237:
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.
470:
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.
425:
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
4713:
449:) 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
326:
317:, 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.
745:-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.
469:
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
424:
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
600:. 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.
419:
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
310:
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
236:
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
248:
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
1935:
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".
643:
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).
740:
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
549:. 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
406:
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.
553:
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
498:
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
456:
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.
1431:
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".
1300:
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".
1614:
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).
3474:
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".
341:
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
2473:
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).
3315:"Aedes Anphevirus: an Insect-Specific Virus Distributed Worldwide in Aedes aegypti Mosquitoes That Has Complex Interplays with Wolbachia and Dengue Virus Infection in Cells"
337:
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
3838:"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"
624:
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.
109:
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
3411:
3299:
3234:
3168:
3103:
3026:
306:. 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
369:
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
3364:"Density-dependent enhanced replication of a densovirus in Wolbachia-infected Aedes cells is associated with production of piRNAs and higher virus-derived siRNAs"
2667:
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).
2089:, 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".
184:. 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.
616:
effects. The activity of specific piRNAs in the epigenetic process also requires interactions between piwi proteins and HP1a, as well as other factors.
1371:
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".
3668:
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".
596:
ovarian somatic sheet (OSS) cells. Subsequent experimental studies have demonstrated that the piRNA pathway is not required for antiviral defence in
4308:
31:
4046:
365:
from the 5’ end of the primary piRNA, producing the secondary piRNA. These secondary piRNAs are targeted toward sequences that possess an
1616:"Piwi and piRNAs act upstream of an endogenous siRNA pathway to suppress Tc3 transposon mobility in the Caenorhabditis elegans germline"
3748:
Girard A, Sachidanandam R, Hannon GJ, Carmell MA (July 2006). "A germline-specific class of small RNAs binds mammalian Piwi proteins".
2142:
Girard A, Sachidanandam R, Hannon GJ, Carmell MA (July 2006). "A germline-specific class of small RNAs binds mammalian Piwi proteins".
1987:
Tam OH, Aravin AA, Stein P, Girard A, Murchison EP, Cheloufi S, Hodges E, Anger M, Sachidanandam R, Schultz RM, Hannon GJ (May 2008).
855:"Computational Identification of piRNAs Using Features Based on RNA Sequence, Structure, Thermodynamic and Physicochemical Properties"
166:
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
154:
2916:
353:. Also proposed is a ‘Ping Pong’ mechanism wherein primary piRNAs recognise their complementary targets and cause the recruitment of
90:(miRNA) in size (26–31 nucleotides as opposed to 21–24 nt), lack of sequence conservation, increased complexity, and independence of
4394:
68:
4681:
4152:
1002:"Drosophila rasiRNA pathway mutations disrupt embryonic axis specification through activation of an ATR/Chk2 DNA damage response"
4686:
4080:
126:
1195:"Double-stranded RNA-mediated silencing of genomic tandem repeats and transposable elements in the D. melanogaster germline"
436:
was reported to coordinate the loading of Ago3 with piRNA, in addition to interacting with both Aub and Ago3. However, the
279:, although they only seem to be required in males. In invertebrates, piRNAs have been detected in both the male and female
4583:
4469:
3425:
Kirino Y, Kim N, de
Planell-Saguer M, Khandros E, Chiorean S, Klein PS, Rigoutsos I, Jongens TA, Mourelatos Z (May 2009).
953:"Delving into the diversity of silencing pathways. Symposium on MicroRNAs and siRNAs: biological functions and mechanisms"
526:
75:
and other spurious or repeat-derived transcripts, but can also be involved in the regulation of other genetic elements in
482:
function across species contributes to the difficulty in establishing the functionality of piRNAs. However, like other
4696:
4691:
4676:
4301:
2359:
Grimson A, Srivastava M, Fahey B, Woodcroft BJ, Chiang HR, King N, Degnan BM, Rokhsar DS, Bartel DP (October 2008).
4039:
733:
905:
Siomi MC, Sato K, Pezic D, Aravin AA (April 2011). "PIWI-interacting small RNAs: the vanguard of genome defence".
742:
313:
311:
possible that active rasiRNA exist and have yet to be observed in other animals. RasiRNAs have been observed in
3617:"Non-coding RNA fragments account for the majority of annotated piRNAs expressed in somatic non-gonadal tissues"
2806:
Le Thomas A, Rogers AK, Webster A, Marinov GK, Liao SE, Perkins EM, Hur JK, Aravin AA, Tóth KF (February 2013).
2308:
1046:"History of the discovery of a master locus producing piRNAs: the flamenco/COM locus in Drosophila melanogaster"
294:, suggesting that piRNA pathways may function in both of these areas and, therefore, may have multiple effects.
4265:
370:
346:
145:
and modes of action do vary somewhat between species, a number of features are conserved. piRNAs have no clear
1139:
Xie Z, Johansen LK, Gustafson AM, Kasschau KD, Lellis AD, Zilberman D, Jacobsen SE, Carrington JC (May 2004).
2579:"Aub and Ago3 Are Recruited to Nuage through Two Mechanisms to Form a Ping-Pong Complex Assembled by Krimper"
2253:
Aravin AA, Sachidanandam R, Bourc'his D, Schaefer C, Pezic D, Toth KF, Bestor T, Hannon GJ (September 2008).
2193:
Tomari Y, Du T, Haley B, Schwarz DS, Bennett R, Cook HA, Koppetsch BS, Theurkauf WE, Zamore PD (March 2004).
1582:
Lin H, Yin H, Beyret E, Findley S, Deng W (2008). "The role of the piRNA pathway in stem cell self-renewal".
4742:
2924:
1484:
Kirino Y, Mourelatos Z (April 2007). "Mouse Piwi-interacting RNAs are 2′-O-methylated at their 3′ termini".
392:
204:
168:
2710:"Noncoding RNA. piRNA-guided slicing specifies transcripts for Zucchini-dependent, phased piRNA biogenesis"
2085:
Houwing S, Kamminga LM, Berezikov E, Cronembold D, Girard A, van den Elst H, Filippov DV, Blaser H, Raz E,
432:
The molecular mechanism that underpins Ping-Pong likely involves several piRNA pathway associated factors.
82:
piRNAs are mostly created from loci that function as transposon traps which provide a kind of RNA-mediated
4716:
4409:
4294:
4090:
4007:
505:
379:
162:
4593:
4437:
4182:
4032:
3405:
3293:
3228:
3162:
3097:
3020:
566:
2808:"Piwi induces piRNA-guided transcriptional silencing and establishment of a repressive chromatin state"
2759:"Noncoding RNA. piRNA-guided transposon cleavage initiates Zucchini-dependent, phased piRNA production"
2525:
Zhang Z, Xu J, Koppetsch BS, Wang J, Tipping C, Ma S, Weng Z, Theurkauf WE, Zamore PD (November 2011).
584:
which are required to recognise and silence transposons, but this relationship is not well understood.
2416:
Gunawardane LS, Saito K, Nishida KM, Miyoshi K, Kawamura Y, Nagami T, Siomi H, Siomi MC (March 2007).
1324:"Large-scale sequencing reveals 21U-RNAs and additional microRNAs and endogenous siRNAs in C. elegans"
4628:
4611:
4019:
3757:
3677:
3057:
2980:
2871:
2429:
2372:
2151:
2000:
1945:
1683:
1380:
1206:
569:(endo-siRNA) may have comparable and even redundant functionality in transposon control in mammalian
420:
piRNA from Mili and Miwi2 proteins isolated from mouse testes. The proposed function of Ping-Pong in
72:
2626:
Sato K, Iwasaki YW, Shibuya A, Carninci P, Tsuchizawa Y, Ishizu H, Siomi MC, Siomi H (August 2015).
4623:
4606:
4464:
4347:
4162:
4111:
3879:
Carmell MA, Girard A, van de Kant HJ, Bourc'his D, Bestor TH, de Rooij DG, Hannon GJ (April 2007).
1837:"Reexamining the P-Element Invasion of Drosophila melanogaster Through the Lens of piRNA Silencing"
592:
In
Dipterans viral-derived piRNAs derived from positive-sense RNA viruses were first identified in
403:
216:
211:
146:
3427:"Arginine methylation of Piwi proteins catalysed by dPRMT5 is required for Ago3 and Aub stability"
4517:
4512:
4387:
4192:
4187:
3881:"MIWI2 is essential for spermatogenesis and repression of transposons in the mouse male germline"
3781:
3701:
3499:
3393:
2949:
2455:
2341:
2224:
2175:
2124:
1969:
1509:
1466:
1404:
1353:
1232:
1000:
Klattenhoff C, Bratu DP, McGinnis-Schultz N, Koppetsch BS, Cook HA, Theurkauf WE (January 2007).
930:
737:
397:
396:
contain adenine at their tenth position, and this has been interpreted as possible evidence of a
2255:"A piRNA pathway primed by individual transposons is linked to de novo DNA methylation in mice"
4601:
4563:
4556:
4507:
4459:
4197:
4106:
3982:
3947:
3902:
3867:
3824:
3773:
3736:
3693:
3646:
3597:
3548:
3491:
3456:
3385:
3344:
3281:
3216:
3150:
3085:
3008:
2941:
2897:
2837:
2788:
2739:
2690:
2649:
2628:"Krimper Enforces an Antisense Bias on piRNA Pools by Binding AGO3 in the Drosophila Germline"
2608:
2577:
Webster A, Li S, Hur JK, Wachsmuth M, Bois JS, Perkins EM, Patel DJ, Aravin AA (August 2015).
2556:
2504:
2447:
2398:
2333:
2309:"Discrete small RNA-generating loci as master regulators of transposon activity in Drosophila"
2284:
2216:
2167:
2116:
2067:
2026:
1961:
1917:
1866:
1817:
1761:
1709:
1645:
1561:
1501:
1458:
1396:
1345:
1282:
1224:
1172:
1121:
1077:
1023:
982:
922:
884:
832:
577:
83:
383:, for instance, does have piRNAs, but does not appear to use the ping pong mechanism at all.
187:
More than 50,000 unique piRNA sequences have been discovered in mice and more than 13,000 in
4427:
3972:
3937:
3927:
3892:
3857:
3849:
3836:
Watanabe T, Takeda A, Tsukiyama T, Mise K, Okuno T, Sasaki H, Minami N, Imai H (July 2006).
3814:
3806:
3765:
3726:
3685:
3636:
3628:
3587:
3579:
3538:
3530:
3483:
3446:
3438:
3375:
3334:
3326:
3271:
3261:
3206:
3198:
3185:
Varjak M, Maringer K, Watson M, Sreenu VB, Fredericks AC, Pondeville E; et al. (2017).
3140:
3130:
3075:
3065:
2998:
2988:
2933:
2887:
2879:
2827:
2819:
2778:
2770:
2729:
2721:
2680:
2639:
2598:
2590:
2546:
2538:
2494:
2486:
2437:
2388:
2380:
2323:
2274:
2266:
2206:
2159:
2106:
2098:
2057:
2016:
2008:
1953:
1907:
1897:
1856:
1848:
1807:
1799:
1751:
1743:
1699:
1691:
1635:
1627:
1591:
1551:
1543:
1493:
1448:
1440:
1388:
1335:
1272:
1214:
1162:
1152:
1111:
1067:
1057:
1013:
972:
964:
914:
874:
866:
822:
784:
2527:"Heterotypic piRNA Ping-Pong requires qin, a protein with both E3 ligase and Tudor domains"
2307:
Brennecke J, Aravin AA, Stark A, Dus M, Kellis M, Sachidanandam R, Hannon GJ (March 2007).
4551:
4522:
4234:
2475:"Collapse of germline piRNAs in the absence of Argonaute3 reveals somatic piRNAs in flies"
2111:
1886:"proTRAC—a software for probabilistic piRNA cluster detection, visualization and analysis"
1453:
550:
510:
233:
232:
proposed that double-stranded (ds) RNA-mediated silencing is implicated in the control of
149:
motifs, due to the fact that the length of a piRNA varies between species (from 21 to 31
3761:
3681:
3061:
2984:
2875:
2433:
2418:"A slicer-mediated mechanism for repeat-associated siRNA 5′ end formation in Drosophila"
2376:
2155:
2004:
1949:
1687:
1670:
Brennecke J, Malone CD, Aravin AA, Sachidanandam R, Stark A, Hannon GJ (November 2008).
1384:
1322:
Ruby JG, Jan C, Player C, Axtell MJ, Lee W, Nusbaum C, Ge H, Bartel DP (December 2006).
1210:
191:. It is thought that there are many hundreds of thousands of different piRNA species in
4643:
4546:
4363:
3942:
3915:
3862:
3837:
3819:
3794:
3641:
3616:
3592:
3567:
3543:
3518:
3451:
3426:
3339:
3314:
3276:
3249:
3211:
3186:
3145:
3118:
3080:
3041:
3003:
2969:"Virus discovery by deep sequencing and assembly of virus-derived small silencing RNAs"
2968:
2892:
2859:
2832:
2807:
2783:
2758:
2734:
2709:
2603:
2578:
2551:
2526:
2499:
2474:
2393:
2360:
2279:
2254:
2021:
1988:
1912:
1885:
1861:
1836:
1812:
1787:
1756:
1731:
1704:
1671:
1640:
1615:
1556:
1531:
1072:
1045:
977:
952:
879:
854:
487:
253:
45:
3119:"Comparative genomics of small RNA regulatory pathway components in vector mosquitoes"
2211:
2194:
1219:
1194:
1167:
1140:
870:
4731:
4578:
4568:
4449:
4404:
4382:
4218:
4147:
4075:
4070:
2086:
1989:"Pseudogene-derived small interfering RNAs regulate gene expression in mouse oocytes"
522:
495:
437:
358:
3705:
3397:
3187:"Aedes aegypti Piwi4 Is a Noncanonical PIWI Protein Involved in Antiviral Responses"
2953:
2669:"RNA clamping by Vasa assembles a piRNA amplifier complex on transposon transcripts"
2459:
2228:
2128:
1513:
1470:
1408:
1357:
1236:
4737:
4661:
4481:
4399:
4318:
4275:
4085:
4016:– a software for probabilistic piRNA cluster detection, visualization, and analysis
3785:
3503:
2345:
2179:
1973:
934:
640:
546:
400:
287:
138:
2062:
2045:
1116:
1099:
3897:
3880:
3266:
2644:
2627:
2594:
2542:
2270:
1631:
1157:
1018:
1001:
827:
810:
4573:
3715:"Small RNAs just got bigger: Piwi-interacting RNAs (piRNAs) in mammalian testes"
3380:
3363:
1852:
1595:
1193:
Aravin AA, Naumova NM, Tulin AV, Vagin VV, Rozovsky YM, Gvozdev VA (July 2001).
581:
224:. The site of the mutations that made these Gypsies "dance" was thus called the
64:
4001:
3583:
3534:
2685:
2668:
2490:
2361:"Early origins and evolution of microRNAs and Piwi-interacting RNAs in animals"
2328:
2102:
1803:
1747:
1547:
1444:
1340:
1323:
789:
772:
4633:
4342:
4337:
4332:
4239:
4213:
4013:
3632:
2937:
2883:
613:
563:
491:
362:
334:
150:
142:
134:
3202:
1100:"Identification and characterization of small RNAs involved in RNA silencing"
1062:
968:
732:
Major advances in the study of piRNA have been achieved thanks to the use of
453:
protein coordinates the Ping-Pong mechanism of
Silkmoth Aub (Siwi) and Ago3.
302:
There are at least three
Argonaute (Ago) subfamilies that have been found in
4497:
4270:
4244:
4126:
4116:
3689:
3135:
3070:
2993:
2774:
2725:
2442:
2417:
1695:
1672:"An epigenetic role for maternally inherited piRNAs in transposon silencing"
1392:
693:
555:
542:
538:
534:
483:
450:
387:
342:
325:
303:
291:
260:
and vertebrate piRNAs have been located in areas lacking any protein-coding
241:
173:
76:
60:
3986:
3951:
3906:
3871:
3828:
3777:
3740:
3697:
3650:
3601:
3552:
3495:
3460:
3389:
3348:
3285:
3220:
3154:
3089:
3012:
2945:
2901:
2841:
2823:
2792:
2743:
2694:
2653:
2612:
2560:
2508:
2451:
2402:
2337:
2288:
2220:
2171:
2120:
2071:
2030:
1965:
1921:
1902:
1870:
1821:
1765:
1713:
1649:
1565:
1505:
1462:
1400:
1349:
1286:
1228:
1176:
1125:
1081:
1027:
986:
926:
888:
836:
4004:– a software for finding ping-pong signatures and ping-pong cycle activity
4651:
4502:
4432:
4286:
4167:
3330:
445:
280:
221:
101:
Double-stranded RNAs capable of silencing repeat elements, then known as
87:
3853:
3810:
3769:
3731:
3714:
2384:
2163:
2012:
1957:
1141:"Genetic and functional diversification of small RNA pathways in plants"
429:
ovaries that are mutant for Ago3 and the Tudor-domain protein Kumo/Qin.
4656:
4177:
4121:
3977:
3960:
1277:
1260:
374:
366:
350:
276:
157:
52:
3487:
3040:
Petit M, Mongelli V, Frangeul L, Blanc H, Jiggins F, Saleh MC (2016).
4671:
4666:
4616:
4249:
4157:
3916:"To be or not to be a piRNA: genomic origin and processing of piRNAs"
2860:"A C. elegans Piwi, PRG-1, regulates 21U-RNAs during spermatogenesis"
1497:
570:
500:
272:
207:
192:
160:
is common to piRNAs in both vertebrates and invertebrates. piRNAs in
17:
3442:
918:
329:
The ping-pong mechanism for the biogenesis of the 5′ end of rasiRNA.
203:
In the early 1980s, it was discovered that a single mutation in the
86:
against transposon expansions and invasions. They are distinct from
3932:
3519:"A PCR-based method for detection and quantification of small RNAs"
2046:"Tiny RNA: Where do we come from? What are we? Where are we going?"
4474:
4131:
324:
261:
124:
114:
110:
91:
537:. These are active in the testes of mammals and are required for
240:
A historical example of invasion and Piwi response is known: the
530:
479:
354:
177:
125:
95:
56:
4290:
4028:
4024:
3250:"PIWIs Go Viral: Arbovirus-Derived piRNAs in Vector Mosquitoes"
951:
Dorner S, Eulalio A, Huntzinger E, Izaurralde E (August 2007).
608:
piRNAs can be transmitted maternally, and based on research in
415:
The piRNA Ping-Pong pathway was first proposed from studies in
4536:
4454:
4419:
4374:
4357:
4056:
2915:
Ozata DM, Gainetdinov I, Zoch A, Phillip D, Zamore PD (2019).
2195:"RISC assembly defects in the Drosophila RNAi mutant armitage"
338:
286:
At the cellular level, piRNAs have been found within both the
181:
48:
2967:
Wu Q, Luo Y, Lu R, Lau N, Lai EC, Li WX; et al. (2010).
373:. One or both of these mechanisms may be acting in different
1732:"Mighty Piwis defend the germline against genome intruders"
63:
proteins. These piRNA complexes are mostly involved in the
3795:"A novel class of small RNAs in mouse spermatogenic cells"
3566:
Tang F, Hayashi K, Kaneda M, Lao K, Surani MA (May 2008).
533:
proteins that are part of a family of proteins called the
252:
piRNA clusters in genomes can now readily be detected via
129:
Proposed piRNA structure, with the 3′ end 2′-O-methylation
27:
Largest class of small non-coding RNA molecules in animals
3517:
Ro S, Park C, Jin J, Sanders KM, Yan W (December 2006).
3042:"piRNA pathway is not required for antiviral defense in
2917:"PIWI-interacting RNAs: small RNAs with big functions"
117:, while rasiRNA is now considered a piRNA subspecies.
30:"piRNA" redirects here. For the software package, see
51:
molecules expressed in animal cells. piRNAs form RNA-
2757:
Han BW, Wang W, Li C, Weng Z, Zamore PD (May 2015).
4642:
4592:
4535:
4490:
4418:
4373:
4356:
4325:
4258:
4227:
4206:
4140:
4099:
4063:
4010:– a web resource on classified and clustered piRNAs
3572:
3523:
Biochemical and Biophysical Research Communications
3568:"A sensitive multiplex assay for piRNA expression"
268:have been identified amidst protein-coding genes.
349:resulting in piRNAs with a tendency to target 5’
3914:Le Thomas A, Tóth KF, Aravin AA (January 2014).
3793:Grivna ST, Beyret E, Wang Z, Lin H (July 2006).
2572:
2570:
1044:Goriaux C, Théron E, Brasset E, Vaury C (2014).
345:piRNAs; in this mechanism, piRNA precursors are
3362:Parry R, Bishop C, De Hayr L, Asgari S (2019).
3180:
3178:
3117:Campbell CL, Black WC, Hess AM, Foy BD (2008).
2520:
2518:
1302:Journal of Bioinformatics and Sequence Analysis
635:Many factors required for the piRNA pathway in
612:, piRNAs may be involved in maternally derived
1254:
1252:
1250:
1248:
1246:
900:
898:
4302:
4040:
2302:
2300:
2298:
1261:"Biogenesis and germline functions of piRNAs"
386:A significant number of piRNAs identified in
8:
3410:: CS1 maint: multiple names: authors list (
3298:: CS1 maint: multiple names: authors list (
3233:: CS1 maint: multiple names: authors list (
3167:: CS1 maint: multiple names: authors list (
3102:: CS1 maint: multiple names: authors list (
3025:: CS1 maint: multiple names: authors list (
1665:
1663:
1661:
1659:
210:could specifically activate all copies of a
94:for biogenesis, at least in animals. (Plant
3615:Tosar JP, Rovira C, Cayota A (2018-01-22).
2708:Mohn F, Handler D, Brennecke J (May 2015).
1725:
1723:
1259:Klattenhoff C, Theurkauf W (January 2008).
767:
765:
763:
761:
759:
757:
98:may play a role in rasi/piRNA biogenesis.)
4370:
4309:
4295:
4287:
4047:
4033:
4025:
2248:
2246:
2244:
2242:
2240:
2238:
1525:
1523:
1426:
1424:
1422:
1420:
1418:
809:Seto AG, Kingston RE, Lau NC (June 2007).
478:The wide variation in piRNA sequences and
3976:
3941:
3931:
3896:
3861:
3818:
3730:
3640:
3591:
3542:
3450:
3379:
3338:
3275:
3265:
3210:
3144:
3134:
3079:
3069:
3002:
2992:
2891:
2831:
2782:
2733:
2684:
2643:
2602:
2550:
2498:
2441:
2392:
2327:
2278:
2210:
2110:
2061:
2020:
1911:
1901:
1884:Rosenkranz D, Zischler H (January 2012).
1860:
1811:
1781:
1779:
1777:
1775:
1755:
1703:
1639:
1555:
1530:Faehnle CR, Joshua-Tor L (October 2007).
1486:Nature Structural & Molecular Biology
1452:
1339:
1276:
1218:
1188:
1186:
1166:
1156:
1115:
1071:
1061:
1039:
1037:
1017:
976:
878:
826:
788:
3248:Miesen P, Joosten J, van Rij RP (2016).
946:
944:
562:. Further, it is thought that piRNA and
2853:
2851:
1788:"Small RNAs as guardians of the genome"
1317:
1315:
848:
846:
753:
620:Accessory proteins of the piRNA pathway
486:, piRNAs are thought to be involved in
103:repeat associated small interfering RNA
32:Partition function for Interacting RNAs
3403:
3291:
3226:
3160:
3095:
3018:
1786:Malone CD, Hannon GJ (February 2009).
1609:
1607:
1605:
1577:
1575:
1093:
1091:
804:
802:
800:
3961:"piRNAs: from biogenesis to function"
3959:Weick EM, Miska EA (September 2014).
1730:O'Donnell KA, Boeke JD (April 2007).
907:Nature Reviews Molecular Cell Biology
853:Monga I, Banerjee I (November 2019).
811:"The coming of age for Piwi proteins"
736:techniques, such as Solexa, 454, and
509:and humans, piRNAs are necessary for
271:In mammals, piRNAs are found both in
7:
1532:"Argonautes confront new small RNAs"
133:piRNAs have been identified in both
55:complexes through interactions with
1536:Current Opinion in Chemical Biology
1098:Aravin A, Tuschl T (October 2005).
576:piRNAs appear to affect particular
25:
4433:Micro
871:10.2174/1389202920666191129112705
361:of the transcript at a point ten
4712:
4711:
4153:Cis-natural antisense transcript
490:, specifically the silencing of
44:) is the largest class of small
4388:precursor, heterogenous nuclear
4081:Signal recognition particle RNA
4518:Trans-acting small interfering
4482:Enhancer RNAs
4400:Transfer
2858:Wang G, Reinke V (June 2008).
2112:11858/00-001M-0000-0012-E169-6
1454:11858/00-001M-0000-0012-E169-6
706:nuclear piRNA pathway proteins
357:proteins. This results in the
249:Piwi-interacting RNA pathway.
1:
4405:Ribosomal
4383:Messenger
2212:10.1016/S0092-8674(04)00218-1
2063:10.1016/j.tplants.2008.05.005
1220:10.1016/S0960-9822(01)00299-8
1117:10.1016/j.febslet.2005.08.009
529:(RISC). piRNAs interact with
527:RNA-induced silencing complex
494:. The majority of piRNAs are
3898:10.1016/j.devcel.2007.03.001
3267:10.1371/journal.ppat.1006017
2645:10.1016/j.molcel.2015.06.024
2595:10.1016/j.molcel.2015.07.017
2543:10.1016/j.molcel.2011.10.011
2271:10.1016/j.molcel.2008.09.003
1632:10.1016/j.molcel.2008.06.003
1158:10.1371/journal.pbio.0020104
1019:10.1016/j.devcel.2006.12.001
828:10.1016/j.molcel.2007.05.021
105:(rasiRNA), were proposed in
3381:10.1016/j.virol.2018.12.006
1853:10.1534/genetics.115.184119
1835:Kelleher ES (August 2016).
1596:10.1016/j.ydbio.2008.05.048
4759:
4584:Multicopy single-stranded
4428:Interferential
4240:Reverse transcribing virus
3584:10.1016/j.bbrc.2008.03.035
3535:10.1016/j.bbrc.2006.10.105
3313:Parry R, Asgari S (2018).
2686:10.1016/j.cell.2014.05.018
2491:10.1016/j.cell.2009.04.027
2329:10.1016/j.cell.2007.01.043
2103:10.1016/j.cell.2007.03.026
1804:10.1016/j.cell.2009.01.045
1748:10.1016/j.cell.2007.03.028
1548:10.1016/j.cbpa.2007.08.032
1445:10.1016/j.cell.2007.03.026
1341:10.1016/j.cell.2006.10.040
790:10.1016/j.cell.2006.07.012
773:"Molecular Biology Select"
734:next-generation sequencing
29:
4707:
4498:Guide
3633:10.1038/s42003-017-0001-7
2938:10.1038/s41576-018-0073-3
2884:10.1016/j.cub.2008.05.009
783:(2): 223–225. July 2006.
314:Schizosaccharomyces pombe
4460:Small nuclear
4020:piRNA cluster – database
3203:10.1128/mSphere.00144-17
3050:Proc Natl Acad Sci U S A
2973:Proc Natl Acad Sci U S A
1063:10.3389/fgene.2014.00257
969:10.1038/sj.embor.7401015
525:via the formation of an
4574:Genomic
4207:Cis-regulatory elements
4178:Repeat-associated siRNA
3842:Genes & Development
3799:Genes & Development
3719:Genes & Development
3690:10.1126/science.1130164
3136:10.1186/1471-2164-9-425
3071:10.1073/pnas.1607952113
3044:Drosophila melanogaster
2994:10.1073/pnas.0911353107
2925:Nature Reviews Genetics
2812:Genes & Development
2775:10.1126/science.aaa1264
2726:10.1126/science.aaa1039
2443:10.1126/science.1140494
2050:Trends in Plant Science
1696:10.1126/science.1165171
1393:10.1126/science.1129333
598:Drosophila melanogaster
246:Drosophila melanogaster
212:retrovirus-like element
169:Drosophila melanogaster
4677:Artificial chromosomes
4465:Small nucleolar
4091:Transfer-messenger RNA
3713:Kim VN (August 2006).
3621:Communications Biology
2824:10.1101/gad.209841.112
2044:Ruvkun G (July 2008).
1903:10.1186/1471-2105-13-5
688:piRNA pathway proteins
330:
163:Caenorhabditis elegans
153:), and the bias for a
130:
4470:Small Cajal Body RNAs
4183:Small interfering RNA
1584:Developmental Biology
1050:Frontiers in Genetics
631:Tudor domain proteins
567:small interfering RNA
328:
244:transposon invaded a
128:
73:transposable elements
4523:Subgenomic messenger
4438:Small interfering
4410:Transfer-messenger
4173:Piwi-interacting RNA
3331:10.1128/JVI.00224-18
676:Brother of Yb (BoYB)
521:piRNA has a role in
69:post-transcriptional
38:Piwi-interacting RNA
4112:Small nucleolar RNA
3854:10.1101/gad.1425706
3811:10.1101/gad.1434406
3770:10.1038/nature04917
3762:2006Natur.442..199G
3732:10.1101/gad.1456106
3682:2006Sci...313..363L
3062:2016PNAS..113E4218P
2985:2010PNAS..107.1606W
2876:2008CBio...18..861W
2434:2007Sci...315.1587G
2428:(5818): 1587–1590.
2385:10.1038/nature07415
2377:2008Natur.455.1193G
2371:(7217): 1193–1197.
2164:10.1038/nature04917
2156:2006Natur.442..199G
2013:10.1038/nature06904
2005:2008Natur.453..534T
1958:10.1038/nature04916
1950:2006Natur.442..203A
1688:2008Sci...322.1387B
1682:(5906): 1387–1392.
1385:2006Sci...313..320V
1211:2001CBio...11.1017A
679:Sister of Yb (SoYB)
147:secondary structure
4552:Chloroplast
4395:modified Messenger
4358:Ribonucleic acids
4193:Trans-acting siRNA
4188:Small temporal RNA
4163:Long noncoding RNA
3978:10.1242/dev.094037
3885:Developmental Cell
1890:BMC Bioinformatics
1278:10.1242/dev.006486
1006:Developmental Cell
604:Epigenetic effects
578:methyltransferases
331:
228:. In 2001, Aravin
131:
4725:
4724:
4602:Xeno
4564:Complementary
4537:Deoxyribonucleic
4531:
4530:
4508:Small hairpin
4284:
4283:
4198:Short hairpin RNA
4107:Small nuclear RNA
4064:Protein synthesis
3971:(18): 3458–3471.
3848:(13): 1732–1743.
3805:(13): 1709–1714.
3756:(7099): 199–202.
3725:(15): 1993–1997.
3676:(5785): 363–367.
3488:10.1242/dev.01809
2769:(6236): 817–821.
2720:(6236): 812–817.
2150:(7099): 199–202.
1999:(7194): 534–538.
1944:(7099): 203–207.
1379:(5785): 320–324.
1205:(13): 1017–1027.
1110:(26): 5830–5840.
588:Antiviral effects
580:that perform the
84:adaptive immunity
16:(Redirected from
4750:
4715:
4714:
4692:Yeast
4513:Small temporal
4443:Piwi-interacting
4371:
4367:
4348:Deoxynucleotides
4311:
4304:
4297:
4288:
4049:
4042:
4035:
4026:
3990:
3980:
3955:
3945:
3935:
3910:
3900:
3875:
3865:
3832:
3822:
3789:
3744:
3734:
3709:
3655:
3654:
3644:
3612:
3606:
3605:
3595:
3578:(4): 1190–1194.
3563:
3557:
3556:
3546:
3514:
3508:
3507:
3471:
3465:
3464:
3454:
3422:
3416:
3415:
3409:
3401:
3383:
3359:
3353:
3352:
3342:
3310:
3304:
3303:
3297:
3289:
3279:
3269:
3260:(12): e1006017.
3245:
3239:
3238:
3232:
3224:
3214:
3182:
3173:
3172:
3166:
3158:
3148:
3138:
3114:
3108:
3107:
3101:
3093:
3083:
3073:
3056:(29): E4218-27.
3037:
3031:
3030:
3024:
3016:
3006:
2996:
2964:
2958:
2957:
2921:
2912:
2906:
2905:
2895:
2855:
2846:
2845:
2835:
2803:
2797:
2796:
2786:
2754:
2748:
2747:
2737:
2705:
2699:
2698:
2688:
2679:(7): 1698–1711.
2664:
2658:
2657:
2647:
2623:
2617:
2616:
2606:
2574:
2565:
2564:
2554:
2522:
2513:
2512:
2502:
2470:
2464:
2463:
2445:
2413:
2407:
2406:
2396:
2356:
2350:
2349:
2331:
2322:(6): 1089–1103.
2313:
2304:
2293:
2292:
2282:
2250:
2233:
2232:
2214:
2190:
2184:
2183:
2139:
2133:
2132:
2114:
2082:
2076:
2075:
2065:
2041:
2035:
2034:
2024:
1984:
1978:
1977:
1932:
1926:
1925:
1915:
1905:
1881:
1875:
1874:
1864:
1847:(4): 1513–1531.
1832:
1826:
1825:
1815:
1783:
1770:
1769:
1759:
1727:
1718:
1717:
1707:
1667:
1654:
1653:
1643:
1611:
1600:
1599:
1579:
1570:
1569:
1559:
1527:
1518:
1517:
1498:10.1038/nsmb1218
1481:
1475:
1474:
1456:
1428:
1413:
1412:
1368:
1362:
1361:
1343:
1334:(6): 1193–1207.
1319:
1310:
1309:
1297:
1291:
1290:
1280:
1256:
1241:
1240:
1222:
1190:
1181:
1180:
1170:
1160:
1136:
1130:
1129:
1119:
1095:
1086:
1085:
1075:
1065:
1041:
1032:
1031:
1021:
997:
991:
990:
980:
948:
939:
938:
902:
893:
892:
882:
859:Current Genomics
850:
841:
840:
830:
806:
795:
794:
792:
769:
720:SetDB1 (Eggless)
698:Maelstrom (Mael)
654:Spindle-E (SpnE)
448:
234:retrotransposons
199:History and loci
21:
4758:
4757:
4753:
4752:
4751:
4749:
4748:
4747:
4728:
4727:
4726:
4721:
4703:
4644:Cloning vectors
4638:
4624:Locked
4588:
4538:
4527:
4486:
4414:
4361:
4360:
4352:
4321:
4315:
4285:
4280:
4254:
4235:Retrotransposon
4223:
4202:
4141:Gene regulation
4136:
4095:
4059:
4053:
3998:
3993:
3958:
3913:
3878:
3835:
3792:
3747:
3712:
3667:
3663:
3661:Further reading
3658:
3614:
3613:
3609:
3565:
3564:
3560:
3516:
3515:
3511:
3473:
3472:
3468:
3443:10.1038/ncb1872
3424:
3423:
3419:
3402:
3361:
3360:
3356:
3312:
3311:
3307:
3290:
3247:
3246:
3242:
3225:
3184:
3183:
3176:
3159:
3116:
3115:
3111:
3094:
3039:
3038:
3034:
3017:
2966:
2965:
2961:
2919:
2914:
2913:
2909:
2870:(12): 861–867.
2864:Current Biology
2857:
2856:
2849:
2805:
2804:
2800:
2756:
2755:
2751:
2707:
2706:
2702:
2666:
2665:
2661:
2625:
2624:
2620:
2576:
2575:
2568:
2524:
2523:
2516:
2472:
2471:
2467:
2415:
2414:
2410:
2358:
2357:
2353:
2311:
2306:
2305:
2296:
2252:
2251:
2236:
2192:
2191:
2187:
2141:
2140:
2136:
2084:
2083:
2079:
2043:
2042:
2038:
1986:
1985:
1981:
1934:
1933:
1929:
1883:
1882:
1878:
1834:
1833:
1829:
1785:
1784:
1773:
1729:
1728:
1721:
1669:
1668:
1657:
1613:
1612:
1603:
1581:
1580:
1573:
1529:
1528:
1521:
1483:
1482:
1478:
1430:
1429:
1416:
1370:
1369:
1365:
1321:
1320:
1313:
1299:
1298:
1294:
1258:
1257:
1244:
1199:Current Biology
1192:
1191:
1184:
1138:
1137:
1133:
1097:
1096:
1089:
1043:
1042:
1035:
999:
998:
994:
950:
949:
942:
919:10.1038/nrm3089
904:
903:
896:
852:
851:
844:
808:
807:
798:
771:
770:
755:
751:
730:
708:
690:
633:
622:
610:D. melanogaster
606:
590:
560:D. melanogaster
551:gene expression
545:development in
519:
511:spermatogenesis
476:
463:
444:
413:
393:D. melanogaster
323:
300:
258:D. melanogaster
256:methods. While
201:
189:D. melanogaster
141:, and although
123:
121:Characteristics
35:
28:
23:
22:
15:
12:
11:
5:
4756:
4754:
4746:
4745:
4743:Non-coding RNA
4740:
4730:
4729:
4723:
4722:
4720:
4719:
4708:
4705:
4704:
4702:
4701:
4700:
4699:
4694:
4689:
4684:
4674:
4669:
4664:
4659:
4654:
4648:
4646:
4640:
4639:
4637:
4636:
4631:
4629:Peptide
4626:
4621:
4620:
4619:
4614:
4609:
4607:Glycol
4598:
4596:
4590:
4589:
4587:
4586:
4581:
4576:
4571:
4566:
4561:
4560:
4559:
4554:
4543:
4541:
4533:
4532:
4529:
4528:
4526:
4525:
4520:
4515:
4510:
4505:
4500:
4494:
4492:
4488:
4487:
4485:
4484:
4479:
4478:
4477:
4472:
4467:
4462:
4452:
4447:
4446:
4445:
4440:
4435:
4424:
4422:
4416:
4415:
4413:
4412:
4407:
4402:
4397:
4392:
4391:
4390:
4379:
4377:
4368:
4354:
4353:
4351:
4350:
4345:
4340:
4335:
4329:
4327:
4323:
4322:
4319:nucleic acids
4316:
4314:
4313:
4306:
4299:
4291:
4282:
4281:
4279:
4278:
4273:
4268:
4266:Telomerase RNA
4262:
4260:
4256:
4255:
4253:
4252:
4247:
4242:
4237:
4231:
4229:
4225:
4224:
4222:
4221:
4216:
4210:
4208:
4204:
4203:
4201:
4200:
4195:
4190:
4185:
4180:
4175:
4170:
4165:
4160:
4155:
4150:
4144:
4142:
4138:
4137:
4135:
4134:
4129:
4124:
4119:
4114:
4109:
4103:
4101:
4100:RNA processing
4097:
4096:
4094:
4093:
4088:
4083:
4078:
4073:
4067:
4065:
4061:
4060:
4054:
4052:
4051:
4044:
4037:
4029:
4023:
4022:
4017:
4011:
4005:
3997:
3996:External links
3994:
3992:
3991:
3956:
3933:10.1186/gb4154
3920:Genome Biology
3911:
3891:(4): 503–514.
3876:
3833:
3790:
3745:
3710:
3664:
3662:
3659:
3657:
3656:
3607:
3558:
3529:(3): 756–763.
3509:
3482:(9): 2167–77.
3466:
3431:Nat. Cell Biol
3417:
3354:
3305:
3240:
3174:
3109:
3032:
2979:(4): 1606–11.
2959:
2907:
2847:
2818:(4): 390–399.
2798:
2749:
2700:
2659:
2638:(4): 553–563.
2632:Molecular Cell
2618:
2589:(4): 564–575.
2583:Molecular Cell
2566:
2537:(4): 572–584.
2531:Molecular Cell
2514:
2485:(3): 509–521.
2465:
2408:
2351:
2294:
2265:(6): 785–799.
2259:Molecular Cell
2234:
2205:(6): 831–841.
2185:
2134:
2077:
2056:(7): 313–316.
2036:
1979:
1927:
1876:
1827:
1798:(4): 656–668.
1771:
1719:
1655:
1620:Molecular Cell
1601:
1571:
1542:(5): 569–577.
1519:
1492:(4): 347–348.
1476:
1414:
1363:
1311:
1292:
1242:
1182:
1131:
1087:
1033:
992:
963:(8): 723–729.
940:
913:(4): 246–258.
894:
865:(7): 508–518.
842:
821:(5): 603–609.
815:Molecular Cell
796:
752:
750:
747:
729:
726:
725:
724:
721:
718:
715:
712:
707:
701:
700:
699:
696:
689:
682:
681:
680:
677:
674:
667:
664:
663:Vreteno (Vret)
661:
658:
655:
652:
649:
632:
626:
621:
618:
605:
602:
589:
586:
518:
515:
503:, and in both
488:gene silencing
475:
472:
462:
459:
412:
409:
322:
319:
299:
298:Classification
296:
254:bioinformatics
226:flamenco locus
220:in the female
200:
197:
122:
119:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
4755:
4744:
4741:
4739:
4736:
4735:
4733:
4718:
4710:
4709:
4706:
4698:
4695:
4693:
4690:
4688:
4685:
4683:
4680:
4679:
4678:
4675:
4673:
4670:
4668:
4665:
4663:
4660:
4658:
4655:
4653:
4650:
4649:
4647:
4645:
4641:
4635:
4632:
4630:
4627:
4625:
4622:
4618:
4615:
4613:
4612:Threose
4610:
4608:
4605:
4604:
4603:
4600:
4599:
4597:
4595:
4591:
4585:
4582:
4580:
4577:
4575:
4572:
4570:
4569:Deoxyribozyme
4567:
4565:
4562:
4558:
4557:Mitochondrial
4555:
4553:
4550:
4549:
4548:
4545:
4544:
4542:
4540:
4534:
4524:
4521:
4519:
4516:
4514:
4511:
4509:
4506:
4504:
4501:
4499:
4496:
4495:
4493:
4489:
4483:
4480:
4476:
4473:
4471:
4468:
4466:
4463:
4461:
4458:
4457:
4456:
4453:
4451:
4448:
4444:
4441:
4439:
4436:
4434:
4431:
4430:
4429:
4426:
4425:
4423:
4421:
4417:
4411:
4408:
4406:
4403:
4401:
4398:
4396:
4393:
4389:
4386:
4385:
4384:
4381:
4380:
4378:
4376:
4375:Translational
4372:
4369:
4365:
4359:
4355:
4349:
4346:
4344:
4341:
4339:
4336:
4334:
4331:
4330:
4328:
4324:
4320:
4312:
4307:
4305:
4300:
4298:
4293:
4292:
4289:
4277:
4274:
4272:
4269:
4267:
4264:
4263:
4261:
4257:
4251:
4248:
4246:
4243:
4241:
4238:
4236:
4233:
4232:
4230:
4226:
4220:
4219:SECIS element
4217:
4215:
4212:
4211:
4209:
4205:
4199:
4196:
4194:
4191:
4189:
4186:
4184:
4181:
4179:
4176:
4174:
4171:
4169:
4166:
4164:
4161:
4159:
4156:
4154:
4151:
4149:
4148:Antisense RNA
4146:
4145:
4143:
4139:
4133:
4130:
4128:
4125:
4123:
4120:
4118:
4115:
4113:
4110:
4108:
4105:
4104:
4102:
4098:
4092:
4089:
4087:
4084:
4082:
4079:
4077:
4076:Ribosomal RNA
4074:
4072:
4071:Messenger RNA
4069:
4068:
4066:
4062:
4058:
4050:
4045:
4043:
4038:
4036:
4031:
4030:
4027:
4021:
4018:
4015:
4012:
4009:
4006:
4003:
4000:
3999:
3995:
3988:
3984:
3979:
3974:
3970:
3966:
3962:
3957:
3953:
3949:
3944:
3939:
3934:
3929:
3925:
3921:
3917:
3912:
3908:
3904:
3899:
3894:
3890:
3886:
3882:
3877:
3873:
3869:
3864:
3859:
3855:
3851:
3847:
3843:
3839:
3834:
3830:
3826:
3821:
3816:
3812:
3808:
3804:
3800:
3796:
3791:
3787:
3783:
3779:
3775:
3771:
3767:
3763:
3759:
3755:
3751:
3746:
3742:
3738:
3733:
3728:
3724:
3720:
3716:
3711:
3707:
3703:
3699:
3695:
3691:
3687:
3683:
3679:
3675:
3671:
3666:
3665:
3660:
3652:
3648:
3643:
3638:
3634:
3630:
3626:
3622:
3618:
3611:
3608:
3603:
3599:
3594:
3589:
3585:
3581:
3577:
3573:
3569:
3562:
3559:
3554:
3550:
3545:
3540:
3536:
3532:
3528:
3524:
3520:
3513:
3510:
3505:
3501:
3497:
3493:
3489:
3485:
3481:
3477:
3470:
3467:
3462:
3458:
3453:
3448:
3444:
3440:
3436:
3432:
3428:
3421:
3418:
3413:
3407:
3399:
3395:
3391:
3387:
3382:
3377:
3373:
3369:
3365:
3358:
3355:
3350:
3346:
3341:
3336:
3332:
3328:
3324:
3320:
3316:
3309:
3306:
3301:
3295:
3287:
3283:
3278:
3273:
3268:
3263:
3259:
3255:
3251:
3244:
3241:
3236:
3230:
3222:
3218:
3213:
3208:
3204:
3200:
3196:
3192:
3188:
3181:
3179:
3175:
3170:
3164:
3156:
3152:
3147:
3142:
3137:
3132:
3128:
3124:
3120:
3113:
3110:
3105:
3099:
3091:
3087:
3082:
3077:
3072:
3067:
3063:
3059:
3055:
3051:
3047:
3045:
3036:
3033:
3028:
3022:
3014:
3010:
3005:
3000:
2995:
2990:
2986:
2982:
2978:
2974:
2970:
2963:
2960:
2955:
2951:
2947:
2943:
2939:
2935:
2932:(2): 89–108.
2931:
2927:
2926:
2918:
2911:
2908:
2903:
2899:
2894:
2889:
2885:
2881:
2877:
2873:
2869:
2865:
2861:
2854:
2852:
2848:
2843:
2839:
2834:
2829:
2825:
2821:
2817:
2813:
2809:
2802:
2799:
2794:
2790:
2785:
2780:
2776:
2772:
2768:
2764:
2760:
2753:
2750:
2745:
2741:
2736:
2731:
2727:
2723:
2719:
2715:
2711:
2704:
2701:
2696:
2692:
2687:
2682:
2678:
2674:
2670:
2663:
2660:
2655:
2651:
2646:
2641:
2637:
2633:
2629:
2622:
2619:
2614:
2610:
2605:
2600:
2596:
2592:
2588:
2584:
2580:
2573:
2571:
2567:
2562:
2558:
2553:
2548:
2544:
2540:
2536:
2532:
2528:
2521:
2519:
2515:
2510:
2506:
2501:
2496:
2492:
2488:
2484:
2480:
2476:
2469:
2466:
2461:
2457:
2453:
2449:
2444:
2439:
2435:
2431:
2427:
2423:
2419:
2412:
2409:
2404:
2400:
2395:
2390:
2386:
2382:
2378:
2374:
2370:
2366:
2362:
2355:
2352:
2347:
2343:
2339:
2335:
2330:
2325:
2321:
2317:
2310:
2303:
2301:
2299:
2295:
2290:
2286:
2281:
2276:
2272:
2268:
2264:
2260:
2256:
2249:
2247:
2245:
2243:
2241:
2239:
2235:
2230:
2226:
2222:
2218:
2213:
2208:
2204:
2200:
2196:
2189:
2186:
2181:
2177:
2173:
2169:
2165:
2161:
2157:
2153:
2149:
2145:
2138:
2135:
2130:
2126:
2122:
2118:
2113:
2108:
2104:
2100:
2096:
2092:
2088:
2081:
2078:
2073:
2069:
2064:
2059:
2055:
2051:
2047:
2040:
2037:
2032:
2028:
2023:
2018:
2014:
2010:
2006:
2002:
1998:
1994:
1990:
1983:
1980:
1975:
1971:
1967:
1963:
1959:
1955:
1951:
1947:
1943:
1939:
1931:
1928:
1923:
1919:
1914:
1909:
1904:
1899:
1895:
1891:
1887:
1880:
1877:
1872:
1868:
1863:
1858:
1854:
1850:
1846:
1842:
1838:
1831:
1828:
1823:
1819:
1814:
1809:
1805:
1801:
1797:
1793:
1789:
1782:
1780:
1778:
1776:
1772:
1767:
1763:
1758:
1753:
1749:
1745:
1741:
1737:
1733:
1726:
1724:
1720:
1715:
1711:
1706:
1701:
1697:
1693:
1689:
1685:
1681:
1677:
1673:
1666:
1664:
1662:
1660:
1656:
1651:
1647:
1642:
1637:
1633:
1629:
1625:
1621:
1617:
1610:
1608:
1606:
1602:
1597:
1593:
1589:
1585:
1578:
1576:
1572:
1567:
1563:
1558:
1553:
1549:
1545:
1541:
1537:
1533:
1526:
1524:
1520:
1515:
1511:
1507:
1503:
1499:
1495:
1491:
1487:
1480:
1477:
1472:
1468:
1464:
1460:
1455:
1450:
1446:
1442:
1438:
1434:
1427:
1425:
1423:
1421:
1419:
1415:
1410:
1406:
1402:
1398:
1394:
1390:
1386:
1382:
1378:
1374:
1367:
1364:
1359:
1355:
1351:
1347:
1342:
1337:
1333:
1329:
1325:
1318:
1316:
1312:
1308:(2): 031–040.
1307:
1303:
1296:
1293:
1288:
1284:
1279:
1274:
1270:
1266:
1262:
1255:
1253:
1251:
1249:
1247:
1243:
1238:
1234:
1230:
1226:
1221:
1216:
1212:
1208:
1204:
1200:
1196:
1189:
1187:
1183:
1178:
1174:
1169:
1164:
1159:
1154:
1150:
1146:
1142:
1135:
1132:
1127:
1123:
1118:
1113:
1109:
1105:
1101:
1094:
1092:
1088:
1083:
1079:
1074:
1069:
1064:
1059:
1055:
1051:
1047:
1040:
1038:
1034:
1029:
1025:
1020:
1015:
1011:
1007:
1003:
996:
993:
988:
984:
979:
974:
970:
966:
962:
958:
954:
947:
945:
941:
936:
932:
928:
924:
920:
916:
912:
908:
901:
899:
895:
890:
886:
881:
876:
872:
868:
864:
860:
856:
849:
847:
843:
838:
834:
829:
824:
820:
816:
812:
805:
803:
801:
797:
791:
786:
782:
778:
774:
768:
766:
764:
762:
760:
758:
754:
748:
746:
744:
739:
735:
728:Investigation
727:
722:
719:
716:
713:
710:
709:
705:
702:
697:
695:
692:
691:
687:
683:
678:
675:
672:
668:
665:
662:
659:
656:
653:
650:
647:
646:
645:
642:
641:Tudor domains
638:
630:
627:
625:
619:
617:
615:
611:
603:
601:
599:
595:
587:
585:
583:
579:
574:
572:
568:
565:
561:
557:
552:
548:
547:invertebrates
544:
540:
536:
532:
528:
524:
523:RNA silencing
517:RNA silencing
516:
514:
512:
508:
507:
502:
497:
493:
489:
485:
481:
473:
471:
468:
461:piRNA Phasing
460:
458:
454:
452:
447:
442:
439:
438:Tudor protein
435:
430:
428:
423:
418:
410:
408:
405:
402:
399:
395:
394:
389:
384:
382:
381:
376:
372:
371:transcription
368:
364:
360:
356:
352:
348:
344:
340:
336:
327:
320:
318:
316:
315:
309:
305:
297:
295:
293:
289:
284:
282:
278:
274:
269:
267:
263:
259:
255:
250:
247:
243:
238:
235:
231:
227:
223:
219:
218:
213:
209:
206:
198:
196:
194:
190:
185:
183:
179:
175:
171:
170:
165:
164:
159:
156:
152:
148:
144:
140:
139:invertebrates
136:
127:
120:
118:
116:
112:
108:
104:
99:
97:
93:
89:
85:
80:
78:
74:
71:silencing of
70:
66:
62:
58:
54:
50:
47:
43:
39:
33:
19:
4687:Bacterial
4662:Lambda phage
4442:
4326:Constituents
4276:List of RNAs
4172:
4086:Transfer RNA
3968:
3964:
3923:
3919:
3888:
3884:
3845:
3841:
3802:
3798:
3753:
3749:
3722:
3718:
3673:
3669:
3624:
3620:
3610:
3575:
3571:
3561:
3526:
3522:
3512:
3479:
3475:
3469:
3437:(5): 652–8.
3434:
3430:
3420:
3406:cite journal
3371:
3367:
3357:
3322:
3318:
3308:
3294:cite journal
3257:
3253:
3243:
3229:cite journal
3194:
3190:
3163:cite journal
3126:
3123:BMC Genomics
3122:
3112:
3098:cite journal
3053:
3049:
3043:
3035:
3021:cite journal
2976:
2972:
2962:
2929:
2923:
2910:
2867:
2863:
2815:
2811:
2801:
2766:
2762:
2752:
2717:
2713:
2703:
2676:
2672:
2662:
2635:
2631:
2621:
2586:
2582:
2534:
2530:
2482:
2478:
2468:
2425:
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2368:
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2354:
2319:
2315:
2262:
2258:
2202:
2198:
2188:
2147:
2143:
2137:
2097:(1): 69–82.
2094:
2090:
2080:
2053:
2049:
2039:
1996:
1992:
1982:
1941:
1937:
1930:
1893:
1889:
1879:
1844:
1840:
1830:
1795:
1791:
1742:(1): 37–44.
1739:
1735:
1679:
1675:
1626:(1): 79–90.
1623:
1619:
1587:
1583:
1539:
1535:
1489:
1485:
1479:
1439:(1): 69–82.
1436:
1432:
1376:
1372:
1366:
1331:
1327:
1305:
1301:
1295:
1268:
1264:
1202:
1198:
1148:
1145:PLOS Biology
1144:
1134:
1107:
1104:FEBS Letters
1103:
1053:
1049:
1012:(1): 45–55.
1009:
1005:
995:
960:
957:EMBO Reports
956:
910:
906:
862:
858:
818:
814:
780:
776:
731:
711:Rhino (HP1D)
703:
685:
670:
636:
634:
628:
623:
609:
607:
597:
593:
591:
582:methylations
575:
559:
520:
504:
477:
466:
464:
455:
440:
433:
431:
426:
421:
416:
414:
401:biosynthetic
391:
385:
378:
332:
312:
307:
301:
285:
270:
265:
264:, piRNAs in
257:
251:
245:
239:
229:
225:
215:
202:
188:
186:
167:
161:
132:
106:
102:
100:
81:
41:
37:
36:
4682:P1-derived
4450:Antisense
4343:Nucleotides
4338:Nucleosides
4333:Nucleobases
4002:PingPongPro
3965:Development
3476:Development
3254:PLOS Pathog
1265:Development
1151:(5): E104.
660:Tejas (Tej)
648:Tudor (Tud)
492:transposons
363:nucleotides
347:transcribed
151:nucleotides
135:vertebrates
59:-subfamily
4732:Categories
4634:Morpholino
4547:Organellar
4455:Processual
4420:Regulatory
4364:non-coding
4214:Riboswitch
4158:CRISPR RNA
4008:piRNA Bank
3926:(1): 204.
3374:: 89–100.
1590:(2): 479.
1271:(1): 3–9.
749:References
704:Drosophila
686:Drosophila
684:Non-Tudor
637:Drosophila
629:Drosophila
614:epigenetic
594:Drosophila
564:endogenous
535:Argonautes
506:C. elegans
484:small RNAs
467:Drosophila
427:Drosophila
422:Drosophila
417:Drosophila
380:C. elegans
335:biogenesis
321:Biogenesis
308:Drosophila
304:eukaryotes
266:C. elegans
143:biogenesis
107:Drosophila
65:epigenetic
46:non-coding
4594:Analogues
4579:Hachimoji
4362:(coding,
4317:Types of
4271:Vault RNA
4245:RNA virus
4228:Parasites
4127:RNase MRP
4117:Guide RNA
4055:Types of
556:fertility
543:stem-cell
539:germ-cell
496:antisense
411:Ping Pong
404:mechanism
398:conserved
388:zebrafish
343:pachytene
292:cytoplasm
281:germlines
242:P-element
205:fruit fly
174:zebrafish
77:germ line
61:Argonaute
4717:Category
4652:Phagemid
4503:Ribozyme
4168:MicroRNA
3987:25183868
3952:24467990
3907:17395546
3872:16766679
3829:16766680
3778:16751776
3741:16882976
3706:21150160
3698:16778019
3651:30271890
3627:(1): 2.
3602:18348866
3553:17084816
3496:15800004
3461:19377467
3398:58572380
3390:30583288
3368:Virology
3349:29950416
3286:28033427
3221:28497119
3155:18801182
3090:27357659
3013:20080648
2954:53565676
2946:30446728
2902:18501605
2842:23392610
2793:25977554
2744:25977553
2695:24910301
2654:26212455
2613:26295961
2561:22099305
2509:19395009
2460:11513777
2452:17322028
2403:18830242
2338:17346786
2289:18922463
2229:17588448
2221:15035985
2172:16751776
2129:13373509
2121:17418787
2087:Moens CB
2072:18562240
2031:18404147
1966:16751777
1922:22233380
1896:(5): 5.
1871:27516614
1841:Genetics
1822:19239887
1766:17418784
1714:19039138
1650:18571451
1566:17928262
1514:31193964
1506:17384647
1471:13373509
1463:17418787
1409:40471466
1401:16809489
1358:16838469
1350:17174894
1287:18032451
1237:14767819
1229:11470406
1177:15024409
1126:16153643
1082:25136352
1028:17199040
987:17599087
927:21427766
889:32655289
837:17560367
738:Illumina
723:SuVar3–9
714:Deadlock
651:Qin/Kumo
639:contain
474:Function
359:cleavage
351:uridines
222:germline
88:microRNA
4657:Plasmid
4122:RNase P
4014:proTRAC
3943:4053809
3863:1522070
3820:1522066
3786:3185036
3758:Bibcode
3678:Bibcode
3670:Science
3642:6052916
3593:3855189
3544:1934510
3504:6810484
3452:2746449
3340:6096813
3319:J Virol
3277:5198996
3212:5415634
3191:mSphere
3146:2566310
3129:: 425.
3081:4961201
3058:Bibcode
3004:2824396
2981:Bibcode
2893:2494713
2872:Bibcode
2833:3589556
2784:4545291
2763:Science
2735:4988486
2714:Science
2604:4545750
2552:3236501
2500:2768572
2430:Bibcode
2422:Science
2394:3837422
2373:Bibcode
2346:2246942
2280:2730041
2180:3185036
2152:Bibcode
2022:2981145
2001:Bibcode
1974:4379895
1946:Bibcode
1913:3293768
1862:4981261
1813:2792755
1757:4122227
1705:2805124
1684:Bibcode
1676:Science
1641:3353317
1557:2077831
1381:Bibcode
1373:Science
1207:Bibcode
1073:4120762
1056:: 257.
978:1978081
935:5710813
880:7327968
671:fs(1)Yb
657:Krimper
571:oocytes
441:krimper
375:species
367:adenine
288:nucleus
277:ovaries
214:called
193:mammals
158:uridine
79:cells.
53:protein
4672:Fosmid
4667:Cosmid
4617:Hexose
4539:acids
4491:Others
4250:Viroid
3985:
3950:
3940:
3905:
3870:
3860:
3827:
3817:
3784:
3776:
3750:Nature
3739:
3704:
3696:
3649:
3639:
3600:
3590:
3551:
3541:
3502:
3494:
3459:
3449:
3396:
3388:
3347:
3337:
3325:(17).
3284:
3274:
3219:
3209:
3153:
3143:
3088:
3078:
3011:
3001:
2952:
2944:
2900:
2890:
2840:
2830:
2791:
2781:
2742:
2732:
2693:
2652:
2611:
2601:
2559:
2549:
2507:
2497:
2458:
2450:
2401:
2391:
2365:Nature
2344:
2336:
2287:
2277:
2227:
2219:
2178:
2170:
2144:Nature
2127:
2119:
2070:
2029:
2019:
1993:Nature
1972:
1964:
1938:Nature
1920:
1910:
1869:
1859:
1820:
1810:
1764:
1754:
1712:
1702:
1648:
1638:
1564:
1554:
1512:
1504:
1469:
1461:
1407:
1399:
1356:
1348:
1285:
1235:
1227:
1175:
1168:350667
1165:
1124:
1080:
1070:
1026:
985:
975:
933:
925:
887:
877:
835:
717:Cutoff
501:embryo
446:A1ZAC4
273:testes
230:et al.
208:genome
180:, and
4697:Human
4475:Y RNA
4259:Other
4132:Y RNA
3782:S2CID
3702:S2CID
3500:S2CID
3394:S2CID
3197:(3).
2950:S2CID
2920:(PDF)
2456:S2CID
2342:S2CID
2312:(PDF)
2225:S2CID
2176:S2CID
2125:S2CID
1970:S2CID
1510:S2CID
1467:S2CID
1405:S2CID
1354:S2CID
1233:S2CID
931:S2CID
262:genes
217:Gypsy
115:siRNA
111:miRNA
92:Dicer
42:piRNA
18:PiRNA
3983:PMID
3948:PMID
3903:PMID
3868:PMID
3825:PMID
3774:PMID
3737:PMID
3694:PMID
3647:PMID
3598:PMID
3549:PMID
3492:PMID
3457:PMID
3412:link
3386:PMID
3345:PMID
3300:link
3282:PMID
3235:link
3217:PMID
3169:link
3151:PMID
3104:link
3086:PMID
3027:link
3009:PMID
2942:PMID
2898:PMID
2838:PMID
2789:PMID
2740:PMID
2691:PMID
2673:Cell
2650:PMID
2609:PMID
2557:PMID
2505:PMID
2479:Cell
2448:PMID
2399:PMID
2334:PMID
2316:Cell
2285:PMID
2217:PMID
2199:Cell
2168:PMID
2117:PMID
2091:Cell
2068:PMID
2027:PMID
1962:PMID
1918:PMID
1867:PMID
1818:PMID
1792:Cell
1762:PMID
1736:Cell
1710:PMID
1646:PMID
1562:PMID
1502:PMID
1459:PMID
1433:Cell
1397:PMID
1346:PMID
1328:Cell
1283:PMID
1225:PMID
1173:PMID
1122:PMID
1078:PMID
1024:PMID
983:PMID
923:PMID
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480:piwi
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1954:doi
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