145:
retroviruses and retrotransposons. When the retroviral DNA is integrated into the host genome they evolve into endogenous retroviruses that influence eukaryotic genomes. So many endogenous retroviruses have inserted themselves into eukaryotic genomes that they allow insight into biology between viral-host interactions and the role of retrotransposons in evolution and disease. Many retrotransposons share features with endogenous retroviruses, the property of recognising and fusing with the host genome. However, there is a key difference between retroviruses and retrotransposons, which is indicated by the env gene. Although similar to the gene carrying out the same function in retroviruses, the env gene is used to determine whether the gene is retroviral or retrotransposon. If the gene is retroviral it can evolve from a retrotransposon into a retrovirus. They differ by the order of sequences in pol genes. Env genes are found in LTR retrotransposon types Ty1-copia (
261:. SINEs exploit LINE transposition components despite LINE-binding proteins prefer binding to LINE RNA. SINEs cannot transpose by themselves because they cannot encode SINE transcripts. They usually consist of parts derived from tRNA and LINEs. The tRNA portion contains an RNA polymerase III promoter which the same kind of enzyme as RNA polymerase II. This makes sure the LINE copies would be transcribed into RNA for further transposition. The LINE component remains so LINE-binding proteins can recognise the LINE part of the SINE.
227:
L1 retrotransposition can disrupt the nature of genes transcribed by pasting themselves inside or near genes which could in turn lead to human disease. LINE1s can only retrotranspose in some cases to form different chromosome structures contributing to differences in genetics between individuals. There is an estimate of 80–100 active L1s in the reference genome of the Human Genome
Project, and an even smaller number of L1s within those active L1s retrotranspose often. L1 insertions have been associated with
107:(TPRT) process, which requires the RNA of the TE to be brought to the cleavage site of the retrotransposon’s integrase, where it is reverse transcribed. In contrast, LTR retrotransposons undergo reverse transcription in the cytoplasm, utilizing two rounds of template switching, and a formation of a pre-integration complex (PIC) composed of double-stranded DNA and an integrase dimer bound to LTRs. This complex then moves into the nucleus for integration into a new genomic location.
361:
288:(hence the name). Their distribution may be important in some genetic diseases and cancers. Copy and pasting Alu RNA requires the Alu's adenine-rich end and the rest of the sequence bound to a signal. The signal-bound Alu can then associate with ribosomes. LINE RNA associates on the same ribosomes as the Alu. Binding to the same ribosome allows Alus of SINEs to interact with LINE. This simultaneous translation of Alu element and LINE allows SINE copy and pasting.
153:) and BEL/Pao. They encode glycoproteins on the retrovirus envelope needed for entry into the host cell. Retroviruses can move between cells whereas LTR retrotransposons can only move themselves into the genome of the same cell. Many vertebrate genes were formed from retroviruses and LTR retrotransposons. One endogenous retrovirus or LTR retrotransposon has the same function and genomic locations in different species, suggesting their role in evolution.
208:
33:
200:
with hydroxyl groups. Reverse transcriptase recognises these hydroxyl groups to synthesise LINE retrotransposon where the DNA is cut. Like with LTR retrotransposons, this new inserted LINE contains eukaryotic genome information so it can be copied and pasted into other genomic regions easily. The information sequences are longer and more variable than those in LTR retrotransposons.
306:
LINE retrotransposition affects brain function. LINE retrotransposition is also a feature of several cancers, but it is unclear whether retrotransposition itself causes cancer instead of just a symptom. Uncontrolled retrotransposition is bad for both the host organism and retrotransposons themselves so they have to be regulated. Retrotransposons are regulated by
122:(ORF) or separated into distinct ORFs. Similar to retroviruses, the gag protein is essential for capsid assembly and the packaging of the TE's RNA and associated proteins. The pol protein is necessary for reverse transcription and includes these crucial domains: PR (protease), RT (reverse transcriptase), RH (
305:
Retrotransposons ensure they are not lost by chance by occurring only in cell genetics that can be passed on from one generation to the next from parent gametes. However, LINEs can transpose into the human embryo cells that eventually develop into the nervous system, raising the question whether this
191:
When a LINE is transcribed, the transcript contains an RNA polymerase II promoter that ensures LINEs can be copied into whichever location it inserts itself into. RNA polymerase II is the enzyme that transcribes genes into mRNA transcripts. The ends of LINE transcripts are rich in multiple adenines,
173:
They mostly fall into two types – LINEs (Long interspersed nuclear elements) and SINEs (Short interspersed nuclear elements). SVA elements are the exception between the two as they share similarities with both LINEs and SINEs, containing Alu elements and different numbers of the same repeat. SVAs are
99:
LTR retrotransposons are characterized by their long terminal repeats (LTRs), which are present at both the 5' and 3' ends of their sequences. These LTRs contain the promoters for these transposable elements (TEs), are essential for TE integration, and can vary in length from just over 100 base pairs
296:
SVA elements are present at lower levels than SINES and LINEs in humans. The starts of SVA and Alu elements are similar, followed by repeats and an end similar to endogenous retrovirus. LINEs bind to sites flanking SVA elements to transpose them. SVA are one of the youngest transposons in great apes
238:
Human L1 actively retrotransposes in the human genome. A recent study identified 1,708 somatic L1 retrotransposition events, especially in colorectal epithelial cells. These events occur from early embryogenesis and retrotransposition rate is substantially increased during colorectal tumourigenesis.
161:
Like LTR retrotransposons, non-LTR retrotransposons contain genes for reverse transcriptase, RNA-binding protein, nuclease, and sometimes ribonuclease H domain but they lack the long terminal repeats. RNA-binding proteins bind the RNA-transposition intermediate and nucleases are enzymes that break
326:
Due to their retrotransposition mechanism, retrotransposons amplify in number quickly, composing 40% of the human genome. The insertion rates for LINE1, Alu and SVA elements are 1/200 – 1/20, 1/20 and 1/900 respectively. The LINE1 insertion rates have varied a lot over the past 35 million years, so
322:
LTR retrotransposons came about later than non-LTR retrotransposons, possibly from an ancestral non-LTR retrotransposon acquiring an integrase from a DNA transposon. Retroviruses gained additional properties to their virus envelopes by taking the relevant genes from other viruses using the power of
226:
LINE-1 (L1) retrotransposons make up a significant portion of the human genome, with an estimated 500,000 copies per genome. Genes encoding for human LINE1 usually have their transcription inhibited by methyl groups binding to its DNA carried out by PIWI proteins and enzymes DNA methyltransferases.
199:
LINEs insert themselves into regions of the eukaryotic genome that are rich in bases AT. At AT regions LINE uses its nuclease to cut one strand of the eukaryotic double-stranded DNA. The adenine-rich sequence in LINE transcript base pairs with the cut strand to flag where the LINE will be inserted
253:
SINEs are much shorter (300bp) than LINEs. They share similarity with genes transcribed by RNA polymerase II, the enzyme that transcribes genes into mRNA transcripts, and the initiation sequence of RNA polymerase III, the enzyme that transcribes genes into ribosomal RNA, tRNA and other small RNA
234:
Each human LINE1 contains two regions from which gene products can be encoded. The first coding region contains a leucine zipper protein involved in protein-protein interactions and a protein that binds to the terminus of nucleic acids. The second coding region has a purine/pyrimidine nuclease,
169:
Although they are retrotransposons, they cannot carry out reverse transcription using an RNA transposition intermediate in the same way as LTR retrotransposons. Those two key components of the retrotransposon are still necessary but the way they are incorporated into the chemical reactions is
144:
An endogenous retrovirus is a retrovirus without virus pathogenic effects that has been integrated into the host genome by inserting their inheritable genetic information into cells that can be passed onto the next generation like a retrotransposon. Because of this, they share features with
195:
The RNA transposition intermediate moves from the nucleus into the cytoplasm for translation. This gives the two coding regions of a LINE that in turn binds back to the RNA it is transcribed from. The LINE RNA then moves back into the nucleus to insert into the eukaryotic genome.
330:
Notably a large number of 100 kilobases in the maize genome show variety due to the presence or absence of retrotransposons. However since maize is unusual genetically as compared to other plants it cannot be used to predict retrotransposition in other plants.
1383:
Nam, Chang Hyun; Youk, Jeonghwan; Kim, Jeong Yeon; Lim, Joonoh; Park, Jung Woo; Oh, Soo A; Lee, Hyun Jung; Park, Ji Won; Won, Hyein; Lee, Yunah; Jeong, Seung-Yong; Lee, Dong-Sung; Oh, Ji Won; Han, Jinju; Lee, Junehawk (2023-05-18).
790:
Wicker T, Sabot F, Hua-Van A, Bennetzen JL, Capy P, Chalhoub B, Flavell A, Leroy P, Morgante M, Panaud O, Paux E, SanMiguel P, Schulman AH (December 2007). "A unified classification system for eukaryotic transposable elements".
203:
Most LINE copies have variable length at the start because reverse transcription usually stops before DNA synthesis is complete. In some cases this causes RNA polymerase II promoter to be lost so LINEs cannot transpose further.
85:(LTRs) and non-long terminal repeats (non-LTRs). Retrotransposons are classified based on sequence and method of transposition. Most retrotransposons in the maize genome are LTR, whereas in humans they are mostly non-LTR.
977:
Yadav VP, Mandal PK, Rao DN, Bhattacharya S (December 2009). "Characterization of the restriction enzyme-like endonuclease encoded by the
Entamoeba histolytica non-long terminal repeat retrotransposon EhLINE1".
103:
LTRs are highly functional sequences, and for that reason LTR and non-LTR retrotransposons differ greatly in their reverse transcription and integration mechanisms. Non-LTR retrotransposons use a
211:
Genetic structure of murine LINE1 and SINEs. Bottom: proposed structure of L1 RNA-protein (RNP) complexes. ORF1 proteins form trimers, exhibiting RNA binding and nucleic acid chaperone activity.
162:
phosphodiester bonds between nucleotides in nucleic acids. Instead of LTRs, non-LTR retrotransposons have short repeats that can have an inverted order of bases next to each other aside from
297:
genome and among the most active and polymorphic in the human population. SVA was created by a fusion between an Alu element, a VNTR (variable number tandem repeat), and an LTR fragment.
55:. Thus, they differ from Class II transposable elements, or DNA transposons, in utilizing an RNA intermediate for the transposition and leaving the transposition donor site unchanged.
2121:
1852:
314:. The short non-coding RNA interacts with protein Argonaute to degrade retrotransposon transcripts and change their DNA histone structure to reduce their transcription.
1291:
Denli AM, Narvaiza I, Kerman BE, Pena M, Benner C, Marchetto MC, Diedrich JK, Aslanian A, Ma J, Moresco JJ, Moore L, Hunter T, Saghatelian A, Gage FH (October 2015).
100:(bp) to more than 1,000 bp. On average, LTR retrotransposons span several thousand base pairs, with the largest known examples reaching up to 30 kilobases (kb).
1909:
285:
235:
reverse transcriptase and protein rich in amino acids cysteines and histidines. The end of the human LINE1, as with other retrotransposons is adenine-rich.
192:
the bases that are added at the end of transcription so that LINE transcripts would not be degraded. This transcript is the RNA transposition intermediate.
1934:
1884:
1944:
177:
While historically viewed as "junk DNA", research suggests in some cases, both LINEs and SINEs were incorporated into novel genes to form new functions.
2066:
1984:
2114:
1974:
1889:
1240:
Doucet AJ, Hulme AE, Sahinovic E, Kulpa DA, Moldovan JB, Kopera HC, Athanikar JN, Hasnaoui M, Bucheton A, Moran JV, Gilbert N (October 2010).
1894:
836:"The Sinbad retrotransposon from the genome of the human blood fluke, Schistosoma mansoni, and the distribution of related Pao-like elements"
2107:
1620:
1989:
281:
are the most common SINE in primates. They are approximately 350 base pairs long, do not encode proteins and can be recognized by the
1954:
1904:
1899:
17:
1929:
1879:
1023:"Ancient exaptation of a CORE-SINE retroposon into a highly conserved mammalian neuronal enhancer of the proopiomelanocortin gene"
1924:
1829:
104:
1949:
1074:"A gene expression restriction network mediated by sense and antisense Alu sequences located on protein-coding messenger RNAs"
1460:
1191:"LINE retrotransposon RNA is an essential structural and functional epigenetic component of a core neocentromeric chromatin"
542:"Evidence that a recent increase in maize genome size was caused by the massive amplification of intergene retrotranposons"
170:
different. This is because unlike LTR retrotransposons, non-LTR retrotransposons do not contain sequences that bind tRNA.
1806:
1793:
1123:
Singer MF (March 1982). "SINEs and LINEs: highly repeated short and long interspersed sequences in mammalian genomes".
254:
molecules. SINEs such as mammalian MIR elements have tRNA gene at the start and adenine-rich at the end like in LINEs.
248:
938:"Not so bad after all: retroviruses and long terminal repeat retrotransposons as a source of new genes in vertebrates"
1816:
524:
258:
186:
2939:
1511:
Dewannieux M, Esnault C, Heidmann T (September 2003). "LINE-mediated retrotransposition of marked Alu sequences".
1613:
257:
SINEs do not encode a functional reverse transcriptase protein and rely on other mobile transposons, especially
2775:
2370:
936:
Naville M, Warren IA, Haftek-Terreau Z, Chalopin D, Brunet F, Levin P, Galiana D, Volff JN (17 February 2016).
2911:
1847:
1693:
1680:
1598:
2559:
2240:
634:"High-frequency recombination between members of an LTR retrotransposon family during transposition bursts"
2245:
2479:
2235:
1166:
Walter M (2015). "Transposon regulation upon dynamic loss of DNA methylation (PDF Download
Available)".
468:"An in vivo assay for the reverse transcriptase of human retrotransposon L1 in Saccharomyces cerevisiae"
130:) protein that is incorporated into the assembled capsid, facilitating attachment to cellular surfaces.
2944:
1964:
1606:
371:
2780:
691:"Becoming a Selfish Clan: Recombination Associated to Reverse-Transcription in LTR Retrotransposons"
2627:
2528:
2445:
1340:
Ohshima K, Okada N (2005). "SINEs and LINEs: symbionts of eukaryotic genomes with a common tail".
2949:
2654:
2430:
1658:
1633:
1189:
Chueh AC, Northrop EL, Brettingham-Moore KH, Choo KH, Wong LH (January 2009). Bickmore WA (ed.).
58:
Through reverse transcription, retrotransposons amplify themselves quickly to become abundant in
2882:
2198:
2088:
2016:
431:
111:
2642:
2615:
2230:
1021:
Santangelo AM, de Souza FS, Franchini LF, Bumaschny VF, Low MJ, Rubinstein M (October 2007).
396:
139:
126:), and INT (integrase). Additionally, some LTR retrotransposons have an ORF for an envelope (
75:
52:
2218:
573:
Lander ES, Linton LM, Birren B, Nusbaum C, Zody MC, Baldwin J, et al. (February 2001).
466:
Dombroski BA, Feng Q, Mathias SL, Sassaman DM, Scott AF, Kazazian HH, Boeke JD (July 1994).
2586:
2581:
2457:
2061:
1775:
1637:
1397:
645:
586:
401:
82:
48:
8:
2822:
2664:
2620:
2576:
2314:
1839:
1710:
411:
406:
94:
1571:
1428:
1401:
1385:
740:"Origin and evolution of retroelements based upon their reverse transcriptase sequences"
649:
590:
2872:
2011:
1919:
1738:
1580:
1555:
1536:
1365:
1322:
1268:
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1100:
1073:
1049:
1022:
1003:
816:
755:
715:
690:
666:
633:
282:
119:
66:(49–78%) and humans (42%). They are only present in eukaryotes but share features with
51:
which move in the host genome by converting their transcribed RNA into DNA through the
2099:
1489:
913:
886:
862:
835:
764:
739:
492:
467:
2867:
2571:
1585:
1528:
1493:
1456:
1433:
1415:
1357:
1314:
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1222:
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1136:
1105:
1054:
995:
991:
959:
918:
867:
808:
769:
720:
671:
614:
520:
497:
231:
by activating cancer-related genes oncogenes and diminishing tumor suppressor genes.
1540:
1369:
1326:
1152:
1007:
820:
2603:
2497:
2130:
2073:
2003:
1641:
1575:
1567:
1520:
1485:
1423:
1405:
1349:
1304:
1263:
1253:
1212:
1202:
1171:
1132:
1095:
1085:
1044:
1034:
987:
949:
908:
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847:
800:
759:
751:
710:
702:
661:
653:
604:
594:
553:
487:
479:
307:
166:
found in LTR retrotransposons that is just one sequence of bases repeating itself.
2860:
2554:
2533:
2265:
1756:
1720:
1258:
1207:
1039:
1175:
2850:
2737:
2608:
2467:
2450:
2255:
2185:
2056:
1914:
1871:
1699:
1689:
1410:
1309:
1292:
657:
437:
311:
123:
954:
937:
2933:
2890:
2855:
2698:
2688:
2659:
2292:
2285:
2021:
1761:
1684:
1675:
1667:
1628:
1476:
Kramerov DA, Vassetzky NS (2005). "Short retroposons in eukaryotic genomes".
1419:
1090:
421:
416:
228:
163:
146:
903:
2877:
2720:
2680:
2647:
2419:
2405:
2303:
2033:
1589:
1532:
1497:
1437:
1361:
1318:
1277:
1226:
1109:
1058:
999:
963:
922:
871:
852:
834:
Copeland CS, Mann VH, Morales ME, Kalinna BH, Brindley PJ (February 2005).
812:
724:
675:
618:
558:
541:
1386:"Widespread somatic L1 retrotransposition in normal colorectal epithelium"
1144:
773:
501:
483:
2901:
2767:
2715:
2710:
2223:
2028:
1939:
1801:
1293:"Primate-specific ORF0 contributes to retrotransposon-mediated diversity"
706:
632:
Sanchez DH, Gaubert H, Drost HG, Zabet NR, Paszkowski J (November 2017).
609:
275:
270:
150:
360:
2809:
2804:
2757:
2750:
2745:
2730:
2725:
2566:
2501:
2250:
2192:
2040:
1857:
442:
127:
67:
1353:
2814:
2799:
2786:
2538:
2326:
2157:
1959:
1650:
599:
574:
426:
115:
59:
1020:
804:
2845:
2791:
2693:
2637:
2632:
2523:
2484:
2440:
2342:
2147:
2083:
1969:
1733:
1629:
1524:
935:
207:
32:
1188:
2827:
2705:
2489:
2474:
2177:
2152:
1979:
36:
Simplified representation of the life cycle of a retrotransposon
2918:
2896:
2593:
2516:
2511:
2276:
2167:
2162:
2078:
447:
2400:
2392:
2210:
2172:
1824:
878:
221:
63:
465:
2906:
2598:
2139:
789:
976:
833:
1510:
1239:
572:
71:
1242:"Characterization of LINE-1 ribonucleoprotein particles"
631:
2129:
1290:
310:. RNA interference is carried out by a bunch of short
575:"Initial sequencing and analysis of the human genome"
929:
533:
1556:"A Field Guide to Eukaryotic Transposable Elements"
110:LTR retrotransposons typically encode the proteins
1455:(5th ed.). Oxford : Oxford University Press.
539:
434:, a powerful method of reconstructing phylogenies.
1475:
884:
2931:
1553:
1450:
334:Mutations caused by retrotransposons include:
81:There are two main types of retrotransposons,
2115:
1614:
1382:
885:Havecker ER, Gao X, Voytas DF (18 May 2004).
737:
1554:Wells, JN; Feschotte, C (23 November 2020).
1547:
1339:
1233:
688:
156:
2122:
2108:
1621:
1607:
1444:
1014:
327:they indicate points in genome evolution.
174:shorter than LINEs but longer than SINEs.
78:-mediated extrachromosomal recombination.
1579:
1427:
1409:
1308:
1267:
1257:
1216:
1206:
1099:
1089:
1065:
1048:
1038:
953:
912:
902:
861:
851:
763:
714:
665:
608:
598:
566:
557:
491:
1071:
785:
783:
351:
300:
206:
133:
31:
887:"The diversity of LTR retrotransposons"
88:
14:
2932:
1165:
1122:
1116:
827:
689:Drost HG, Sanchez DH (December 2019).
118:, which may be combined into a single
2103:
1602:
780:
738:Xiong Y, Eickbush TH (October 1990).
514:
355:
317:
1572:10.1146/annurev-genet-040620-022145
942:Clinical Microbiology and Infection
105:target-primed reverse transcription
24:
1690:Short tandem repeat/Microsatellite
1159:
756:10.1002/j.1460-2075.1990.tb07536.x
540:SanMiguel P, Bennetzen JL (1998).
249:Short interspersed nuclear element
25:
2961:
187:Long interspersed nuclear element
1478:International Review of Cytology
992:10.1111/j.1742-4658.2009.07419.x
519:. Washington (D.C.): ASM press.
359:
1504:
1469:
1451:Stansfield WD, King RC (1997).
1376:
1342:Cytogenetic and Genome Research
1333:
1284:
1182:
970:
291:
264:
2776:Last universal common ancestor
2371:Defective interfering particle
1694:Trinucleotide repeat disorders
731:
682:
625:
508:
472:Molecular and Cellular Biology
459:
13:
1:
2912:Clonally transmissible cancer
2348:Satellite-like nucleic acids
1681:Variable number tandem repeat
1490:10.1016/s0074-7696(05)47004-7
1072:Liang KH, Yeh CT (May 2013).
453:
74:, for example, discontinuous
45:Class I transposable elements
1259:10.1371/journal.pgen.1001150
1208:10.1371/journal.pgen.1000354
1137:10.1016/0092-8674(82)90194-5
1040:10.1371/journal.pgen.0030166
695:Genome Biology and Evolution
7:
1176:10.13140/rg.2.2.18747.21286
390:
215:
10:
2966:
2468:Class II or DNA transposon
2463:Class I or retrotransposon
1411:10.1038/s41586-023-06046-z
1310:10.1016/j.cell.2015.09.025
658:10.1038/s41467-017-01374-x
515:Craig, Nancy Lynn (2015).
347:Acting as DNA repair sites
268:
246:
219:
184:
137:
92:
2838:
2781:Earliest known life forms
2766:
2679:
2655:Repeated sequences in DNA
2547:
2429:
2418:
2391:
2363:
2313:
2302:
2275:
2264:
2209:
2138:
2049:
2002:
1870:
1838:
1815:
1792:
1783:
1774:
1749:
1709:
1666:
1657:
1648:
1560:Annual Review of Genetics
955:10.1016/j.cmi.2016.02.001
27:Type of genetic component
18:Long interspersed element
2628:Endogenous viral element
2446:Horizontal gene transfer
1453:A dictionary of genetics
1091:10.1186/1471-2164-14-325
840:BMC Evolutionary Biology
793:Nature Reviews. Genetics
341:Changing gene regulation
242:
180:
157:Non-LTR retrotransposons
2940:Mobile genetic elements
2325:dsDNA satellite virus (
904:10.1186/gb-2004-5-6-225
432:Retrotransposon markers
2883:Helper dependent virus
2199:Biological dark matter
2089:Protein tandem repeats
2017:Tandemly arrayed genes
853:10.1186/1471-2148-5-20
559:10.1006/anbo.1998.0746
368:This section is empty.
344:Changing gene products
212:
37:
2643:Endogenous retrovirus
2616:Origin of replication
2332:ssDNA satellite virus
2322:ssRNA satellite virus
638:Nature Communications
484:10.1128/mcb.14.7.4485
397:Copy-number variation
352:Role in biotechnology
323:LTR retrotransposon.
301:Role in human disease
210:
140:Endogenous retrovirus
134:Endogenous retrovirus
83:long terminal repeats
76:reverse transcriptase
53:reverse transcription
35:
2587:Secondary chromosome
2582:Extrachromosomal DNA
2458:Transposable element
2062:Pathogenicity island
402:Genomic organization
89:LTR retrotransposons
2823:Model lipid bilayer
2665:Interspersed repeat
1402:2023Natur.617..540N
650:2017NatCo...8.1283S
591:2001Natur.409..860L
412:Interspersed repeat
407:Insertion sequences
95:LTR retrotransposon
2133:organic structures
2012:Gene amplification
707:10.1093/gbe/evz255
552:(Suppl A): 37–44.
283:restriction enzyme
213:
120:open reading frame
38:
2945:Molecular biology
2927:
2926:
2868:Non-cellular life
2675:
2674:
2414:
2413:
2387:
2386:
2341:ssRNA satellite (
2097:
2096:
1998:
1997:
1866:
1865:
1770:
1769:
1659:Repeated sequence
1634:repeated sequence
1462:978-0-19-509441-1
1396:(7961): 540–547.
1354:10.1159/000084981
701:(12): 3382–3392.
585:(6822): 860–921.
388:
387:
338:Gene inactivation
318:Role in evolution
16:(Redirected from
2957:
2604:Gene duplication
2427:
2426:
2423:self-replication
2311:
2310:
2273:
2272:
2131:Self-replicating
2124:
2117:
2110:
2101:
2100:
2074:Low copy repeats
2067:Symbiosis island
2004:Gene duplication
1790:
1789:
1781:
1780:
1664:
1663:
1642:gene duplication
1623:
1616:
1609:
1600:
1599:
1594:
1593:
1583:
1551:
1545:
1544:
1508:
1502:
1501:
1473:
1467:
1466:
1448:
1442:
1441:
1431:
1413:
1380:
1374:
1373:
1337:
1331:
1330:
1312:
1288:
1282:
1281:
1271:
1261:
1252:(10): e1001150.
1237:
1231:
1230:
1220:
1210:
1186:
1180:
1179:
1163:
1157:
1156:
1120:
1114:
1113:
1103:
1093:
1069:
1063:
1062:
1052:
1042:
1018:
1012:
1011:
980:The FEBS Journal
974:
968:
967:
957:
933:
927:
926:
916:
906:
882:
876:
875:
865:
855:
831:
825:
824:
787:
778:
777:
767:
744:The EMBO Journal
735:
729:
728:
718:
686:
680:
679:
669:
629:
623:
622:
612:
602:
600:10.1038/35057062
570:
564:
563:
561:
546:Annals of Botany
537:
531:
530:
512:
506:
505:
495:
463:
383:
380:
370:You can help by
363:
356:
308:RNA interference
62:genomes such as
41:Retrotransposons
21:
2965:
2964:
2960:
2959:
2958:
2956:
2955:
2954:
2930:
2929:
2928:
2923:
2873:Synthetic virus
2861:Artificial cell
2834:
2762:
2671:
2560:RNA replication
2555:DNA replication
2543:
2534:Group II intron
2432:
2422:
2410:
2401:Mammalian prion
2383:
2359:
2338:dsRNA satellite
2335:ssDNA satellite
2305:
2298:
2267:
2260:
2205:
2134:
2128:
2098:
2093:
2045:
1994:
1862:
1834:
1811:
1785:Retrotransposon
1766:
1757:Inverted repeat
1745:
1730:DNA transposon
1726:Retrotransposon
1721:Gene conversion
1712:
1705:
1702:
1653:
1644:
1627:
1597:
1552:
1548:
1513:Nature Genetics
1509:
1505:
1474:
1470:
1463:
1449:
1445:
1381:
1377:
1348:(1–4): 475–90.
1338:
1334:
1289:
1285:
1238:
1234:
1201:(1): e1000354.
1187:
1183:
1164:
1160:
1121:
1117:
1070:
1066:
1033:(10): 1813–26.
1019:
1015:
986:(23): 7070–82.
975:
971:
934:
930:
883:
879:
832:
828:
805:10.1038/nrg2165
788:
781:
750:(10): 3353–62.
736:
732:
687:
683:
630:
626:
571:
567:
538:
534:
527:
513:
509:
464:
460:
456:
393:
384:
378:
375:
354:
320:
312:non-coding RNAs
303:
294:
273:
267:
251:
245:
224:
218:
189:
183:
159:
142:
136:
97:
91:
49:mobile elements
28:
23:
22:
15:
12:
11:
5:
2963:
2953:
2952:
2950:Non-coding DNA
2947:
2942:
2925:
2924:
2922:
2921:
2916:
2915:
2914:
2909:
2899:
2893:
2887:
2886:
2885:
2880:
2870:
2865:
2864:
2863:
2858:
2848:
2842:
2840:
2836:
2835:
2833:
2832:
2831:
2830:
2825:
2817:
2812:
2807:
2802:
2796:
2795:
2794:
2783:
2778:
2772:
2770:
2764:
2763:
2761:
2760:
2755:
2754:
2753:
2748:
2740:
2738:Kappa organism
2735:
2734:
2733:
2728:
2723:
2718:
2713:
2703:
2702:
2701:
2696:
2685:
2683:
2677:
2676:
2673:
2672:
2670:
2669:
2668:
2667:
2662:
2652:
2651:
2650:
2645:
2640:
2635:
2625:
2624:
2623:
2613:
2612:
2611:
2609:Non-coding DNA
2606:
2601:
2591:
2590:
2589:
2584:
2579:
2574:
2564:
2563:
2562:
2551:
2549:
2545:
2544:
2542:
2541:
2536:
2531:
2529:Group I intron
2526:
2521:
2520:
2519:
2509:
2508:
2507:
2504:
2495:
2492:
2487:
2482:
2472:
2471:
2470:
2465:
2455:
2454:
2453:
2451:Genomic island
2448:
2437:
2435:
2431:Mobile genetic
2424:
2416:
2415:
2412:
2411:
2409:
2408:
2403:
2397:
2395:
2389:
2388:
2385:
2384:
2382:
2381:
2380:
2379:
2376:
2367:
2365:
2361:
2360:
2358:
2357:
2356:
2355:
2352:
2346:
2339:
2336:
2333:
2330:
2323:
2319:
2317:
2308:
2300:
2299:
2297:
2296:
2289:
2281:
2279:
2270:
2262:
2261:
2259:
2258:
2256:dsDNA-RT virus
2253:
2251:ssRNA-RT virus
2248:
2246:(−)ssRNA virus
2243:
2241:(+)ssRNA virus
2238:
2233:
2228:
2227:
2226:
2215:
2213:
2207:
2206:
2204:
2203:
2202:
2201:
2196:
2186:Incertae sedis
2182:
2181:
2180:
2175:
2170:
2165:
2155:
2150:
2144:
2142:
2136:
2135:
2127:
2126:
2119:
2112:
2104:
2095:
2094:
2092:
2091:
2086:
2081:
2076:
2071:
2070:
2069:
2064:
2057:Genomic island
2053:
2051:
2047:
2046:
2044:
2043:
2038:
2037:
2036:
2026:
2025:
2024:
2014:
2008:
2006:
2000:
1999:
1996:
1995:
1993:
1992:
1987:
1982:
1977:
1972:
1967:
1962:
1957:
1952:
1947:
1942:
1937:
1932:
1927:
1922:
1917:
1912:
1907:
1902:
1897:
1892:
1887:
1882:
1876:
1874:
1872:DNA transposon
1868:
1867:
1864:
1863:
1861:
1860:
1855:
1850:
1844:
1842:
1836:
1835:
1833:
1832:
1827:
1821:
1819:
1813:
1812:
1810:
1809:
1804:
1798:
1796:
1787:
1778:
1772:
1771:
1768:
1767:
1765:
1764:
1759:
1753:
1751:
1747:
1746:
1744:
1743:
1742:
1741:
1736:
1728:
1723:
1717:
1715:
1707:
1706:
1704:
1703:
1700:Macrosatellite
1697:
1687:
1678:
1672:
1670:
1668:Tandem repeats
1661:
1655:
1654:
1649:
1646:
1645:
1626:
1625:
1618:
1611:
1603:
1596:
1595:
1546:
1525:10.1038/ng1223
1503:
1468:
1461:
1443:
1375:
1332:
1283:
1232:
1181:
1158:
1115:
1064:
1013:
969:
948:(4): 312–323.
928:
891:Genome Biology
877:
826:
799:(12): 973–82.
779:
730:
681:
624:
565:
532:
525:
517:Mobile DNA III
507:
478:(7): 4485–92.
457:
455:
452:
451:
450:
445:
440:
438:Tn3 transposon
435:
429:
424:
419:
414:
409:
404:
399:
392:
389:
386:
385:
366:
364:
353:
350:
349:
348:
345:
342:
339:
319:
316:
302:
299:
293:
290:
269:Main article:
266:
263:
247:Main article:
244:
241:
220:Main article:
217:
214:
185:Main article:
182:
179:
164:direct repeats
158:
155:
149:), Ty3-gypsy (
138:Main article:
135:
132:
93:Main article:
90:
87:
26:
9:
6:
4:
3:
2:
2962:
2951:
2948:
2946:
2943:
2941:
2938:
2937:
2935:
2920:
2917:
2913:
2910:
2908:
2905:
2904:
2903:
2900:
2898:
2894:
2892:
2891:Nanobacterium
2888:
2884:
2881:
2879:
2876:
2875:
2874:
2871:
2869:
2866:
2862:
2859:
2857:
2856:Cell division
2854:
2853:
2852:
2849:
2847:
2844:
2843:
2841:
2837:
2829:
2826:
2824:
2821:
2820:
2818:
2816:
2813:
2811:
2808:
2806:
2803:
2801:
2797:
2793:
2790:
2789:
2788:
2784:
2782:
2779:
2777:
2774:
2773:
2771:
2769:
2765:
2759:
2756:
2752:
2749:
2747:
2744:
2743:
2741:
2739:
2736:
2732:
2729:
2727:
2724:
2722:
2719:
2717:
2714:
2712:
2709:
2708:
2707:
2704:
2700:
2699:Hydrogenosome
2697:
2695:
2692:
2691:
2690:
2689:Mitochondrion
2687:
2686:
2684:
2682:
2681:Endosymbiosis
2678:
2666:
2663:
2661:
2660:Tandem repeat
2658:
2657:
2656:
2653:
2649:
2646:
2644:
2641:
2639:
2636:
2634:
2631:
2630:
2629:
2626:
2622:
2619:
2618:
2617:
2614:
2610:
2607:
2605:
2602:
2600:
2597:
2596:
2595:
2592:
2588:
2585:
2583:
2580:
2578:
2575:
2573:
2570:
2569:
2568:
2565:
2561:
2558:
2557:
2556:
2553:
2552:
2550:
2548:Other aspects
2546:
2540:
2537:
2535:
2532:
2530:
2527:
2525:
2522:
2518:
2515:
2514:
2513:
2510:
2505:
2503:
2499:
2496:
2493:
2491:
2488:
2486:
2483:
2481:
2478:
2477:
2476:
2473:
2469:
2466:
2464:
2461:
2460:
2459:
2456:
2452:
2449:
2447:
2444:
2443:
2442:
2439:
2438:
2436:
2434:
2428:
2425:
2421:
2417:
2407:
2404:
2402:
2399:
2398:
2396:
2394:
2390:
2377:
2374:
2373:
2372:
2369:
2368:
2366:
2362:
2353:
2350:
2349:
2347:
2344:
2340:
2337:
2334:
2331:
2328:
2324:
2321:
2320:
2318:
2316:
2312:
2309:
2307:
2301:
2295:
2294:
2293:Avsunviroidae
2290:
2288:
2287:
2286:Pospiviroidae
2283:
2282:
2280:
2278:
2274:
2271:
2269:
2263:
2257:
2254:
2252:
2249:
2247:
2244:
2242:
2239:
2237:
2234:
2232:
2229:
2225:
2222:
2221:
2220:
2217:
2216:
2214:
2212:
2208:
2200:
2197:
2195:
2194:
2190:
2189:
2188:
2187:
2183:
2179:
2176:
2174:
2171:
2169:
2166:
2164:
2161:
2160:
2159:
2156:
2154:
2151:
2149:
2146:
2145:
2143:
2141:
2140:Cellular life
2137:
2132:
2125:
2120:
2118:
2113:
2111:
2106:
2105:
2102:
2090:
2087:
2085:
2082:
2080:
2077:
2075:
2072:
2068:
2065:
2063:
2060:
2059:
2058:
2055:
2054:
2052:
2048:
2042:
2039:
2035:
2032:
2031:
2030:
2027:
2023:
2022:Ribosomal DNA
2020:
2019:
2018:
2015:
2013:
2010:
2009:
2007:
2005:
2001:
1991:
1988:
1986:
1983:
1981:
1978:
1976:
1973:
1971:
1968:
1966:
1963:
1961:
1958:
1956:
1953:
1951:
1948:
1946:
1943:
1941:
1938:
1936:
1933:
1931:
1928:
1926:
1923:
1921:
1918:
1916:
1913:
1911:
1908:
1906:
1903:
1901:
1898:
1896:
1893:
1891:
1888:
1886:
1883:
1881:
1878:
1877:
1875:
1873:
1869:
1859:
1856:
1854:
1851:
1849:
1846:
1845:
1843:
1841:
1837:
1831:
1828:
1826:
1823:
1822:
1820:
1818:
1814:
1808:
1805:
1803:
1800:
1799:
1797:
1795:
1791:
1788:
1786:
1782:
1779:
1777:
1773:
1763:
1762:Direct repeat
1760:
1758:
1755:
1754:
1752:
1748:
1740:
1737:
1735:
1732:
1731:
1729:
1727:
1724:
1722:
1719:
1718:
1716:
1714:
1708:
1701:
1698:
1695:
1691:
1688:
1686:
1685:Minisatellite
1682:
1679:
1677:
1676:Satellite DNA
1674:
1673:
1671:
1669:
1665:
1662:
1660:
1656:
1652:
1647:
1643:
1639:
1635:
1631:
1624:
1619:
1617:
1612:
1610:
1605:
1604:
1601:
1591:
1587:
1582:
1577:
1573:
1569:
1565:
1561:
1557:
1550:
1542:
1538:
1534:
1530:
1526:
1522:
1518:
1514:
1507:
1499:
1495:
1491:
1487:
1483:
1479:
1472:
1464:
1458:
1454:
1447:
1439:
1435:
1430:
1425:
1421:
1417:
1412:
1407:
1403:
1399:
1395:
1391:
1387:
1379:
1371:
1367:
1363:
1359:
1355:
1351:
1347:
1343:
1336:
1328:
1324:
1320:
1316:
1311:
1306:
1303:(3): 583–93.
1302:
1298:
1294:
1287:
1279:
1275:
1270:
1265:
1260:
1255:
1251:
1247:
1246:PLOS Genetics
1243:
1236:
1228:
1224:
1219:
1214:
1209:
1204:
1200:
1196:
1195:PLOS Genetics
1192:
1185:
1177:
1173:
1169:
1162:
1154:
1150:
1146:
1142:
1138:
1134:
1130:
1126:
1119:
1111:
1107:
1102:
1097:
1092:
1087:
1083:
1079:
1075:
1068:
1060:
1056:
1051:
1046:
1041:
1036:
1032:
1028:
1027:PLOS Genetics
1024:
1017:
1009:
1005:
1001:
997:
993:
989:
985:
981:
973:
965:
961:
956:
951:
947:
943:
939:
932:
924:
920:
915:
910:
905:
900:
896:
892:
888:
881:
873:
869:
864:
859:
854:
849:
845:
841:
837:
830:
822:
818:
814:
810:
806:
802:
798:
794:
786:
784:
775:
771:
766:
761:
757:
753:
749:
745:
741:
734:
726:
722:
717:
712:
708:
704:
700:
696:
692:
685:
677:
673:
668:
663:
659:
655:
651:
647:
643:
639:
635:
628:
620:
616:
611:
610:2027.42/62798
606:
601:
596:
592:
588:
584:
580:
576:
569:
560:
555:
551:
547:
543:
536:
528:
526:9781555819200
522:
518:
511:
503:
499:
494:
489:
485:
481:
477:
473:
469:
462:
458:
449:
446:
444:
441:
439:
436:
433:
430:
428:
425:
423:
422:Paleovirology
420:
418:
417:Paleogenetics
415:
413:
410:
408:
405:
403:
400:
398:
395:
394:
382:
373:
369:
365:
362:
358:
357:
346:
343:
340:
337:
336:
335:
332:
328:
324:
315:
313:
309:
298:
289:
287:
284:
280:
278:
272:
262:
260:
255:
250:
240:
236:
232:
230:
229:tumorigenesis
223:
209:
205:
201:
197:
193:
188:
178:
175:
171:
167:
165:
154:
152:
148:
147:Pseudoviridae
141:
131:
129:
125:
121:
117:
113:
108:
106:
101:
96:
86:
84:
79:
77:
73:
69:
65:
61:
56:
54:
50:
46:
43:(also called
42:
34:
30:
19:
2878:Viral vector
2721:Gerontoplast
2648:Transpoviron
2462:
2420:Nucleic acid
2406:Fungal prion
2304:Helper-virus
2291:
2284:
2191:
2184:
2034:Gene cluster
1802:Alu sequence
1784:
1725:
1711:Interspersed
1563:
1559:
1549:
1516:
1512:
1506:
1481:
1477:
1471:
1452:
1446:
1393:
1389:
1378:
1345:
1341:
1335:
1300:
1296:
1286:
1249:
1245:
1235:
1198:
1194:
1184:
1168:ResearchGate
1167:
1161:
1131:(3): 433–4.
1128:
1124:
1118:
1081:
1078:BMC Genomics
1077:
1067:
1030:
1026:
1016:
983:
979:
972:
945:
941:
931:
897:(225): 225.
894:
890:
880:
843:
839:
829:
796:
792:
747:
743:
733:
698:
694:
684:
641:
637:
627:
582:
578:
568:
549:
545:
535:
516:
510:
475:
471:
461:
379:January 2021
376:
372:adding to it
367:
333:
329:
325:
321:
304:
295:
292:SVA elements
276:
274:
265:Alu elements
256:
252:
237:
233:
225:
202:
198:
194:
190:
176:
172:
168:
160:
143:
109:
102:
98:
80:
68:retroviruses
57:
44:
40:
39:
29:
2902:Cancer cell
2768:Abiogenesis
2716:Chromoplast
2711:Chloroplast
2494:Degradative
2236:dsRNA virus
2231:ssDNA virus
2224:Giant virus
2219:dsDNA virus
2029:Gene family
1940:Tc1/mariner
1895:EnSpm/CACTA
1566:: 539–561.
1519:(1): 41–8.
1484:: 165–221.
644:(1): 1283.
271:Alu element
151:Metaviridae
2934:Categories
2810:Proteinoid
2805:Coacervate
2758:Nitroplast
2751:Trophosome
2746:Bacteriome
2731:Apicoplast
2726:Leucoplast
2567:Chromosome
2485:Resistance
2193:Parakaryon
2041:Pseudogene
1858:retroposon
1776:Transposon
1638:transposon
454:References
443:Transposon
60:eukaryotic
2819:Research
2800:Protocell
2539:Retrozyme
2498:Virulence
2480:Fertility
2327:Virophage
2315:Satellite
2306:dependent
2158:Eukaryota
1960:P element
1910:Harbinger
1651:Repeatome
1420:0028-0836
846:(1): 20.
427:RetrOryza
2846:Organism
2839:See also
2815:Sulphobe
2792:Ribozyme
2787:RNA life
2694:Mitosome
2638:Prophage
2633:Provirus
2621:Replicon
2577:Circular
2524:Phagemid
2441:Mobilome
2433:elements
2343:Virusoid
2266:Subviral
2178:Protista
2163:Animalia
2148:Bacteria
2084:Telomere
2050:See also
1990:Zisupton
1970:Polinton
1965:PiggyBac
1920:Helitron
1739:Helitron
1734:Polinton
1630:Genetics
1590:32955944
1541:32151696
1533:12897783
1498:16344113
1438:37165195
1429:10191854
1370:42841487
1362:16093701
1327:10525450
1319:26496605
1278:20949108
1227:19180186
1153:22129236
1110:23663499
1059:17922573
1008:30791213
1000:19878305
964:26899828
923:15186483
872:15725362
821:32132898
813:17984973
725:31755923
676:29097664
619:11237011
391:See also
216:Human L1
70:such as
2828:Jeewanu
2742:Organs
2706:Plastid
2506:Cryptic
2475:Plasmid
2173:Plantae
2153:Archaea
1980:Transib
1955:Novosib
1935:Kolobok
1905:Ginger2
1900:Ginger1
1885:Crypton
1581:8293684
1398:Bibcode
1269:2951350
1218:2625447
1145:6280868
1101:3655826
1084:: 325.
1050:2000970
774:1698615
716:6894440
667:5668417
646:Bibcode
587:Bibcode
502:7516468
124:RNase H
2919:Virome
2897:Nanobe
2594:Genome
2572:Linear
2517:Fosmid
2512:Cosmid
2277:Viroid
2268:agents
2079:CRISPR
1945:Merlin
1930:ISL2EU
1880:Academ
1713:repeat
1588:
1578:
1539:
1531:
1496:
1459:
1436:
1426:
1418:
1390:Nature
1368:
1360:
1325:
1317:
1276:
1266:
1225:
1215:
1151:
1143:
1108:
1098:
1057:
1047:
1006:
998:
962:
921:
914:463057
911:
870:
863:554778
860:
819:
811:
772:
765:552073
762:
723:
713:
674:
664:
617:
579:Nature
523:
500:
493:358820
490:
448:Retron
47:) are
2393:Prion
2364:Other
2211:Virus
2168:Fungi
1985:Zator
1925:IS3EU
1830:LINE2
1825:LINE1
1817:LINEs
1794:SINEs
1750:Other
1537:S2CID
1366:S2CID
1323:S2CID
1149:S2CID
1004:S2CID
817:S2CID
259:LINEs
243:SINEs
222:LINE1
181:LINEs
64:maize
2907:HeLa
2851:Cell
2599:Gene
1975:Sola
1950:MuDR
1890:Dada
1853:MER4
1848:HERV
1840:LTRs
1586:PMID
1529:PMID
1494:PMID
1457:ISBN
1434:PMID
1416:ISSN
1358:PMID
1315:PMID
1297:Cell
1274:PMID
1223:PMID
1141:PMID
1125:Cell
1106:PMID
1055:PMID
996:PMID
960:PMID
919:PMID
868:PMID
809:PMID
770:PMID
721:PMID
672:PMID
615:PMID
521:ISBN
498:PMID
286:AluI
114:and
2490:Col
2378:DNA
2375:RNA
2354:DNA
2351:RNA
1915:hAT
1807:MIR
1576:PMC
1568:doi
1521:doi
1486:doi
1482:247
1424:PMC
1406:doi
1394:617
1350:doi
1346:110
1305:doi
1301:163
1264:PMC
1254:doi
1213:PMC
1203:doi
1172:doi
1133:doi
1096:PMC
1086:doi
1045:PMC
1035:doi
988:doi
984:276
950:doi
909:PMC
899:doi
858:PMC
848:doi
801:doi
760:PMC
752:doi
711:PMC
703:doi
662:PMC
654:doi
605:hdl
595:doi
583:409
554:doi
488:PMC
480:doi
374:.
277:Alu
128:env
116:pol
112:gag
72:HIV
2936::
2502:Ti
1640:,
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