
4649:
4647:
4644:
4642:
4639:
4638:
4636:
4634:
4630:
4624:
4621:
4619:
4616:
4614:
4611:
4607:
4604:
4602:
4601:Threose
4599:
4597:
4594:
4593:
4592:
4589:
4588:
4586:
4584:
4580:
4574:
4571:
4569:
4566:
4564:
4561:
4559:
4558:Deoxyribozyme
4556:
4554:
4551:
4547:
4546:Mitochondrial
4544:
4542:
4539:
4538:
4537:
4534:
4533:
4531:
4529:
4523:
4513:
4510:
4508:
4505:
4503:
4500:
4498:
4495:
4493:
4490:
4488:
4485:
4484:
4482:
4478:
4472:
4469:
4465:
4462:
4460:
4457:
4455:
4452:
4450:
4447:
4446:
4445:
4442:
4440:
4437:
4433:
4430:
4428:
4425:
4423:
4420:
4419:
4418:
4415:
4414:
4412:
4410:
4406:
4400:
4397:
4395:
4392:
4390:
4387:
4385:
4382:
4378:
4375:
4374:
4373:
4370:
4369:
4367:
4365:
4364:Translational
4361:
4358:
4354:
4348:
4344:
4338:
4335:
4333:
4330:
4328:
4325:
4323:
4320:
4319:
4317:
4313:
4309:
4301:
4296:
4294:
4289:
4287:
4282:
4281:
4278:
4266:
4263:
4261:
4258:
4256:
4253:
4252:
4250:
4246:
4240:
4237:
4235:
4232:
4230:
4227:
4225:
4222:
4221:
4219:
4215:
4209:
4208:SECIS element
4206:
4204:
4201:
4200:
4198:
4194:
4188:
4185:
4183:
4180:
4178:
4175:
4173:
4170:
4168:
4165:
4163:
4160:
4158:
4155:
4153:
4150:
4148:
4145:
4143:
4140:
4138:
4137:Antisense RNA
4135:
4134:
4132:
4128:
4122:
4119:
4117:
4114:
4112:
4109:
4107:
4104:
4102:
4099:
4097:
4094:
4093:
4091:
4087:
4081:
4078:
4076:
4073:
4071:
4068:
4066:
4065:Ribosomal RNA
4063:
4061:
4060:Messenger RNA
4058:
4057:
4055:
4051:
4047:
4039:
4034:
4032:
4027:
4025:
4020:
4019:
4016:
4010:
4007:
4004:
4001:
3998:
3995:
3992:
3989:
3988:
3977:
3973:
3968:
3963:
3959:
3955:
3951:
3946:
3942:
3938:
3933:
3928:
3923:
3918:
3914:
3910:
3906:
3901:
3897:
3893:
3888:
3883:
3879:
3875:
3871:
3866:
3862:
3858:
3853:
3848:
3844:
3840:
3836:
3832:
3828:
3823:
3819:
3815:
3810:
3805:
3801:
3797:
3793:
3789:
3785:
3780:
3776:
3772:
3768:
3764:
3760:
3756:
3752:
3748:
3744:
3740:
3735:
3731:
3727:
3722:
3717:
3713:
3709:
3705:
3700:
3696:
3692:
3688:
3684:
3680:
3676:
3672:
3668:
3664:
3660:
3655:
3654:
3641:
3637:
3632:
3627:
3623:
3619:
3615:
3611:
3607:
3600:
3592:
3588:
3583:
3578:
3574:
3570:
3566:
3562:
3558:
3551:
3543:
3539:
3534:
3529:
3525:
3521:
3517:
3513:
3509:
3502:
3494:
3490:
3486:
3482:
3478:
3474:
3470:
3466:
3459:
3451:
3447:
3442:
3437:
3433:
3429:
3425:
3421:
3417:
3410:
3402:
3396:
3388:
3384:
3380:
3376:
3371:
3366:
3362:
3358:
3354:
3347:
3339:
3335:
3330:
3325:
3321:
3317:
3313:
3309:
3305:
3298:
3290:
3284:
3276:
3272:
3267:
3262:
3257:
3252:
3248:
3244:
3240:
3233:
3225:
3219:
3211:
3207:
3202:
3197:
3193:
3189:
3185:
3181:
3177:
3170:
3168:
3159:
3153:
3145:
3141:
3136:
3131:
3126:
3121:
3117:
3113:
3109:
3102:
3094:
3088:
3080:
3076:
3071:
3066:
3061:
3056:
3052:
3048:
3044:
3040:
3036:
3034:
3025:
3017:
3011:
3003:
2999:
2994:
2989:
2984:
2979:
2975:
2971:
2967:
2963:
2959:
2952:
2944:
2940:
2936:
2932:
2928:
2924:
2921:(2): 89–108.
2920:
2916:
2915:
2907:
2900:
2892:
2888:
2883:
2878:
2874:
2870:
2866:
2862:
2858:
2854:
2850:
2843:
2841:
2832:
2828:
2823:
2818:
2814:
2810:
2806:
2802:
2798:
2791:
2783:
2779:
2774:
2769:
2765:
2761:
2757:
2753:
2749:
2742:
2734:
2730:
2725:
2720:
2716:
2712:
2708:
2704:
2700:
2693:
2685:
2681:
2676:
2671:
2667:
2663:
2659:
2652:
2644:
2640:
2635:
2630:
2626:
2622:
2618:
2611:
2603:
2599:
2594:
2589:
2585:
2581:
2577:
2573:
2569:
2562:
2560:
2551:
2547:
2542:
2537:
2533:
2529:
2525:
2521:
2517:
2510:
2508:
2499:
2495:
2490:
2485:
2481:
2477:
2473:
2469:
2465:
2458:
2450:
2446:
2442:
2438:
2433:
2428:
2424:
2420:
2416:
2412:
2408:
2401:
2393:
2389:
2384:
2379:
2375:
2371:
2367:
2363:
2359:
2355:
2351:
2344:
2336:
2332:
2328:
2324:
2319:
2314:
2310:
2306:
2299:
2292:
2290:
2288:
2279:
2275:
2270:
2265:
2261:
2257:
2253:
2249:
2245:
2238:
2236:
2234:
2232:
2230:
2228:
2219:
2215:
2211:
2207:
2202:
2197:
2193:
2189:
2185:
2178:
2170:
2166:
2162:
2158:
2154:
2150:
2146:
2142:
2138:
2134:
2127:
2119:
2115:
2111:
2107:
2102:
2097:
2093:
2089:
2085:
2081:
2077:
2070:
2062:
2058:
2053:
2048:
2044:
2040:
2036:
2029:
2021:
2017:
2012:
2007:
2003:
1999:
1995:
1991:
1987:
1983:
1979:
1972:
1964:
1960:
1956:
1952:
1948:
1944:
1940:
1936:
1932:
1928:
1920:
1912:
1908:
1903:
1898:
1893:
1888:
1884:
1880:
1876:
1869:
1861:
1857:
1852:
1847:
1843:
1839:
1835:
1831:
1827:
1820:
1812:
1808:
1803:
1798:
1794:
1790:
1786:
1782:
1778:
1771:
1769:
1767:
1765:
1756:
1752:
1747:
1742:
1738:
1734:
1730:
1726:
1722:
1715:
1713:
1704:
1700:
1695:
1690:
1686:
1682:
1678:
1674:
1670:
1666:
1662:
1655:
1653:
1651:
1649:
1640:
1636:
1631:
1626:
1622:
1618:
1614:
1610:
1606:
1599:
1597:
1595:
1586:
1582:
1578:
1574:
1567:
1565:
1556:
1552:
1547:
1542:
1538:
1534:
1530:
1526:
1522:
1515:
1513:
1504:
1500:
1496:
1492:
1488:
1484:
1480:
1476:
1469:
1461:
1457:
1453:
1449:
1444:
1439:
1435:
1431:
1427:
1423:
1416:
1414:
1412:
1410:
1408:
1399:
1395:
1391:
1387:
1383:
1379:
1375:
1371:
1367:
1363:
1356:
1348:
1344:
1340:
1336:
1331:
1326:
1322:
1318:
1314:
1307:
1305:
1297:(2): 031–040.
1296:
1292:
1285:
1277:
1273:
1268:
1263:
1259:
1255:
1251:
1244:
1242:
1240:
1238:
1236:
1227:
1223:
1219:
1215:
1210:
1205:
1201:
1197:
1193:
1189:
1185:
1178:
1176:
1167:
1163:
1158:
1153:
1148:
1143:
1139:
1135:
1131:
1124:
1116:
1112:
1107:
1102:
1098:
1094:
1090:
1083:
1081:
1072:
1068:
1063:
1058:
1053:
1048:
1044:
1040:
1036:
1029:
1027:
1018:
1014:
1009:
1004:
1000:
996:
992:
985:
977:
973:
968:
963:
959:
955:
951:
947:
943:
936:
934:
925:
921:
917:
913:
909:
905:
901:
897:
890:
888:
879:
875:
870:
865:
861:
857:
853:
849:
845:
838:
836:
827:
823:
818:
813:
809:
805:
801:
794:
792:
790:
780:
775:
771:
767:
763:
757:
755:
753:
751:
749:
747:
742:
735:
733:
728:
724:
717:Investigation
711:
708:
705:
702:
699:
698:
694:
686:
684:
681:
680:
676:
667:
664:
661:
657:
654:
651:
648:
645:
642:
639:
636:
635:
634:
631:
630:Tudor domains
627:
619:
614:
606:
604:
600:
590:
588:
584:
574:
572:
568:
563:
561:
557:
554:
550:
546:
541:
537:
536:invertebrates
533:
529:
525:
521:
517:
513:
512:RNA silencing
506:RNA silencing
503:
501:
497:
496:
491:
486:
482:
478:
474:
470:
460:
457:
450:piRNA Phasing
447:
443:
441:
436:
431:
428:
427:Tudor protein
424:
419:
417:
412:
407:
397:
394:
391:
388:
384:
383:
378:
373:
371:
370:
365:
361:
360:transcription
357:
353:
349:
345:
341:
337:
333:
329:
325:
316:
307:
305:
304:
298:
294:
284:
282:
278:
273:
271:
267:
263:
258:
256:
252:
248:
244:
239:
236:
232:
227:
224:
220:
216:
212:
208:
207:
202:
198:
195:
185:
183:
179:
174:
172:
168:
164:
160:
159:
154:
153:
148:
145:
141:
137:
133:
129:
128:invertebrates
125:
116:
107:
105:
101:
97:
93:
88:
86:
82:
78:
74:
69:
67:
63:
60:silencing of
59:
55:
51:
47:
43:
39:
36:
32:
28:
22:
4676:Bacterial
4651:Lambda phage
4431:
4315:Constituents
4265:List of RNAs
4161:
4075:Transfer RNA
3957:
3953:
3912:
3908:
3877:
3873:
3834:
3830:
3791:
3787:
3742:
3738:
3711:
3707:
3662:
3658:
3613:
3609:
3599:
3564:
3560:
3550:
3515:
3511:
3501:
3468:
3464:
3458:
3426:(5): 652–8.
3423:
3419:
3409:
3395:cite journal
3360:
3356:
3346:
3311:
3307:
3297:
3283:cite journal
3246:
3242:
3232:
3218:cite journal
3183:
3179:
3152:cite journal
3115:
3112:BMC Genomics
3111:
3101:
3087:cite journal
3042:
3038:
3032:
3024:
3010:cite journal
2965:
2961:
2951:
2918:
2912:
2899:
2856:
2852:
2804:
2800:
2790:
2755:
2751:
2741:
2706:
2702:
2692:
2665:
2661:
2651:
2624:
2620:
2610:
2575:
2571:
2523:
2519:
2471:
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2343:
2308:
2304:
2251:
2247:
2191:
2187:
2177:
2136:
2132:
2126:
2086:(1): 69–82.
2083:
2079:
2069:
2042:
2038:
2028:
1985:
1981:
1971:
1930:
1926:
1919:
1882:
1878:
1868:
1833:
1829:
1819:
1784:
1780:
1731:(1): 37–44.
1728:
1724:
1668:
1664:
1615:(1): 79–90.
1612:
1608:
1576:
1572:
1528:
1524:
1478:
1474:
1468:
1428:(1): 69–82.
1425:
1421:
1365:
1361:
1355:
1320:
1316:
1294:
1290:
1284:
1257:
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1191:
1187:
1137:
1134:PLOS Biology
1133:
1123:
1096:
1093:FEBS Letters
1092:
1042:
1038:
1001:(1): 45–55.
998:
994:
984:
949:
946:EMBO Reports
945:
899:
895:
851:
847:
807:
803:
769:
765:
720:
700:Rhino (HP1D)
692:
674:
659:
625:
623:
617:
612:
598:
596:
586:
582:
580:
571:methylations
564:
548:
509:
493:
466:
455:
453:
444:
429:
422:
420:
415:
410:
405:
403:
390:biosynthetic
380:
374:
367:
321:
301:
296:
290:
274:
259:
254:
253:, piRNAs in
246:
240:
234:
228:
218:
214:
204:
191:
177:
175:
156:
150:
121:
95:
91:
89:
70:
30:
26:
25:
4671:P1-derived
4439:Antisense
4332:Nucleotides
4327:Nucleosides
4322:Nucleobases
3991:PingPongPro
3954:Development
3465:Development
3243:PLOS Pathog
1254:Development
1140:(5): E104.
649:Tejas (Tej)
637:Tudor (Tud)
481:transposons
352:nucleotides
336:transcribed
140:nucleotides
124:vertebrates
48:-subfamily
4721:Categories
4623:Morpholino
4536:Organellar
4444:Processual
4409:Regulatory
4353:non-coding
4203:Riboswitch
4147:CRISPR RNA
3997:piRNA Bank
3915:(1): 204.
3363:: 89–100.
1579:(2): 479.
1260:(1): 3–9.
738:References
693:Drosophila
675:Drosophila
673:Non-Tudor
626:Drosophila
618:Drosophila
603:epigenetic
583:Drosophila
553:endogenous
524:Argonautes
495:C. elegans
473:small RNAs
456:Drosophila
416:Drosophila
411:Drosophila
406:Drosophila
369:C. elegans
324:biogenesis
310:Biogenesis
297:Drosophila
293:eukaryotes
255:C. elegans
132:biogenesis
96:Drosophila
54:epigenetic
35:non-coding
4583:Analogues
4568:Hachimoji
4351:(coding,
4306:Types of
4260:Vault RNA
4234:RNA virus
4217:Parasites
4116:RNase MRP
4106:Guide RNA
4044:Types of
545:fertility
532:stem-cell
528:germ-cell
485:antisense
400:Ping Pong
393:mechanism
387:conserved
377:zebrafish
332:pachytene
281:cytoplasm
270:germlines
231:P-element
194:fruit fly
163:zebrafish
66:germ line
50:Argonaute
4706:Category
4641:Phagemid
4492:Ribozyme
4157:MicroRNA
3976:25183868
3941:24467990
3896:17395546
3861:16766679
3818:16766680
3767:16751776
3730:16882976
3695:21150160
3687:16778019
3640:30271890
3616:(1): 2.
3591:18348866
3542:17084816
3485:15800004
3450:19377467
3387:58572380
3379:30583288
3357:Virology
3338:29950416
3275:28033427
3210:28497119
3144:18801182
3079:27357659
3002:20080648
2943:53565676
2935:30446728
2891:18501605
2831:23392610
2782:25977554
2733:25977553
2684:24910301
2643:26212455
2602:26295961
2550:22099305
2498:19395009
2449:11513777
2441:17322028
2392:18830242
2327:17346786
2278:18922463
2218:17588448
2210:15035985
2161:16751776
2118:13373509
2110:17418787
2076:Moens CB
2061:18562240
2020:18404147
1955:16751777
1911:22233380
1885:(5): 5.
1860:27516614
1830:Genetics
1811:19239887
1755:17418784
1703:19039138
1639:18571451
1555:17928262
1503:31193964
1495:17384647
1460:13373509
1452:17418787
1398:40471466
1390:16809489
1347:16838469
1339:17174894
1276:18032451
1226:14767819
1218:11470406
1166:15024409
1115:16153643
1071:25136352
1017:17199040
976:17599087
916:21427766
878:32655289
826:17560367
727:Illumina
712:SuVar3–9
703:Deadlock
640:Qin/Kumo
628:contain
463:Function
348:cleavage
340:uridines
211:germline
77:microRNA
4646:Plasmid
4111:RNase P
4003:proTRAC
3932:4053809
3852:1522070
3809:1522066
3775:3185036
3747:Bibcode
3667:Bibcode
3659:Science
3631:6052916
3582:3855189
3533:1934510
3493:6810484
3441:2746449
3329:6096813
3308:J Virol
3266:5198996
3201:5415634
3180:mSphere
3135:2566310
3118:: 425.
3070:4961201
3047:Bibcode
2993:2824396
2970:Bibcode
2882:2494713
2861:Bibcode
2822:3589556
2773:4545291
2752:Science
2724:4988486
2703:Science
2593:4545750
2541:3236501
2489:2768572
2419:Bibcode
2411:Science
2383:3837422
2362:Bibcode
2335:2246942
2269:2730041
2169:3185036
2141:Bibcode
2011:2981145
1990:Bibcode
1963:4379895
1935:Bibcode
1902:3293768
1851:4981261
1802:2792755
1746:4122227
1694:2805124
1673:Bibcode
1665:Science
1630:3353317
1546:2077831
1370:Bibcode
1362:Science
1196:Bibcode
1062:4120762
1045:: 257.
967:1978081
924:5710813
869:7327968
660:fs(1)Yb
646:Krimper
560:oocytes
430:krimper
364:species
356:adenine
277:nucleus
266:ovaries
203:called
182:mammals
147:uridine
68:cells.
42:protein
4661:Fosmid
4656:Cosmid
4606:Hexose
4528:acids
4480:Others
4239:Viroid
3974:
3939:
3929:
3894:
3859:
3849:
3816:
3806:
3773:
3765:
3739:Nature
3728:
3693:
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3638:
3628:
3589:
3579:
3540:
3530:
3491:
3483:
3448:
3438:
3385:
3377:
3336:
3326:
3314:(17).
3273:
3263:
3208:
3198:
3142:
3132:
3077:
3067:
3000:
2990:
2941:
2933:
2889:
2879:
2829:
2819:
2780:
2770:
2731:
2721:
2682:
2641:
2600:
2590:
2548:
2538:
2496:
2486:
2447:
2439:
2390:
2380:
2354:Nature
2333:
2325:
2276:
2266:
2216:
2208:
2167:
2159:
2133:Nature
2116:
2108:
2059:
2018:
2008:
1982:Nature
1961:
1953:
1927:Nature
1909:
1899:
1858:
1848:
1809:
1799:
1753:
1743:
1701:
1691:
1637:
1627:
1553:
1543:
1501:
1493:
1458:
1450:
1396:
1388:
1345:
1337:
1274:
1224:
1216:
1164:
1157:350667
1154:
1113:
1069:
1059:
1015:
974:
964:
922:
914:
876:
866:
824:
706:Cutoff
490:embryo
435:A1ZAC4
262:testes
219:et al.
197:genome
169:, and
4686:Human
4464:Y RNA
4248:Other
4121:Y RNA
3771:S2CID
3691:S2CID
3489:S2CID
3383:S2CID
3186:(3).
2939:S2CID
2909:(PDF)
2445:S2CID
2331:S2CID
2301:(PDF)
2214:S2CID
2165:S2CID
2114:S2CID
1959:S2CID
1499:S2CID
1456:S2CID
1394:S2CID
1343:S2CID
1222:S2CID
920:S2CID
251:genes
206:Gypsy
104:siRNA
100:miRNA
81:Dicer
31:piRNA
3972:PMID
3937:PMID
3892:PMID
3857:PMID
3814:PMID
3763:PMID
3726:PMID
3683:PMID
3636:PMID
3587:PMID
3538:PMID
3481:PMID
3446:PMID
3401:link
3375:PMID
3334:PMID
3289:link
3271:PMID
3224:link
3206:PMID
3158:link
3140:PMID
3093:link
3075:PMID
3016:link
2998:PMID
2931:PMID
2887:PMID
2827:PMID
2778:PMID
2729:PMID
2680:PMID
2662:Cell
2639:PMID
2598:PMID
2546:PMID
2494:PMID
2468:Cell
2437:PMID
2388:PMID
2323:PMID
2305:Cell
2274:PMID
2206:PMID
2188:Cell
2157:PMID
2106:PMID
2080:Cell
2057:PMID
2016:PMID
1951:PMID
1907:PMID
1856:PMID
1807:PMID
1781:Cell
1751:PMID
1725:Cell
1699:PMID
1635:PMID
1551:PMID
1491:PMID
1448:PMID
1422:Cell
1386:PMID
1335:PMID
1317:Cell
1272:PMID
1214:PMID
1162:PMID
1111:PMID
1067:PMID
1013:PMID
972:PMID
912:PMID
874:PMID
822:PMID
766:Cell
683:Vasa
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530:and
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469:piwi
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