17:
248:
37:
254:, directly at the site of integration: L1 RNP recognize AAAATT hexanucleotides and ORF2 endonuclease activity cleaves the DNA first-strand. L1 polyA tail associate with TTTT overhang and the host DNA is used as a primer to initiate reverse-transcription. ORF2 probably also mediate second-strand cleavage and attachment of newly synthesized cDNA to the DNA template, using again host DNA as a primer for second-strand synthesis.
288:, where the ORF2 endonuclease domain opens the DNA (at TTAAAA hexanucleotide motifs in mammals). Thus, a 3'OH group is freed for the reverse transcriptase to prime reverse transcription of the LINE RNA transcript. Following the reverse transcription the target strand is cleaved and the newly created
236:
elements for their proliferation make up 13% of the human genome and have a copy number of around 1.5 million. They probably originated from the RTE family of LINEs. Recent estimates show the typical human genome contains on average 100 L1 elements with potential for mobilization, however there is a
210:
Remnants of L2 and L3 elements are found in the human genome. It is estimated that L2 and L3 elements were active ~200-300 million years ago. Due to the age of L2 elements found within therian genomes, they lack flanking target site duplications. The L2 (and L3) elements are in the same group as the
110:
class, and is referred to as L1Hs. The human genome contains an estimated 100,000 truncated and 4,000 full-length LINE-1 elements. Due to the accumulation of random mutations, the sequence of many LINEs has degenerated to the extent that they are no longer transcribed or translated. Comparisons of
323:
In plant genomes, epigenetic modification of LINEs can lead to expression changes of nearby genes and even to phenotypic changes: In the oil palm genome, methylation of a Karma-type LINE underlies the somaclonal, 'mantled' variant of this plant, responsible for drastic yield loss.
351:). Hypomethylation of LINES is associated with chromosomal instability and altered gene expression and is found in various cancer cell types in various tissues types. Hypomethylation of a specific L1 located in the MET onco gene is associated with bladder cancer tumorogenesis,
295:
New insertions create short target site duplications (TSDs), and the majority of new inserts are severely 5’-truncated (average insert size of 900bp in humans) and often inverted (Szak et al., 2002). Because they lack their 5’UTR, most of new inserts are non functional.
87:, and encode one or two proteins, ORF1 and ORF2. The functional domains present within ORF1 vary greatly among LINEs, but often exhibit RNA/DNA binding activity. ORF2 is essential to successful retrotransposition, and encodes a protein with both
131:
Based on structural features and the phylogeny of the essential protein ORF2p, LINEs can be separated into six main groups, referred to as R2, RanI, L1, RTE, I and Jockey. These groups can further be subdivided into at least 28 clades.
429:
Huang X, Su G, Wang Z, Shangguan S, Cui X, Zhu J, et al. (March 2014). "Hypomethylation of long interspersed nucleotide element-1 in peripheral mononuclear cells of juvenile systemic lupus erythematosus patients in China".
135:
In plant genomes, so far only LINEs of the L1 and RTE clade have been reported. Whereas L1 elements diversify into several subclades, RTE-type LINEs are highly conserved, often constituting a single family.
2015:
Kazazian HH, Wong C, Youssoufian H, Scott AF, Phillips DG, Antonarakis SE (March 1988). "Haemophilia A resulting from de novo insertion of L1 sequences represents a novel mechanism for mutation in man".
1823:
Luan DD, Korman MH, Jakubczak JL, Eickbush TH (February 1993). "Reverse transcription of R2Bm RNA is primed by a nick at the chromosomal target site: a mechanism for non-LTR retrotransposition".
2679:
154:
domain is occasionally present. Except for the evolutionary ancient R2 and RTE superfamilies, LINEs usually encode for another protein named ORF1, which may contain an
343:. There are nearly 100 examples of known diseases caused by retroelement insertions, including some types of cancer and neurological disorders. Correlation between
331:
mediated restriction of LINE-1 elements were reported and it is due to the interaction between A3C with the ORF1p that affects the reverse transcriptase activity.
237:
fair amount of variation and some individuals may contain a larger number of active L1 elements, making these individuals more prone to L1-induced mutagenesis.
2736:
2761:
2711:
2771:
2382:"Regulation of L1 expression and retrotransposition by melatonin and its receptor: implications for cancer risk associated with light exposure at night"
244:
copy numbers have also been found in the brains of people with schizophrenia, indicating that LINE elements may play a role in some neuronal diseases.
2893:
2811:
106:, with approximately 20.7% of the sequences identified as being derived from LINEs. The only active lineage of LINE found within humans belongs to the
44:. Bottom: proposed structure of L1 RNA-protein (RNP) complexes. ORF1 proteins form trimers, exhibiting RNA binding and nucleic acid chaperone activity.
2801:
2716:
1968:"Human LINE-1 restriction by APOBEC3C is deaminase independent and mediated by an ORF1p interaction that affects LINE reverse transcriptase activity"
1866:
Yang N, Kazazian HH (September 2006). "L1 retrotransposition is suppressed by endogenously encoded small interfering RNAs in human cultured cells".
1625:"Sequencing, identification and mapping of primed L1 elements (SIMPLE) reveals significant variation in full length L1 elements between individuals"
968:"BNR - a LINE family from Beta vulgaris - contains a RRM domain in open reading frame 1 and defines a L1 sub-clade present in diverse plant genomes"
2329:
Spadafora C (April 2015). "A LINE-1-encoded reverse transcriptase-dependent regulatory mechanism is active in embryogenesis and tumorigenesis".
139:
In fungi, Tad, L1, CRE, Deceiver and Inkcap-like elements have been identified, with Tad-like elements appearing exclusively in fungal genomes.
224:
In the first human genome draft the fraction of LINE elements of the human genome was given as 21% and their copy number as 850,000. Of these,
2721:
2447:
2816:
176:
in plants, fungi or insects, but are dominant in vertebrates and especially in mammals, where they represent around 20% of the genome.
304:
It has been shown that host cells regulate L1 retrotransposition activity, for example through epigenetic silencing. For example, the
2781:
2731:
2726:
2756:
2706:
2751:
2656:
2776:
263:
LINE elements propagate by a so-called target primed reverse transcription mechanism (TPRT), which was first described for the
284:(RNP) complex, likely composed of two ORF2s and an unknown number of ORF1 trimers. The complex is transported back into the
2633:
2620:
524:
Singer MF (March 1982). "SINEs and LINEs: highly repeated short and long interspersed sequences in mammalian genomes".
229:
142:
All LINEs encode a least one protein, ORF2, which contains an RT and an endonuclease (EN) domain, either an N-terminal
41:
2946:
1525:
2280:"Hypomethylation of a LINE-1 promoter activates an alternate transcript of the MET oncogene in bladders with cancer"
919:"Simple and fast classification of non-LTR retrotransposons based on phylogeny of their RT domain protein sequences"
2936:
2440:
2221:"LINE-1 hypomethylation in cancer is highly variable and inversely correlated with microsatellite instability"
1140:"Acquisition of an Archaea-like ribonuclease H domain by plant L1 retrotransposons supports modular evolution"
2674:
2520:
2507:
2425:
1584:"The RTE class of non-LTR retrotransposons is widely distributed in animals and is the origin of many SINEs"
194:-element is one of the elements that are still active in the human genome today. It is found in all therian
2941:
2791:
2433:
119:
The first description of an approximately 6.4 kb long LINE-derived sequence was published by J. Adams
352:
147:
867:"A family of long reiterated DNA sequences, one copy of which is next to the human beta globin gene"
1050:"Two-step regulation and continuous retrotransposition of the rice LINE-type retrotransposon Karma"
2172:"Long interspersed nuclear element-1 hypomethylation in cancer: biology and clinical applications"
1674:
Bundo M, Toyoshima M, Okada Y, Akamatsu W, Ueda J, Nemoto-Miyauchi T, et al. (January 2014).
1097:
Heitkam T, Holtgräwe D, Dohm JC, Minoche AE, Himmelbauer H, Weisshaar B, Schmidt T (August 2014).
228:, L2 and L3 elements made up 516,000, 315,000 and 37,000 copies, respectively. The non-autonomous
2485:
2460:
340:
80:
816:
Sheen FM, Sherry ST, Risch GM, Robichaux M, Nasidze I, Stoneking M, et al. (October 2000).
2915:
2843:
1766:"Sequence patterns indicate an enzymatic involvement in integration of mammalian retroposons"
1009:"Evolutionary dynamics and evolutionary history in the RTE clade of non-LTR retrotransposons"
309:
159:
88:
21:
1909:
Ong-Abdullah M, Ordway JM, Jiang N, Ooi SE, Kok SY, Sarpan N, et al. (September 2015).
1449:
Lander ES, Linton LM, Birren B, Nusbaum C, Zody MC, Baldwin J, et al. (February 2001).
1099:"Profiling of extensively diversified plant LINEs reveals distinct plant-specific subclades"
382:"Whole-genome resequencing allows detection of many rare LINE-1 insertion alleles in humans"
2888:
2602:
2464:
2338:
2232:
2025:
1922:
1911:"Loss of Karma transposon methylation underlies the mantled somaclonal variant of oil palm"
1777:
1462:
1302:
1151:
770:
713:
700:
Nurk S, Koren S, Rhie A, Rautiainen M, Bzikadze AV, Mikheenko A, et al. (April 2022).
604:"Human L1 retrotransposon encodes a conserved endonuclease required for retrotransposition"
99:
64:
2380:
deHaro D, Kines KJ, Sokolowski M, Dauchy RT, Streva VA, Hill SM, et al. (July 2014).
1966:
Horn AV, Klawitter S, Held U, Berger A, Vasudevan AA, Bock A, et al. (January 2014).
1600:
1583:
1559:
1542:
1258:
1241:
1025:
1008:
8:
2666:
2537:
818:"Reading between the LINEs: human genomic variation induced by LINE-1 retrotransposition"
173:
2342:
2278:
Wolff EM, Byun HM, Han HF, Sharma S, Nichols PW, Siegmund KD, et al. (April 2010).
2236:
2029:
1926:
1781:
1543:"Evolutionary dynamics in a novel L2 clade of non-LTR retrotransposons in Deuterostomia"
1466:
1306:
1155:
774:
717:
2838:
2746:
2565:
2406:
2381:
2362:
2306:
2279:
2255:
2220:
2196:
2171:
2144:
2119:
2095:
2068:
2049:
1992:
1967:
1943:
1910:
1891:
1848:
1741:
1716:
1651:
1624:
1508:
Kapitonov VV, Pavlicek A, Jurka J (January 2006). "Anthology of Human
Repetitive DNA".
1423:
1398:
1374:
1347:
1323:
1290:
1222:
1174:
1139:
943:
918:
734:
701:
677:
652:
633:
549:
501:
474:
455:
406:
381:
360:
1074:
1049:
891:
866:
842:
817:
653:"The diversity of retrotransposons and the properties of their reverse transcriptases"
620:
603:
580:
2411:
2354:
2311:
2260:
2201:
2149:
2100:
2041:
1997:
1948:
1883:
1840:
1836:
1805:
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1765:
1746:
1697:
1656:
1605:
1564:
1521:
1517:
1490:
1428:
1379:
1328:
1263:
1214:
1179:
1120:
1079:
1030:
989:
984:
967:
948:
896:
847:
798:
793:
758:
739:
682:
625:
584:
541:
537:
506:
447:
411:
281:
276:
ORF2 (and ORF1 when present) proteins primarily associate in cis with their encoding
155:
2366:
2219:
Estécio MR, Gharibyan V, Shen L, Ibrahim AE, Doshi K, He R, et al. (May 2007).
1895:
1852:
1399:"The genome sizes of megabats (Chiroptera: Pteropodidae) are remarkably constrained"
1226:
637:
553:
2900:
2830:
2468:
2401:
2393:
2346:
2301:
2291:
2250:
2240:
2191:
2183:
2139:
2131:
2090:
2080:
2053:
2033:
1987:
1979:
1938:
1930:
1875:
1832:
1795:
1785:
1736:
1728:
1687:
1646:
1636:
1595:
1554:
1513:
1480:
1470:
1418:
1410:
1369:
1359:
1318:
1310:
1253:
1206:
1169:
1159:
1110:
1069:
1061:
1020:
979:
938:
930:
886:
878:
837:
829:
788:
778:
729:
721:
672:
668:
664:
615:
576:
533:
496:
486:
459:
439:
401:
393:
1197:
Novikova O, Fet V, Blinov A (February 2009). "Non-LTR retrotransposons in fungi".
2611:
2583:
2552:
2547:
2296:
2245:
1692:
1675:
1623:
Streva VA, Jordan VE, Linker S, Hedges DJ, Batzer MA, Deininger PL (March 2015).
491:
355:
is associated with increased cancer risk because light exposure at night reduces
168:
143:
68:
339:
A historic example of L1-conferred disease is
Haemophilia A, which is caused by
2883:
2741:
2698:
2526:
2516:
1770:
Proceedings of the
National Academy of Sciences of the United States of America
1144:
Proceedings of the
National Academy of Sciences of the United States of America
934:
763:
Proceedings of the
National Academy of Sciences of the United States of America
475:"Long interspersed element-1 (LINE-1): passenger or driver in human neoplasms?"
264:
151:
76:
16:
2187:
1641:
1364:
1210:
865:
Adams JW, Kaufman RE, Kretschmer PJ, Harrison M, Nienhuis AW (December 1980).
2930:
2848:
2588:
2511:
2502:
2494:
2455:
1715:
Babushok DV, Ostertag EM, Courtney CE, Choi JM, Kazazian HH (February 2006).
1286:
783:
277:
247:
84:
1164:
882:
725:
443:
111:
LINE DNA sequences can be used to date transposon insertions in the genome.
2860:
2415:
2358:
2315:
2264:
2205:
2153:
2104:
2001:
1952:
1887:
1790:
1750:
1701:
1660:
1568:
1494:
1432:
1414:
1383:
1332:
1267:
1218:
1183:
1124:
1083:
1034:
993:
952:
851:
743:
686:
510:
451:
415:
347:
mobilization and oncogenesis has been reported for epithelial cell cancer (
285:
103:
92:
25:
2045:
1844:
1809:
1609:
900:
802:
629:
588:
545:
397:
2855:
2766:
2628:
2397:
1983:
1676:"Increased l1 retrotransposition in the neuronal genome in schizophrenia"
1485:
1282:
269:
1934:
1314:
1291:"Genome analysis of the platypus reveals unique signatures of evolution"
1138:
Smyshlyaev G, Voigt F, Blinov A, Barabas O, Novikova O (December 2013).
2867:
2684:
2350:
2135:
1732:
1065:
72:
1115:
1098:
833:
2786:
2477:
2037:
1879:
1475:
1450:
356:
348:
172:), and/or an esterase. LINE elements are relatively rare compared to
2910:
2796:
2560:
2456:
1348:"Horizontal transfer of BovB and L1 retrotransposons in eukaryotes"
567:
Jurka J (June 1998). "Repeats in genomic DNA: mining and meaning".
328:
163:
36:
2085:
1280:
2806:
1345:
199:
1346:
Ivancevic AM, Kortschak RD, Bertozzi T, Adelson DL (July 2018).
2905:
2069:"Mobile elements in the human genome: implications for disease"
195:
864:
2651:
1510:
Encyclopedia of
Molecular Cell Biology and Molecular Medicine
1242:"The age and evolution of non-LTR retrotransposable elements"
344:
317:
313:
241:
233:
225:
191:
185:
107:
29:
2014:
1714:
1137:
2379:
2218:
1822:
1673:
1096:
305:
289:
2117:
1908:
1448:
815:
602:
Feng Q, Moran JV, Kazazian HH, Boeke JD (November 1996).
2120:"L1 retrotransposons, cancer stem cells and oncogenesis"
2118:
Carreira PE, Richardson SR, Faulkner GJ (January 2014).
1622:
1965:
1047:
699:
252:
Mechanism of target-primed reverse transcription (TPRT)
2169:
1507:
1006:
650:
601:
1451:"Initial sequencing and analysis of the human genome"
359:, a hormone that has been shown to reduce L1-induced
916:
428:
1540:
2277:
1239:
1196:
1048:Komatsu M, Shimamoto K, Kyozuka J (August 2003).
1007:Zupunski V, Gubensek F, Kordis D (October 2001).
917:Kapitonov VV, Tempel S, Jurka J (December 2009).
2928:
1541:Lovsin N, Gubensek F, Kordi D (December 2001).
912:
910:
759:"Translation of the human LINE-1 element, L1Hs"
2066:
965:
756:
299:
2441:
1581:
1444:
1442:
1240:Malik HS, Burke WD, Eickbush TH (June 1999).
379:
2373:
2170:Kitkumthorn N, Mutirangura A (August 2011).
2111:
1865:
1717:"L1 integration in a transgenic mouse model"
1396:
907:
651:Eickbush TH, Jamburuthugoda VK (June 2008).
1281:Warren WC, Hillier LW, Marshall Graves JA,
432:International Journal of Rheumatic Diseases
334:
2448:
2434:
2331:Annals of the New York Academy of Sciences
1439:
472:
126:
71:that are widespread in the genome of many
2405:
2328:
2305:
2295:
2254:
2244:
2195:
2143:
2094:
2084:
1991:
1942:
1868:Nature Structural & Molecular Biology
1799:
1789:
1740:
1691:
1650:
1640:
1599:
1558:
1484:
1474:
1422:
1373:
1363:
1322:
1257:
1173:
1163:
1114:
1073:
1024:
983:
942:
890:
858:
841:
792:
782:
733:
702:"The complete sequence of a human genome"
676:
619:
517:
500:
490:
405:
1816:
1582:Malik HS, Eickbush TH (September 1998).
1512:. Wiley-VCH Verlag GmbH & Co. KGaA.
1274:
757:McMillan JP, Singer MF (December 1993).
560:
246:
35:
15:
2067:Solyom S, Kazazian HH (February 2012).
966:Heitkam T, Schmidt T (September 2009).
114:
2929:
1289:, GrĂĽtzner F, et al. (May 2008).
523:
40:Genetic structure of murine LINE1 and
2429:
2165:
2163:
1763:
1601:10.1093/oxfordjournals.molbev.a026020
1560:10.1093/oxfordjournals.molbev.a003768
1259:10.1093/oxfordjournals.molbev.a026164
1199:Functional & Integrative Genomics
1026:10.1093/oxfordjournals.molbev.a003727
569:Current Opinion in Structural Biology
566:
57:long interspersed nucleotide elements
380:Ewing AD, Kazazian HH (June 2011).
316:sequences can cause suppression of
13:
2517:Short tandem repeat/Microsatellite
2271:
2212:
2160:
1397:Smith JD, Gregory TR (June 2009).
49:Long interspersed nuclear elements
14:
2958:
205:
1518:10.1002/3527600906.mcb.200300166
985:10.1111/j.1365-313x.2009.03923.x
473:Rodić N, Burns KH (March 2013).
2322:
2060:
2008:
1959:
1902:
1859:
1757:
1708:
1667:
1616:
1588:Molecular Biology and Evolution
1575:
1547:Molecular Biology and Evolution
1534:
1501:
1390:
1339:
1246:Molecular Biology and Evolution
1233:
1190:
1131:
1090:
1041:
1013:Molecular Biology and Evolution
1000:
959:
809:
2521:Trinucleotide repeat disorders
750:
693:
669:10.1016/j.virusres.2007.12.010
644:
595:
466:
422:
373:
258:
179:
1:
2508:Variable number tandem repeat
621:10.1016/s0092-8674(00)81997-2
581:10.1016/S0959-440X(98)80067-5
366:
2297:10.1371/journal.pgen.1000917
2246:10.1371/journal.pone.0000399
1837:10.1016/0092-8674(93)90078-5
1693:10.1016/j.neuron.2013.10.053
538:10.1016/0092-8674(82)90194-5
492:10.1371/journal.pgen.1003402
214:
98:LINEs are the most abundant
75:. LINEs contain an internal
7:
300:Regulation of LINE activity
219:
10:
2963:
935:10.1016/j.gene.2009.07.019
267:element from the silkworm
183:
63:) are a group of non-LTR (
61:long interspersed elements
2876:
2829:
2697:
2665:
2642:
2619:
2610:
2601:
2576:
2536:
2493:
2484:
2475:
2188:10.1007/s13148-011-0032-8
1642:10.1186/s12864-015-1374-y
1365:10.1186/s13059-018-1456-7
1211:10.1007/s10142-008-0093-8
353:Shift work sleep disorder
232:elements which depend on
20:ORF2 protein (exhibiting
2947:Repetitive DNA sequences
784:10.1073/pnas.90.24.11533
335:Association with disease
2937:Mobile genetic elements
1165:10.1073/pnas.1310958110
726:10.1126/science.abj6987
444:10.1111/1756-185X.12239
341:insertional mutagenesis
306:RNA interference (RNAi)
127:Classification of LINEs
2916:Protein tandem repeats
2844:Tandemly arrayed genes
2386:Nucleic Acids Research
1972:Nucleic Acids Research
1791:10.1073/pnas.94.5.1872
1764:Jurka J (March 1997).
1415:10.1098/rsbl.2009.0016
871:Nucleic Acids Research
310:small interfering RNAs
255:
45:
33:
883:10.1093/nar/8.24.6113
398:10.1101/gr.114777.110
250:
89:reverse transcriptase
79:promoter to initiate
39:
28:activity) from human
22:reverse transcriptase
19:
2889:Pathogenicity island
2176:Clinical Epigenetics
320:retrotransposition.
174:LTR-retrotransposons
115:History of discovery
100:transposable element
65:long terminal repeat
2343:2015NYASA1341..164S
2237:2007PLoSO...2..399E
2030:1988Natur.332..164K
1935:10.1038/nature15365
1927:2015Natur.525..533O
1782:1997PNAS...94.1872J
1467:2001Natur.409..860L
1315:10.1038/nature06936
1307:2008Natur.453..175W
1156:2013PNAS..11020140S
1150:(50): 20140–20145.
775:1993PNAS...9011533M
769:(24): 11533–11537.
718:2022Sci...376...44N
211:CR1 clade, Jockey.
2839:Gene amplification
2398:10.1093/nar/gku503
2351:10.1111/nyas.12637
2136:10.1111/febs.12601
1984:10.1093/nar/gkt898
1733:10.1101/gr.4571606
1066:10.1105/tpc.011809
361:genome instability
256:
150:or rarely both. A
46:
34:
2942:Molecular biology
2924:
2923:
2825:
2824:
2693:
2692:
2597:
2596:
2486:Repeated sequence
2461:repeated sequence
2392:(12): 7694–7707.
2024:(6160): 164–166.
1921:(7570): 533–537.
1553:(12): 2213–2224.
1461:(6822): 860–921.
1301:(7192): 175–183.
1116:10.1111/tpj.12565
1103:The Plant Journal
1019:(10): 1849–1863.
972:The Plant Journal
877:(24): 6113–6128.
834:10.1101/gr.149400
828:(10): 1496–1508.
282:ribonucleoprotein
55:) (also known as
2954:
2901:Low copy repeats
2894:Symbiosis island
2831:Gene duplication
2617:
2616:
2608:
2607:
2491:
2490:
2469:gene duplication
2450:
2443:
2436:
2427:
2426:
2420:
2419:
2409:
2377:
2371:
2370:
2326:
2320:
2319:
2309:
2299:
2275:
2269:
2268:
2258:
2248:
2216:
2210:
2209:
2199:
2167:
2158:
2157:
2147:
2124:The FEBS Journal
2115:
2109:
2108:
2098:
2088:
2064:
2058:
2057:
2038:10.1038/332164a0
2012:
2006:
2005:
1995:
1963:
1957:
1956:
1946:
1906:
1900:
1899:
1880:10.1038/nsmb1141
1863:
1857:
1856:
1820:
1814:
1813:
1803:
1793:
1776:(5): 1872–1877.
1761:
1755:
1754:
1744:
1712:
1706:
1705:
1695:
1671:
1665:
1664:
1654:
1644:
1620:
1614:
1613:
1603:
1594:(9): 1123–1134.
1579:
1573:
1572:
1562:
1538:
1532:
1531:
1505:
1499:
1498:
1488:
1478:
1476:10.1038/35057062
1446:
1437:
1436:
1426:
1394:
1388:
1387:
1377:
1367:
1343:
1337:
1336:
1326:
1278:
1272:
1271:
1261:
1237:
1231:
1230:
1194:
1188:
1187:
1177:
1167:
1135:
1129:
1128:
1118:
1094:
1088:
1087:
1077:
1060:(8): 1934–1944.
1045:
1039:
1038:
1028:
1004:
998:
997:
987:
963:
957:
956:
946:
914:
905:
904:
894:
862:
856:
855:
845:
813:
807:
806:
796:
786:
754:
748:
747:
737:
697:
691:
690:
680:
663:(1–2): 221–234.
648:
642:
641:
623:
599:
593:
592:
564:
558:
557:
521:
515:
514:
504:
494:
470:
464:
463:
426:
420:
419:
409:
377:
146:or a C-terminal
69:retrotransposons
2962:
2961:
2957:
2956:
2955:
2953:
2952:
2951:
2927:
2926:
2925:
2920:
2872:
2821:
2689:
2661:
2638:
2612:Retrotransposon
2593:
2584:Inverted repeat
2572:
2557:DNA transposon
2553:Retrotransposon
2548:Gene conversion
2539:
2532:
2529:
2480:
2471:
2454:
2424:
2423:
2378:
2374:
2327:
2323:
2290:(4): e1000917.
2276:
2272:
2217:
2213:
2168:
2161:
2116:
2112:
2073:Genome Medicine
2065:
2061:
2013:
2009:
1964:
1960:
1907:
1903:
1864:
1860:
1821:
1817:
1762:
1758:
1721:Genome Research
1713:
1709:
1672:
1668:
1621:
1617:
1580:
1576:
1539:
1535:
1528:
1506:
1502:
1447:
1440:
1403:Biology Letters
1395:
1391:
1344:
1340:
1279:
1275:
1238:
1234:
1195:
1191:
1136:
1132:
1095:
1091:
1046:
1042:
1005:
1001:
964:
960:
915:
908:
863:
859:
822:Genome Research
814:
810:
755:
751:
712:(6588): 44–53.
698:
694:
649:
645:
600:
596:
565:
561:
522:
518:
485:(3): e1003402.
471:
467:
427:
423:
386:Genome Research
378:
374:
369:
337:
302:
261:
222:
217:
208:
188:
182:
129:
117:
12:
11:
5:
2960:
2950:
2949:
2944:
2939:
2922:
2921:
2919:
2918:
2913:
2908:
2903:
2898:
2897:
2896:
2891:
2884:Genomic island
2880:
2878:
2874:
2873:
2871:
2870:
2865:
2864:
2863:
2853:
2852:
2851:
2841:
2835:
2833:
2827:
2826:
2823:
2822:
2820:
2819:
2814:
2809:
2804:
2799:
2794:
2789:
2784:
2779:
2774:
2769:
2764:
2759:
2754:
2749:
2744:
2739:
2734:
2729:
2724:
2719:
2714:
2709:
2703:
2701:
2699:DNA transposon
2695:
2694:
2691:
2690:
2688:
2687:
2682:
2677:
2671:
2669:
2663:
2662:
2660:
2659:
2654:
2648:
2646:
2640:
2639:
2637:
2636:
2631:
2625:
2623:
2614:
2605:
2599:
2598:
2595:
2594:
2592:
2591:
2586:
2580:
2578:
2574:
2573:
2571:
2570:
2569:
2568:
2563:
2555:
2550:
2544:
2542:
2534:
2533:
2531:
2530:
2527:Macrosatellite
2524:
2514:
2505:
2499:
2497:
2495:Tandem repeats
2488:
2482:
2481:
2476:
2473:
2472:
2453:
2452:
2445:
2438:
2430:
2422:
2421:
2372:
2337:(1): 164–171.
2321:
2270:
2211:
2182:(2): 315–330.
2159:
2110:
2059:
2007:
1978:(1): 396–416.
1958:
1901:
1874:(9): 763–771.
1858:
1831:(4): 595–605.
1815:
1756:
1727:(2): 240–250.
1707:
1686:(2): 306–313.
1666:
1615:
1574:
1533:
1526:
1500:
1438:
1409:(3): 347–351.
1389:
1352:Genome Biology
1338:
1273:
1252:(6): 793–805.
1232:
1189:
1130:
1109:(3): 385–397.
1089:
1054:The Plant Cell
1040:
999:
978:(6): 872–882.
958:
929:(2): 207–213.
906:
857:
808:
749:
692:
657:Virus Research
643:
614:(5): 905–916.
594:
575:(3): 333–337.
559:
532:(3): 433–434.
516:
465:
438:(3): 280–290.
421:
392:(6): 985–990.
371:
370:
368:
365:
336:
333:
301:
298:
292:is integrated
260:
257:
221:
218:
216:
213:
207:
206:Other elements
204:
184:Main article:
181:
178:
152:ribonuclease H
128:
125:
116:
113:
9:
6:
4:
3:
2:
2959:
2948:
2945:
2943:
2940:
2938:
2935:
2934:
2932:
2917:
2914:
2912:
2909:
2907:
2904:
2902:
2899:
2895:
2892:
2890:
2887:
2886:
2885:
2882:
2881:
2879:
2875:
2869:
2866:
2862:
2859:
2858:
2857:
2854:
2850:
2849:Ribosomal DNA
2847:
2846:
2845:
2842:
2840:
2837:
2836:
2834:
2832:
2828:
2818:
2815:
2813:
2810:
2808:
2805:
2803:
2800:
2798:
2795:
2793:
2790:
2788:
2785:
2783:
2780:
2778:
2775:
2773:
2770:
2768:
2765:
2763:
2760:
2758:
2755:
2753:
2750:
2748:
2745:
2743:
2740:
2738:
2735:
2733:
2730:
2728:
2725:
2723:
2720:
2718:
2715:
2713:
2710:
2708:
2705:
2704:
2702:
2700:
2696:
2686:
2683:
2681:
2678:
2676:
2673:
2672:
2670:
2668:
2664:
2658:
2655:
2653:
2650:
2649:
2647:
2645:
2641:
2635:
2632:
2630:
2627:
2626:
2624:
2622:
2618:
2615:
2613:
2609:
2606:
2604:
2600:
2590:
2589:Direct repeat
2587:
2585:
2582:
2581:
2579:
2575:
2567:
2564:
2562:
2559:
2558:
2556:
2554:
2551:
2549:
2546:
2545:
2543:
2541:
2535:
2528:
2525:
2522:
2518:
2515:
2513:
2512:Minisatellite
2509:
2506:
2504:
2503:Satellite DNA
2501:
2500:
2498:
2496:
2492:
2489:
2487:
2483:
2479:
2474:
2470:
2466:
2462:
2458:
2451:
2446:
2444:
2439:
2437:
2432:
2431:
2428:
2417:
2413:
2408:
2403:
2399:
2395:
2391:
2387:
2383:
2376:
2368:
2364:
2360:
2356:
2352:
2348:
2344:
2340:
2336:
2332:
2325:
2317:
2313:
2308:
2303:
2298:
2293:
2289:
2285:
2284:PLOS Genetics
2281:
2274:
2266:
2262:
2257:
2252:
2247:
2242:
2238:
2234:
2230:
2226:
2222:
2215:
2207:
2203:
2198:
2193:
2189:
2185:
2181:
2177:
2173:
2166:
2164:
2155:
2151:
2146:
2141:
2137:
2133:
2129:
2125:
2121:
2114:
2106:
2102:
2097:
2092:
2087:
2086:10.1186/gm311
2082:
2078:
2074:
2070:
2063:
2055:
2051:
2047:
2043:
2039:
2035:
2031:
2027:
2023:
2019:
2011:
2003:
1999:
1994:
1989:
1985:
1981:
1977:
1973:
1969:
1962:
1954:
1950:
1945:
1940:
1936:
1932:
1928:
1924:
1920:
1916:
1912:
1905:
1897:
1893:
1889:
1885:
1881:
1877:
1873:
1869:
1862:
1854:
1850:
1846:
1842:
1838:
1834:
1830:
1826:
1819:
1811:
1807:
1802:
1797:
1792:
1787:
1783:
1779:
1775:
1771:
1767:
1760:
1752:
1748:
1743:
1738:
1734:
1730:
1726:
1722:
1718:
1711:
1703:
1699:
1694:
1689:
1685:
1681:
1677:
1670:
1662:
1658:
1653:
1648:
1643:
1638:
1634:
1630:
1626:
1619:
1611:
1607:
1602:
1597:
1593:
1589:
1585:
1578:
1570:
1566:
1561:
1556:
1552:
1548:
1544:
1537:
1529:
1527:9783527600908
1523:
1519:
1515:
1511:
1504:
1496:
1492:
1487:
1486:2027.42/62798
1482:
1477:
1472:
1468:
1464:
1460:
1456:
1452:
1445:
1443:
1434:
1430:
1425:
1420:
1416:
1412:
1408:
1404:
1400:
1393:
1385:
1381:
1376:
1371:
1366:
1361:
1357:
1353:
1349:
1342:
1334:
1330:
1325:
1320:
1316:
1312:
1308:
1304:
1300:
1296:
1292:
1288:
1284:
1277:
1269:
1265:
1260:
1255:
1251:
1247:
1243:
1236:
1228:
1224:
1220:
1216:
1212:
1208:
1204:
1200:
1193:
1185:
1181:
1176:
1171:
1166:
1161:
1157:
1153:
1149:
1145:
1141:
1134:
1126:
1122:
1117:
1112:
1108:
1104:
1100:
1093:
1085:
1081:
1076:
1071:
1067:
1063:
1059:
1055:
1051:
1044:
1036:
1032:
1027:
1022:
1018:
1014:
1010:
1003:
995:
991:
986:
981:
977:
973:
969:
962:
954:
950:
945:
940:
936:
932:
928:
924:
920:
913:
911:
902:
898:
893:
888:
884:
880:
876:
872:
868:
861:
853:
849:
844:
839:
835:
831:
827:
823:
819:
812:
804:
800:
795:
790:
785:
780:
776:
772:
768:
764:
760:
753:
745:
741:
736:
731:
727:
723:
719:
715:
711:
707:
703:
696:
688:
684:
679:
674:
670:
666:
662:
658:
654:
647:
639:
635:
631:
627:
622:
617:
613:
609:
605:
598:
590:
586:
582:
578:
574:
570:
563:
555:
551:
547:
543:
539:
535:
531:
527:
520:
512:
508:
503:
498:
493:
488:
484:
480:
479:PLOS Genetics
476:
469:
461:
457:
453:
449:
445:
441:
437:
433:
425:
417:
413:
408:
403:
399:
395:
391:
387:
383:
376:
372:
364:
362:
358:
354:
350:
346:
342:
332:
330:
325:
321:
319:
315:
312:derived from
311:
308:mechanism of
307:
297:
293:
291:
287:
283:
279:
274:
273:
271:
266:
253:
249:
245:
243:
238:
235:
231:
227:
212:
203:
201:
197:
193:
187:
177:
175:
171:
170:
165:
161:
157:
153:
149:
145:
140:
137:
133:
124:
122:
112:
109:
105:
101:
96:
94:
90:
86:
82:
81:transcription
78:
74:
70:
66:
62:
58:
54:
50:
43:
38:
31:
27:
23:
18:
2861:Gene cluster
2643:
2629:Alu sequence
2538:Interspersed
2389:
2385:
2375:
2334:
2330:
2324:
2287:
2283:
2273:
2228:
2224:
2214:
2179:
2175:
2130:(1): 63–73.
2127:
2123:
2113:
2076:
2072:
2062:
2021:
2017:
2010:
1975:
1971:
1961:
1918:
1914:
1904:
1871:
1867:
1861:
1828:
1824:
1818:
1773:
1769:
1759:
1724:
1720:
1710:
1683:
1679:
1669:
1632:
1629:BMC Genomics
1628:
1618:
1591:
1587:
1577:
1550:
1546:
1536:
1509:
1503:
1458:
1454:
1406:
1402:
1392:
1355:
1351:
1341:
1298:
1294:
1276:
1249:
1245:
1235:
1205:(1): 27–42.
1202:
1198:
1192:
1147:
1143:
1133:
1106:
1102:
1092:
1057:
1053:
1043:
1016:
1012:
1002:
975:
971:
961:
926:
922:
874:
870:
860:
825:
821:
811:
766:
762:
752:
709:
705:
695:
660:
656:
646:
611:
607:
597:
572:
568:
562:
529:
525:
519:
482:
478:
468:
435:
431:
424:
389:
385:
375:
338:
326:
322:
303:
294:
280:, forming a
275:
268:
262:
251:
239:
223:
209:
189:
167:
141:
138:
134:
130:
120:
118:
104:human genome
97:
93:endonuclease
60:
56:
52:
48:
47:
26:endonuclease
2856:Gene family
2767:Tc1/mariner
2722:EnSpm/CACTA
2231:(5): e399.
270:Bombyx mori
259:Propagation
180:L1 elements
160:L1-like RRM
156:Gag-knuckle
102:within the
95:activity.
2931:Categories
2868:Pseudogene
2685:retroposon
2603:Transposon
2465:transposon
1635:(1): 220.
1287:Ponting CP
367:References
240:Increased
73:eukaryotes
2787:P element
2737:Harbinger
2478:Repeatome
2079:(2): 12.
1358:(1): 85.
357:melatonin
349:carcinoma
215:Incidence
192:LINE-1/L1
169:IPR035300
123:in 1980.
2911:Telomere
2877:See also
2817:Zisupton
2797:Polinton
2792:PiggyBac
2747:Helitron
2566:Helitron
2561:Polinton
2457:Genetics
2416:24914052
2367:22881053
2359:25586649
2316:20421991
2265:17476321
2225:PLOS ONE
2206:22704344
2154:24286172
2105:22364178
2002:24101588
1953:26352475
1896:32601334
1888:16936727
1853:42587840
1751:16365384
1702:24389010
1661:25887476
1569:11719571
1495:11237011
1433:19324635
1384:29983116
1333:18464734
1283:Birney E
1268:10368957
1227:23319640
1219:18677522
1184:24277848
1125:24862340
1084:12897263
1035:11557792
994:19473321
953:19651192
852:11042149
744:35357919
687:18261821
638:17897241
554:22129236
511:23555307
452:24330152
416:20980553
329:APOBEC3C
220:In human
200:megabats
164:InterPro
2807:Transib
2782:Novosib
2762:Kolobok
2732:Ginger2
2727:Ginger1
2712:Crypton
2407:4081101
2339:Bibcode
2307:2858672
2256:1851990
2233:Bibcode
2197:3365388
2145:4160015
2096:3392758
2054:4259071
2046:2831458
2026:Bibcode
1993:3874205
1944:4857894
1923:Bibcode
1845:7679954
1810:9050872
1778:Bibcode
1742:1361720
1652:4381410
1610:9729877
1463:Bibcode
1424:2679926
1375:6036668
1324:2803040
1303:Bibcode
1175:3864347
1152:Bibcode
944:2829327
901:6258162
803:8265584
771:Bibcode
735:9186530
714:Bibcode
706:Science
678:2695964
630:8945517
589:9666329
546:6280868
502:3610623
460:6530689
407:3106331
286:nucleus
198:except
196:mammals
166::
2906:CRISPR
2772:Merlin
2757:ISL2EU
2707:Academ
2540:repeat
2414:
2404:
2365:
2357:
2314:
2304:
2263:
2253:
2204:
2194:
2152:
2142:
2103:
2093:
2052:
2044:
2018:Nature
2000:
1990:
1951:
1941:
1915:Nature
1894:
1886:
1851:
1843:
1808:
1798:
1749:
1739:
1700:
1680:Neuron
1659:
1649:
1608:
1567:
1524:
1493:
1455:Nature
1431:
1421:
1382:
1372:
1331:
1321:
1295:Nature
1266:
1225:
1217:
1182:
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