29:
576:, with particularly extensive insertions found in colorectal and lung cancers. It is currently unclear if these insertions are causes or secondary effects of cancer progression. However, at least two cases have found somatic L1 insertions causative of cancer by disrupting the coding sequences of genes
599:
are used as diagnostic biomarkers in some types of cancers. L1 hypomethylation of colon tumor samples is correlated with cancer stage progression. Furthermore, less invasive blood assays for L1 copy number or methylation levels are indicative of breast or bladder cancer progression and may serve as
700:
mutant cancer cells, which would possibly explain why some patients test PCR positive for SARS-CoV-2 even after clearance of the virus. These results however have been criticized as "mechanistically plausible but likely very rare", misleading and infrequent or artefactual.
629:, which carry MeCP2 mutations, exhibit increased L1 transposition, suggesting a link between L1 activity and neurological disorders. Current studies are aimed at investigating the potential roles of L1 activity in various neuropsychiatric disorders including
516:. The structure of the ORF2 protein was solved in 2023. Its protein core contains three domains of unknown functions, termed "tower/EN-linker" and "wrist/RNA-binding domain" that bind Alu RNA's polyA tail and C-terminal domain that binds Alu RNA stem loop.
1565:
Kaur D, Agrahari M, Singh SS, Mandal PK, Bhattacharya A, Bhattacharya S (March 2021). "Transcriptomic analysis of
Entamoeba histolytica reveals domain-specific sense strand expression of LINE-encoded ORFs with massive antisense expression of RT domain".
153:
monomers of around 200 bp, followed by a short non-monomeric region. Human 5’ UTRs are ~900 bp in length and do not contain repeated motifs. All families of human L1s harbor in their most 5’ extremity a binding motif for the transcription factor
2230:
Chen J, Rattner A, Nathans J (July 2006). "Effects of L1 retrotransposon insertion on transcript processing, localization and accumulation: lessons from the retinal degeneration 7 mouse and implications for the genomic ecology of L1 elements".
612:
compared to other organs. Studies of animal models and human cell lines have shown that L1s become active in neural progenitor cells (NPCs), and that experimental deregulation of or overexpression of L1 increases somatic
2685:
Zheng F, Kawabe Y, Murakami M, Takahashi M, Nishihata K, Yoshida S, et al. (July 2021). "LINE-1 vectors mediate recombinant antibody gene transfer by retrotransposition in
Chinese hamster ovary cells".
1105:
2723:
Jachowicz JW, Bing X, Pontabry J, Bošković A, Rando OJ, Torres-Padilla ME (October 2017). "LINE-1 activation after fertilization regulates global chromatin accessibility in the early mouse embryo".
32:
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:-family transcription factors, and both YY1 and SOX sites were shown to be required for human L1 transcription initiation and activation. Both mouse and human 5’ UTRs also contain a weak
67:(LINEs). L1 transposons are most ubiquitous in mammals, where they make up a significant fraction of the total genome length, for example they comprise approximately 17% of the
3137:
2762:"A comparative analysis of L1 retrotransposition activities in human genomes suggests an ongoing increase in L1 number despite an evolutionary trend towards lower activity"
347:
that mediates the formation of ORF1 trimeric complexes. ORF1 trimers have RNA-binding and nucleic acid chaperone activity that are necessary for retrotransposition.
137:(UTRs) and target site duplications. In humans, ORF2 is thought to be translated by an unconventional termination/reinitiation mechanism, while mouse L1s contain an
1601:
Tubio JM, Li Y, Ju YS, Martincorena I, Cooke SL, Tojo M, et al. (ICGC Breast Cancer Group; ICGC Bone Cancer Group; ICGC Prostate Cancer Group) (August 2014).
1109:
3194:
1230:"The dicistronic RNA from the mouse LINE-1 retrotransposon contains an internal ribosome entry site upstream of each ORF: implications for retrotransposition"
3219:
3169:
549:
As with other transposable elements, the host organism keeps a heavy check on LINE1 to prevent it from becoming overly active. In the primitive eukaryote
3229:
1416:"An antisense promoter in mouse L1 retrotransposon open reading frame-1 initiates expression of diverse fusion transcripts and limits retrotransposition"
1758:
Sunami E, Vu AT, Nguyen SL, Giuliano AE, Hoon DS (August 2008). "Quantification of LINE1 in circulating DNA as a molecular biomarker of breast cancer".
3351:
3269:
3259:
3174:
384:
203:
544:
3179:
677:
2268:"Reverse-transcribed SARS-CoV-2 RNA can integrate into the genome of cultured human cells and can be expressed in patient-derived tissues"
2905:
3274:
3239:
3189:
3184:
832:
Lander ES, Linton LM, Birren B, Nusbaum C, Zody MC, et al. (International Human Genome
Sequencing Consortium) (February 2001).
3214:
3164:
1279:"Binding of the ubiquitous nuclear transcription factor YY1 to a cis regulatory sequence in the human LINE-1 transposable element"
3209:
3114:
3234:
389:
208:
3394:
3399:
2376:"No evidence of SARS-CoV-2 reverse transcription and integration as the origin of chimeric transcripts in patient tissues"
592:
227:
106:
induced by L1 and its non-autonomous counterparts have been found to cause a variety of heritable and somatic diseases.
3091:
3078:
1603:"Mobile DNA in cancer. Extensive transduction of nonrepetitive DNA mediated by L1 retrotransposition in cancer genomes"
99:
75:. L1 activity has contributed to the instability and evolution of genomes and is tightly regulated in the germline by
3414:
3101:
64:
3404:
396:
215:
2898:
535:. Unlike viral RTs, L1 ORF2p can be primed by RNA, including RNA hairpin primers produced by the Alu element.
3132:
2978:
2965:
2599:"RHOX10 drives mouse spermatogonial stem cell establishment through a transcription factor signaling cascade"
2883:
736:"LINEs between Species: Evolutionary Dynamics of LINE-1 Retrotransposons across the Eukaryotic Tree of Life"
138:
653:. L1s are also highly expressed in octopus brain, suggesting a convergent mechanism in complex cognition.
401:
220:
1915:
McConnell MJ, Lindberg MR, Brennand KJ, Piper JC, Voet T, Cowing-Zitron C, et al. (November 2013).
339:
protein that lacks homology with any protein of known function. In vertebrates, it contains a conserved
3409:
3249:
2891:
2083:"Lower LINE-1 methylation in first-episode schizophrenia patients with the history of childhood trauma"
577:
2325:
Smits N, Rasmussen J, Bodea GO, Amarilla AA, Gerdes P, Sanchez-Luque FJ, et al. (August 2021).
114:
2805:
Garcia-Perez JL, Marchetto MC, Muotri AR, Coufal NG, Gage FH, O'Shea KS, et al. (July 2007).
2482:
Penzkofer T, Jäger M, Figlerowicz M, Badge R, Mundlos S, Robinson PN, et al. (January 2017).
2943:
2918:
2433:
Yan B, Chakravorty S, Mirabelli C, Wang L, Trujillo-Ochoa JL, Chauss D, et al. (July 2021).
2124:"Identification of LINE retrotransposons and long non-coding RNAs expressed in the octopus brain"
1650:
Ogino S, Nosho K, Kirkner GJ, Kawasaki T, Chan AT, Schernhammer ES, et al. (December 2008).
676:
The naturally occurring mouse retinal degeneration model rd7 is caused by an L1 insertion in the
20:
3373:
3301:
2024:
Muotri AR, Marchetto MC, Coufal NG, Oefner R, Yeo G, Nakashima K, et al. (November 2010).
71:. These active L1s can interrupt the genome through insertions, deletions, rearrangements, and
1132:"Opportunity and means: horizontal gene transfer from the human host to a bacterial pathogen"
551:
524:
509:
273:
88:
72:
52:
277:
3346:
3060:
2922:
2557:
2536:
Tan K, Kim ME, Song HW, Skarbrevik D, Babajanian E, Bedrosian TA, et al. (June 2021).
2387:
2279:
2186:
2122:
Petrosino G, Ponte G, Volpe M, Zarrella I, Ansaloni F, Langella C, et al. (May 2022).
2037:
1928:
1871:
1767:
1712:
958:
845:
666:
596:
442:
80:
478:
309:
8:
3124:
2995:
2327:"No evidence of human genome integration of SARS-CoV-2 found by long-read DNA sequencing"
1516:"The ORF1 protein encoded by LINE-1: structure and function during L1 retrotransposition"
528:
134:
2561:
2435:"Host-Virus Chimeric Events in SARS-CoV-2-Infected Cells Are Infrequent and Artifactual"
2391:
2283:
2190:
2041:
1932:
1875:
1858:
Coufal NG, Garcia-Perez JL, Peng GE, Yeo GW, Mu Y, Lovci MT, et al. (August 2009).
1771:
1716:
1021:
962:
849:
3296:
3204:
3023:
2866:
2841:
2788:
2761:
2748:
2711:
2668:
2641:
2623:
2598:
2580:
2537:
2508:
2483:
2459:
2434:
2410:
2375:
2351:
2326:
2302:
2267:
2207:
2174:
2173:
Kaneko H, Dridi S, Tarallo V, Gelfand BD, Fowler BJ, Cho WG, et al. (March 2011).
2150:
2123:
2058:
2025:
2001:
1976:
1949:
1916:
1892:
1859:
1835:
1810:
1791:
1735:
1700:
1676:
1651:
1627:
1602:
1542:
1515:
1491:
1464:
1440:
1415:
1254:
1229:
1205:
1180:
1156:
1131:
1081:
1054:
1030:
1005:
981:
946:
922:
897:
809:
784:
760:
735:
689:
501:
446:
332:
130:
1809:
Wilhelm CS, Kelsey KT, Butler R, Plaza S, Gagne L, Zens MS, et al. (March 2010).
1391:
1366:
1342:
1317:
625:
and methylation of the L1 5' UTR. Human cell lines modeling the neurological disorder
2871:
2828:
2793:
2740:
2715:
2703:
2673:
2628:
2585:
2513:
2464:
2415:
2356:
2307:
2248:
2212:
2155:
2104:
2063:
2006:
1954:
1897:
1840:
1783:
1740:
1681:
1632:
1583:
1547:
1496:
1445:
1396:
1347:
1298:
1259:
1210:
1161:
1086:
1035:
986:
927:
873:
814:
765:
646:
614:
408:
239:
1795:
3358:
3288:
2926:
2861:
2853:
2818:
2783:
2773:
2752:
2732:
2695:
2663:
2653:
2618:
2610:
2575:
2565:
2503:
2495:
2454:
2446:
2405:
2395:
2346:
2338:
2297:
2287:
2240:
2202:
2194:
2145:
2135:
2094:
2053:
2045:
1996:
1988:
1944:
1936:
1887:
1879:
1830:
1822:
1775:
1730:
1720:
1671:
1663:
1622:
1614:
1575:
1537:
1527:
1486:
1476:
1435:
1427:
1386:
1378:
1367:"A YY1-binding site is required for accurate human LINE-1 transcription initiation"
1337:
1329:
1290:
1249:
1241:
1200:
1192:
1151:
1143:
1076:
1066:
1025:
1017:
976:
966:
917:
909:
863:
853:
804:
796:
755:
747:
642:
532:
437:
268:
1826:
1579:
3069:
3041:
3010:
3005:
2614:
2342:
1725:
1071:
634:
159:
76:
2081:
Misiak B, Szmida E, Karpiński P, Loska O, Sąsiadek MM, Frydecka D (2015-12-01).
1977:"Mobile DNA elements in the generation of diversity and complexity in the brain"
1652:"A cohort study of tumoral LINE-1 hypomethylation and prognosis in colon cancer"
1055:"The NF1 gene contains hotspots for L1 endonuclease-dependent de novo insertion"
488:
319:
3341:
3199:
3156:
2984:
2974:
2857:
2842:"Homotypic clustering of L1 and B1/Alu repeats compartmentalizes the 3D genome"
2778:
2550:
Proceedings of the
National Academy of Sciences of the United States of America
2380:
Proceedings of the
National Academy of Sciences of the United States of America
2272:
Proceedings of the
National Academy of Sciences of the United States of America
2140:
1532:
1481:
951:
Proceedings of the
National Academy of Sciences of the United States of America
713:, a database of functional annotations and predictions of active LINE1 elements
3388:
3306:
3046:
2969:
2960:
2952:
2913:
2266:
Zhang L, Richards A, Barrasa MI, Hughes SH, Young RA, Jaenisch R (May 2021).
2175:"DICER1 deficit induces Alu RNA toxicity in age-related macular degeneration"
1414:
Li J, Kannan M, Trivett AL, Liao H, Wu X, Akagi K, et al. (April 2014).
630:
626:
556:
150:
2570:
2400:
2292:
1940:
1779:
1618:
1294:
971:
688:
In 2021, a study proposed that L1 elements may be responsible for potential
3318:
2875:
2832:
2797:
2744:
2707:
2699:
2677:
2632:
2589:
2517:
2468:
2419:
2360:
2311:
2252:
2216:
2159:
2108:
2067:
2010:
1958:
1901:
1844:
1787:
1744:
1685:
1636:
1587:
1551:
1500:
1465:"L1 antisense promoter drives tissue-specific transcription of human genes"
1449:
1400:
1351:
1333:
1263:
1214:
1165:
1090:
1039:
990:
931:
877:
818:
769:
520:
505:
68:
2658:
2484:"L1Base 2: more retrotransposition-active LINE-1s, more mammalian genomes"
1667:
1302:
1147:
913:
459:
290:
3313:
3224:
3086:
2823:
2806:
2499:
2450:
2244:
1431:
1382:
1245:
1179:
Alisch RS, Garcia-Perez JL, Muotri AR, Gage FH, Moran JV (January 2006).
868:
800:
751:
662:
585:
581:
425:
256:
95:
2198:
2049:
1883:
1196:
665:, which requires L1 proteins, are associated with a form of age-related
3325:
3142:
693:
513:
420:
344:
340:
251:
2840:
Lu JY, Chang L, Li T, Wang T, Yin Y, Zhan G, et al. (June 2021).
2099:
2082:
413:
244:
3244:
2935:
858:
833:
650:
163:
126:
110:
60:
2736:
1992:
1811:"Implications of LINE1 methylation for bladder cancer risk in women"
1318:"Members of the SRY family regulate the human LINE retrotransposons"
1278:
734:
Ivancevic AM, Kortschak RD, Bertozzi T, Adelson DL (December 2016).
545:
Long interspersed nuclear element § Regulation of LINE activity
3368:
3254:
3018:
2914:
1106:"Gonorrhea has picked up human DNA (and that's just the beginning)"
638:
454:
285:
103:
28:
19:
This article is about long interspersed element 1. For other, see
3264:
2804:
1181:"Unconventional translation of mammalian LINE-1 retrotransposons"
733:
109:
In 2011, human L1 was reportedly discovered in the genome of the
87:. L1s can further impact genome variation through mispairing and
1701:"LINE-1 hypomethylation during primary colon cancer progression"
232:
3363:
1228:
Li PW, Li J, Timmerman SL, Krushel LA, Martin SL (2006-01-01).
710:
573:
2481:
2374:
Parry R, Gifford RJ, Lytras S, Ray SC, Coin LJ (August 2021).
1914:
1053:
Wimmer K, Callens T, Wernstedt A, Messiaen L (November 2011).
1052:
149:
The 5' UTRs of mouse L1s contain a variable number of GC-rich
2722:
622:
609:
559:, resulting in no detectable amounts of its protein product.
84:
2684:
1699:
Sunami E, de Maat M, Vu A, Turner RR, Hoon DS (April 2011).
1178:
512:
activity. The encoded protein has a molecular weight of 150
2642:"Factors Regulating the Activity of LINE1 Retrotransposons"
2432:
2324:
2121:
2023:
1277:
Becker KG, Swergold GD, Ozato K, Thayer RE (October 1993).
697:
618:
2265:
2080:
1003:
94:
L1 gene products are also required by many non-autonomous
2807:"LINE-1 retrotransposition in human embryonic stem cells"
2172:
1857:
1649:
1276:
831:
670:
336:
155:
56:
2026:"L1 retrotransposition in neurons is modulated by MeCP2"
1860:"L1 retrotransposition in human neural progenitor cells"
1808:
2640:
Protasova MS, Andreeva TV, Rogaev EI (September 2021).
2639:
1564:
608:
Higher L1 copy numbers have been observed in the human
1757:
1315:
2373:
1698:
1600:
1006:"LINE-1 elements in structural variation and disease"
1004:
Beck CR, Garcia-Perez JL, Badge RM, Moran JV (2011).
834:"Initial sequencing and analysis of the human genome"
2535:
1227:
91:
during meiosis due to its repetitive DNA sequences.
63:, including animals and plants, classified with the
1413:
1364:
947:"Tracking LINE1 retrotransposition in the germline"
572:L1 activity has been observed in numerous types of
158:. Younger families also have two binding sites for
2229:
1974:
1316:Tchénio T, Casella JF, Heidmann T (January 2000).
3386:
2596:
1462:
782:
1975:Erwin JA, Marchetto MC, Gage FH (August 2014).
1917:"Mosaic copy number variation in human neurons"
1365:Athanikar JN, Badge RM, Moran JV (2004-01-01).
1129:
129:(bp) long and consists of two non-overlapping
2899:
2839:
2759:
895:
617:. This phenomenon is negatively regulated by
603:
2426:
2367:
2318:
2259:
1010:Annual Review of Genomics and Human Genetics
177:LINE-1 (L1.2) retrotransposable element ORF1
125:A typical L1 element is approximately 6,000
113:bacteria, evidently having arrived there by
2906:
2892:
2597:Tan K, Song HW, Wilkinson MF (July 2021).
1760:Annals of the New York Academy of Sciences
1520:Journal of Biomedicine & Biotechnology
1469:Journal of Biomedicine & Biotechnology
2865:
2822:
2787:
2777:
2667:
2657:
2622:
2579:
2569:
2507:
2458:
2409:
2399:
2350:
2301:
2291:
2206:
2149:
2139:
2098:
2057:
2000:
1948:
1891:
1834:
1734:
1724:
1675:
1626:
1541:
1531:
1490:
1480:
1439:
1390:
1341:
1253:
1204:
1155:
1080:
1070:
1029:
980:
970:
921:
867:
857:
808:
759:
621:, which is downregulated in NPCs, and by
343:domain and a highly variable coiled-coil
1656:Journal of the National Cancer Institute
825:
785:"The Evolution of LINE-1 in Vertebrates"
783:Boissinot S, Sookdeo A (December 2016).
531:structures likely present on the active
27:
3387:
2760:Wehbi SS, Zu Dohna H (November 2021).
1513:
527:activities of L1 ORF2p are boosted by
2887:
1970:
1968:
1108:. National Geographic. Archived from
358:LINE-1 retrotransposable element ORF2
1103:
944:
891:
889:
887:
729:
727:
591:Quantification of L1 copy number by
1463:Mätlik K, Redik K, Speek M (2006).
1022:10.1146/annurev-genom-082509-141802
902:The New England Journal of Medicine
896:Kazazian HH, Moran JV (July 2017).
562:
335:of L1 encodes a 500-amino acid, 40-
41:Long interspersed nuclear element-1
13:
2975:Short tandem repeat/Microsatellite
2528:
1965:
898:"Mobile DNA in Health and Disease"
656:
65:long interspersed nuclear elements
14:
3426:
2542:gene cluster suppresses germline
884:
724:
669:, a neurological disorder of the
555:, ORF2 is massively expressed in
504:of L1 encodes a protein that has
51:) is a family of related class I
1130:Anderson MT, Seifert HS (2011).
2475:
2223:
2166:
2115:
2074:
2017:
1908:
1851:
1802:
1751:
1692:
1643:
1594:
1558:
1507:
1456:
1407:
1358:
1309:
1270:
1221:
1172:
2979:Trinucleotide repeat disorders
1123:
1097:
1046:
997:
938:
776:
595:or L1 methylation levels with
166:promoter of unknown function.
16:Group of transposable elements
1:
2966:Variable number tandem repeat
1827:10.1158/1078-0432.CCR-09-2983
1580:10.1016/j.plasmid.2021.102560
717:
600:methods for early detection.
538:
141:(IRES) upstream of each ORF.
3395:Genes on human chromosome 22
2615:10.1016/j.celrep.2021.109423
2343:10.1016/j.celrep.2021.109530
1981:Nature Reviews. Neuroscience
1726:10.1371/journal.pone.0018884
1072:10.1371/journal.pgen.1002371
789:Genome Biology and Evolution
740:Genome Biology and Evolution
139:internal ribosome entry site
133:(ORFs) which are flanked by
120:
7:
3400:Genes on human chromosome 1
704:
683:
10:
3431:
2858:10.1038/s41422-020-00466-6
2779:10.1186/s13100-021-00255-x
2141:10.1186/s12915-022-01303-5
604:Neuropsychiatric disorders
542:
18:
3334:
3287:
3155:
3123:
3100:
3077:
3068:
3059:
3034:
2994:
2951:
2942:
2933:
635:autism spectrum disorders
567:
484:
474:
469:
465:
453:
436:
431:
419:
407:
395:
383:
375:
367:
362:
357:
315:
305:
300:
296:
284:
267:
262:
250:
238:
226:
214:
202:
194:
186:
181:
176:
144:
3415:Repetitive DNA sequences
2811:Human Molecular Genetics
2233:Human Molecular Genetics
1815:Clinical Cancer Research
1283:Human Molecular Genetics
661:Increased RNA levels of
115:horizontal gene transfer
3405:Mobile genetic elements
2571:10.1073/pnas.2024785118
2401:10.1073/pnas.2109066118
2293:10.1073/pnas.2105968118
1941:10.1126/science.1243472
1780:10.1196/annals.1448.011
1619:10.1126/science.1251343
1185:Genes & Development
972:10.1073/pnas.1709067114
350:
169:
21:Line 1 (disambiguation)
3374:Protein tandem repeats
3302:Tandemly arrayed genes
2700:10.1002/biot.202000620
2488:Nucleic Acids Research
1533:10.1155/jbb/2006/45621
1482:10.1155/JBB/2006/71753
1420:Nucleic Acids Research
1371:Nucleic Acids Research
1322:Nucleic Acids Research
1234:Nucleic Acids Research
588:cancer, respectively.
73:copy number variations
33:
2688:Biotechnology Journal
2659:10.3390/genes12101562
1295:10.1093/hmg/2.10.1697
1148:10.1128/mBio.00005-11
1104:Yong E (2011-02-16).
945:Wang PJ (July 2017).
914:10.1056/NEJMra1510092
552:Entamoeba histolytica
525:reverse transcriptase
510:reverse transcriptase
89:unequal crossing over
81:histone modifications
53:transposable elements
31:
3347:Pathogenicity island
2451:10.1128/JVI.00294-21
1334:10.1093/nar/28.2.411
667:macular degeneration
597:bisulfite sequencing
135:untranslated regions
39:(an abbreviation of
2562:2021PNAS..11824785T
2556:(23): e2024785118.
2439:Journal of Virology
2392:2021PNAS..11809066P
2386:(33): e2109066118.
2284:2021PNAS..11805968Z
2278:(21): e2105968118.
2199:10.1038/nature09830
2191:2011Natur.471..325K
2050:10.1038/nature09544
2042:2010Natur.468..443M
1933:2013Sci...342..632M
1884:10.1038/nature08248
1876:2009Natur.460.1127C
1772:2008NYASA1137..171S
1717:2011PLoSO...618884S
1668:10.1093/jnci/djn359
1197:10.1101/gad.1380406
1112:on December 5, 2019
963:2017PNAS..114.7194W
850:2001Natur.409..860L
529:single-stranded DNA
131:open reading frames
3297:Gene amplification
2824:10.1093/hmg/ddm105
2500:10.1093/nar/gkw925
2245:10.1093/hmg/ddl138
1514:Martin SL (2006).
1432:10.1093/nar/gku091
1383:10.1093/nar/gkh698
1246:10.1093/nar/gkj490
801:10.1093/gbe/evw247
752:10.1093/gbe/evw243
102:retrotransposons.
59:of many groups of
34:
3410:Molecular biology
3382:
3381:
3283:
3282:
3151:
3150:
3055:
3054:
2944:Repeated sequence
2919:repeated sequence
2817:(13): 1569–1577.
2731:(10): 1502–1510.
2100:10.2217/epi.15.68
1870:(7259): 1127–31.
1613:(6196): 1251343.
957:(28): 7194–7196.
844:(6822): 860–921.
795:(12): 3485–3507.
746:(11): 3301–3322.
647:Tourette syndrome
533:replication forks
498:
497:
494:
493:
329:
328:
325:
324:
151:tandemly repeated
3422:
3359:Low copy repeats
3352:Symbiosis island
3289:Gene duplication
3075:
3074:
3066:
3065:
2949:
2948:
2927:gene duplication
2908:
2901:
2894:
2885:
2884:
2879:
2869:
2836:
2826:
2801:
2791:
2781:
2756:
2719:
2681:
2671:
2661:
2636:
2626:
2593:
2583:
2573:
2522:
2521:
2511:
2479:
2473:
2472:
2462:
2445:(15): e0029421.
2430:
2424:
2423:
2413:
2403:
2371:
2365:
2364:
2354:
2322:
2316:
2315:
2305:
2295:
2263:
2257:
2256:
2227:
2221:
2220:
2210:
2185:(7338): 325–30.
2170:
2164:
2163:
2153:
2143:
2119:
2113:
2112:
2102:
2093:(8): 1275–1285.
2078:
2072:
2071:
2061:
2021:
2015:
2014:
2004:
1972:
1963:
1962:
1952:
1912:
1906:
1905:
1895:
1855:
1849:
1848:
1838:
1806:
1800:
1799:
1755:
1749:
1748:
1738:
1728:
1696:
1690:
1689:
1679:
1647:
1641:
1640:
1630:
1598:
1592:
1591:
1562:
1556:
1555:
1545:
1535:
1511:
1505:
1504:
1494:
1484:
1460:
1454:
1453:
1443:
1411:
1405:
1404:
1394:
1362:
1356:
1355:
1345:
1313:
1307:
1306:
1289:(10): 1697–702.
1274:
1268:
1267:
1257:
1225:
1219:
1218:
1208:
1176:
1170:
1169:
1159:
1142:(1): e00005-11.
1127:
1121:
1120:
1118:
1117:
1101:
1095:
1094:
1084:
1074:
1065:(11): e1002371.
1050:
1044:
1043:
1033:
1001:
995:
994:
984:
974:
942:
936:
935:
925:
893:
882:
881:
871:
861:
859:10.1038/35057062
829:
823:
822:
812:
780:
774:
773:
763:
731:
643:bipolar disorder
563:Roles in disease
467:
466:
355:
354:
298:
297:
174:
173:
43:, also known as
3430:
3429:
3425:
3424:
3423:
3421:
3420:
3419:
3385:
3384:
3383:
3378:
3330:
3279:
3147:
3119:
3096:
3070:Retrotransposon
3051:
3042:Inverted repeat
3030:
3015:DNA transposon
3011:Retrotransposon
3006:Gene conversion
2997:
2990:
2987:
2938:
2929:
2912:
2882:
2737:10.1038/ng.3945
2725:Nature Genetics
2694:(7): e2000620.
2531:
2529:Further reading
2526:
2525:
2494:(D1): D68–D73.
2480:
2476:
2431:
2427:
2372:
2368:
2323:
2319:
2264:
2260:
2239:(13): 2146–56.
2228:
2224:
2171:
2167:
2120:
2116:
2079:
2075:
2036:(7322): 443–6.
2022:
2018:
1993:10.1038/nrn3730
1973:
1966:
1927:(6158): 632–7.
1913:
1909:
1856:
1852:
1807:
1803:
1756:
1752:
1697:
1693:
1648:
1644:
1599:
1595:
1563:
1559:
1512:
1508:
1461:
1457:
1412:
1408:
1377:(13): 3846–55.
1363:
1359:
1314:
1310:
1275:
1271:
1226:
1222:
1177:
1173:
1128:
1124:
1115:
1113:
1102:
1098:
1051:
1047:
1002:
998:
943:
939:
894:
885:
830:
826:
781:
777:
732:
725:
720:
707:
686:
659:
657:Retinal disease
606:
570:
565:
547:
541:
353:
172:
147:
123:
77:DNA methylation
24:
17:
12:
11:
5:
3428:
3418:
3417:
3412:
3407:
3402:
3397:
3380:
3379:
3377:
3376:
3371:
3366:
3361:
3356:
3355:
3354:
3349:
3342:Genomic island
3338:
3336:
3332:
3331:
3329:
3328:
3323:
3322:
3321:
3311:
3310:
3309:
3299:
3293:
3291:
3285:
3284:
3281:
3280:
3278:
3277:
3272:
3267:
3262:
3257:
3252:
3247:
3242:
3237:
3232:
3227:
3222:
3217:
3212:
3207:
3202:
3197:
3192:
3187:
3182:
3177:
3172:
3167:
3161:
3159:
3157:DNA transposon
3153:
3152:
3149:
3148:
3146:
3145:
3140:
3135:
3129:
3127:
3121:
3120:
3118:
3117:
3112:
3106:
3104:
3098:
3097:
3095:
3094:
3089:
3083:
3081:
3072:
3063:
3057:
3056:
3053:
3052:
3050:
3049:
3044:
3038:
3036:
3032:
3031:
3029:
3028:
3027:
3026:
3021:
3013:
3008:
3002:
3000:
2992:
2991:
2989:
2988:
2985:Macrosatellite
2982:
2972:
2963:
2957:
2955:
2953:Tandem repeats
2946:
2940:
2939:
2934:
2931:
2930:
2911:
2910:
2903:
2896:
2888:
2881:
2880:
2852:(6): 613–630.
2837:
2802:
2757:
2720:
2682:
2637:
2594:
2546:transposition"
2532:
2530:
2527:
2524:
2523:
2474:
2425:
2366:
2317:
2258:
2222:
2165:
2114:
2073:
2016:
1987:(8): 497–506.
1964:
1907:
1850:
1801:
1750:
1691:
1662:(23): 1734–8.
1642:
1593:
1557:
1506:
1455:
1426:(7): 4546–62.
1406:
1357:
1308:
1269:
1220:
1171:
1122:
1096:
1045:
1016:(1): 187–215.
996:
937:
908:(4): 361–370.
883:
824:
775:
722:
721:
719:
716:
715:
714:
706:
703:
690:endogenisation
685:
682:
658:
655:
605:
602:
569:
566:
564:
561:
540:
537:
496:
495:
492:
491:
486:
482:
481:
476:
472:
471:
463:
462:
457:
451:
450:
440:
434:
433:
429:
428:
423:
417:
416:
411:
405:
404:
399:
393:
392:
387:
381:
380:
377:
373:
372:
369:
365:
364:
360:
359:
352:
349:
327:
326:
323:
322:
317:
313:
312:
307:
303:
302:
294:
293:
288:
282:
281:
271:
265:
264:
260:
259:
254:
248:
247:
242:
236:
235:
230:
224:
223:
218:
212:
211:
206:
200:
199:
196:
192:
191:
188:
184:
183:
179:
178:
171:
168:
146:
143:
122:
119:
15:
9:
6:
4:
3:
2:
3427:
3416:
3413:
3411:
3408:
3406:
3403:
3401:
3398:
3396:
3393:
3392:
3390:
3375:
3372:
3370:
3367:
3365:
3362:
3360:
3357:
3353:
3350:
3348:
3345:
3344:
3343:
3340:
3339:
3337:
3333:
3327:
3324:
3320:
3317:
3316:
3315:
3312:
3308:
3307:Ribosomal DNA
3305:
3304:
3303:
3300:
3298:
3295:
3294:
3292:
3290:
3286:
3276:
3273:
3271:
3268:
3266:
3263:
3261:
3258:
3256:
3253:
3251:
3248:
3246:
3243:
3241:
3238:
3236:
3233:
3231:
3228:
3226:
3223:
3221:
3218:
3216:
3213:
3211:
3208:
3206:
3203:
3201:
3198:
3196:
3193:
3191:
3188:
3186:
3183:
3181:
3178:
3176:
3173:
3171:
3168:
3166:
3163:
3162:
3160:
3158:
3154:
3144:
3141:
3139:
3136:
3134:
3131:
3130:
3128:
3126:
3122:
3116:
3113:
3111:
3108:
3107:
3105:
3103:
3099:
3093:
3090:
3088:
3085:
3084:
3082:
3080:
3076:
3073:
3071:
3067:
3064:
3062:
3058:
3048:
3047:Direct repeat
3045:
3043:
3040:
3039:
3037:
3033:
3025:
3022:
3020:
3017:
3016:
3014:
3012:
3009:
3007:
3004:
3003:
3001:
2999:
2993:
2986:
2983:
2980:
2976:
2973:
2971:
2970:Minisatellite
2967:
2964:
2962:
2961:Satellite DNA
2959:
2958:
2956:
2954:
2950:
2947:
2945:
2941:
2937:
2932:
2928:
2924:
2920:
2916:
2909:
2904:
2902:
2897:
2895:
2890:
2889:
2886:
2877:
2873:
2868:
2863:
2859:
2855:
2851:
2847:
2846:Cell Research
2843:
2838:
2834:
2830:
2825:
2820:
2816:
2812:
2808:
2803:
2799:
2795:
2790:
2785:
2780:
2775:
2771:
2767:
2763:
2758:
2754:
2750:
2746:
2742:
2738:
2734:
2730:
2726:
2721:
2717:
2713:
2709:
2705:
2701:
2697:
2693:
2689:
2683:
2679:
2675:
2670:
2665:
2660:
2655:
2651:
2647:
2643:
2638:
2634:
2630:
2625:
2620:
2616:
2612:
2609:(3): 109423.
2608:
2604:
2600:
2595:
2591:
2587:
2582:
2577:
2572:
2567:
2563:
2559:
2555:
2551:
2547:
2545:
2541:
2534:
2533:
2519:
2515:
2510:
2505:
2501:
2497:
2493:
2489:
2485:
2478:
2470:
2466:
2461:
2456:
2452:
2448:
2444:
2440:
2436:
2429:
2421:
2417:
2412:
2407:
2402:
2397:
2393:
2389:
2385:
2381:
2377:
2370:
2362:
2358:
2353:
2348:
2344:
2340:
2337:(7): 109530.
2336:
2332:
2328:
2321:
2313:
2309:
2304:
2299:
2294:
2289:
2285:
2281:
2277:
2273:
2269:
2262:
2254:
2250:
2246:
2242:
2238:
2234:
2226:
2218:
2214:
2209:
2204:
2200:
2196:
2192:
2188:
2184:
2180:
2176:
2169:
2161:
2157:
2152:
2147:
2142:
2137:
2133:
2129:
2125:
2118:
2110:
2106:
2101:
2096:
2092:
2088:
2084:
2077:
2069:
2065:
2060:
2055:
2051:
2047:
2043:
2039:
2035:
2031:
2027:
2020:
2012:
2008:
2003:
1998:
1994:
1990:
1986:
1982:
1978:
1971:
1969:
1960:
1956:
1951:
1946:
1942:
1938:
1934:
1930:
1926:
1922:
1918:
1911:
1903:
1899:
1894:
1889:
1885:
1881:
1877:
1873:
1869:
1865:
1861:
1854:
1846:
1842:
1837:
1832:
1828:
1824:
1821:(5): 1682–9.
1820:
1816:
1812:
1805:
1797:
1793:
1789:
1785:
1781:
1777:
1773:
1769:
1765:
1761:
1754:
1746:
1742:
1737:
1732:
1727:
1722:
1718:
1714:
1711:(4): e18884.
1710:
1706:
1702:
1695:
1687:
1683:
1678:
1673:
1669:
1665:
1661:
1657:
1653:
1646:
1638:
1634:
1629:
1624:
1620:
1616:
1612:
1608:
1604:
1597:
1589:
1585:
1581:
1577:
1573:
1569:
1561:
1553:
1549:
1544:
1539:
1534:
1529:
1525:
1521:
1517:
1510:
1502:
1498:
1493:
1488:
1483:
1478:
1474:
1470:
1466:
1459:
1451:
1447:
1442:
1437:
1433:
1429:
1425:
1421:
1417:
1410:
1402:
1398:
1393:
1388:
1384:
1380:
1376:
1372:
1368:
1361:
1353:
1349:
1344:
1339:
1335:
1331:
1327:
1323:
1319:
1312:
1304:
1300:
1296:
1292:
1288:
1284:
1280:
1273:
1265:
1261:
1256:
1251:
1247:
1243:
1240:(3): 853–64.
1239:
1235:
1231:
1224:
1216:
1212:
1207:
1202:
1198:
1194:
1191:(2): 210–24.
1190:
1186:
1182:
1175:
1167:
1163:
1158:
1153:
1149:
1145:
1141:
1137:
1133:
1126:
1111:
1107:
1100:
1092:
1088:
1083:
1078:
1073:
1068:
1064:
1060:
1059:PLOS Genetics
1056:
1049:
1041:
1037:
1032:
1027:
1023:
1019:
1015:
1011:
1007:
1000:
992:
988:
983:
978:
973:
968:
964:
960:
956:
952:
948:
941:
933:
929:
924:
919:
915:
911:
907:
903:
899:
892:
890:
888:
879:
875:
870:
869:2027.42/62798
865:
860:
855:
851:
847:
843:
839:
835:
828:
820:
816:
811:
806:
802:
798:
794:
790:
786:
779:
771:
767:
762:
757:
753:
749:
745:
741:
737:
730:
728:
723:
712:
709:
708:
702:
699:
695:
691:
681:
679:
674:
672:
668:
664:
654:
652:
648:
644:
640:
636:
632:
631:schizophrenia
628:
627:Rett syndrome
624:
620:
616:
611:
601:
598:
594:
589:
587:
584:in colon and
583:
579:
575:
560:
558:
554:
553:
546:
536:
534:
530:
526:
522:
517:
515:
511:
507:
503:
490:
487:
483:
480:
477:
473:
468:
464:
461:
458:
456:
452:
449:
448:
444:
441:
439:
435:
430:
427:
424:
422:
418:
415:
412:
410:
406:
403:
400:
398:
394:
391:
388:
386:
382:
378:
374:
370:
366:
361:
356:
348:
346:
342:
338:
334:
321:
318:
314:
311:
308:
304:
299:
295:
292:
289:
287:
283:
280:
279:
275:
272:
270:
266:
261:
258:
255:
253:
249:
246:
243:
241:
237:
234:
231:
229:
225:
222:
219:
217:
213:
210:
207:
205:
201:
197:
193:
189:
185:
180:
175:
167:
165:
161:
157:
152:
142:
140:
136:
132:
128:
118:
116:
112:
107:
105:
101:
97:
92:
90:
86:
82:
78:
74:
70:
66:
62:
58:
54:
50:
46:
42:
38:
30:
26:
22:
3319:Gene cluster
3109:
3087:Alu sequence
2996:Interspersed
2849:
2845:
2814:
2810:
2769:
2765:
2728:
2724:
2691:
2687:
2652:(10): 1562.
2649:
2645:
2606:
2603:Cell Reports
2602:
2553:
2549:
2543:
2539:
2491:
2487:
2477:
2442:
2438:
2428:
2383:
2379:
2369:
2334:
2331:Cell Reports
2330:
2320:
2275:
2271:
2261:
2236:
2232:
2225:
2182:
2178:
2168:
2131:
2127:
2117:
2090:
2086:
2076:
2033:
2029:
2019:
1984:
1980:
1924:
1920:
1910:
1867:
1863:
1853:
1818:
1814:
1804:
1766:(1): 171–4.
1763:
1759:
1753:
1708:
1704:
1694:
1659:
1655:
1645:
1610:
1606:
1596:
1571:
1567:
1560:
1526:(1): 45621.
1523:
1519:
1509:
1475:(1): 71753.
1472:
1468:
1458:
1423:
1419:
1409:
1374:
1370:
1360:
1328:(2): 411–5.
1325:
1321:
1311:
1286:
1282:
1272:
1237:
1233:
1223:
1188:
1184:
1174:
1139:
1135:
1125:
1114:. Retrieved
1110:the original
1099:
1062:
1058:
1048:
1013:
1009:
999:
954:
950:
940:
905:
901:
841:
837:
827:
792:
788:
778:
743:
739:
687:
675:
660:
607:
590:
571:
550:
548:
518:
506:endonuclease
499:
445:
376:Alt. symbols
330:
276:
195:Alt. symbols
148:
124:
108:
93:
69:human genome
48:
44:
40:
36:
35:
25:
3314:Gene family
3225:Tc1/mariner
3180:EnSpm/CACTA
2128:BMC Biology
2087:Epigenomics
649:, and drug
586:endometrial
500:The second
479:Swiss-model
363:Identifiers
310:Swiss-model
182:Identifiers
3389:Categories
3326:Pseudogene
3143:retroposon
3061:Transposon
2923:transposon
2766:Mobile DNA
2134:(1): 116.
1574:: 102560.
1116:2016-07-14
718:References
696:genome in
694:SARS-CoV-2
543:See also:
539:Regulation
475:Structures
470:Search for
432:Other data
345:N-terminus
341:C-terminus
331:The first
306:Structures
301:Search for
263:Other data
127:base pairs
61:eukaryotes
3245:P element
3195:Harbinger
2936:Repeatome
2772:(1): 26.
2716:233484152
651:addiction
615:mosaicism
557:antisense
460:Q18028649
385:NCBI gene
291:Q18028646
204:NCBI gene
164:antisense
121:Structure
111:gonorrhea
104:Mutations
3369:Telomere
3335:See also
3275:Zisupton
3255:Polinton
3250:PiggyBac
3205:Helitron
3024:Helitron
3019:Polinton
2915:Genetics
2876:33514913
2833:17468180
2798:34782009
2745:28846101
2708:33938150
2678:34680956
2633:34289349
2590:34083437
2518:27924012
2469:33980601
2420:34344759
2361:34380018
2312:33958444
2253:16723373
2217:21297615
2160:35581640
2109:26212695
2068:21085180
2011:25005482
1959:24179226
1902:19657334
1845:20179218
1796:32676787
1788:18837943
1745:21533144
1705:PLOS ONE
1686:19033568
1637:25082706
1588:33482228
1552:16877816
1501:16877819
1450:24493738
1401:15272086
1352:10606637
1264:16464823
1215:16418485
1166:21325040
1091:22125493
1040:21801021
991:28663337
932:28745987
878:11237011
819:28175298
770:27702814
705:See also
684:COVID-19
639:epilepsy
489:InterPro
455:Wikidata
320:InterPro
286:Wikidata
98:and SVA
3265:Transib
3240:Novosib
3220:Kolobok
3190:Ginger2
3185:Ginger1
3170:Crypton
2867:8169921
2789:8594186
2753:5213902
2669:8535693
2624:8357189
2581:8201764
2558:Bibcode
2509:5210629
2460:8274596
2411:8379926
2388:Bibcode
2352:8316065
2303:8166107
2280:Bibcode
2208:3077055
2187:Bibcode
2151:9115989
2059:3059197
2038:Bibcode
2002:4443810
1950:3975283
1929:Bibcode
1921:Science
1893:2909034
1872:Bibcode
1836:2831156
1768:Bibcode
1736:3077413
1713:Bibcode
1677:2639290
1628:4380235
1607:Science
1568:Plasmid
1543:1510943
1492:1559930
1441:3985663
1303:8268924
1255:1361618
1206:1356112
1157:3042738
1082:3219598
1031:4124830
982:5514774
959:Bibcode
923:5980640
846:Bibcode
810:5381506
761:5203782
692:of the
574:cancers
521:nicking
485:Domains
421:UniProt
379:L1ORF2p
316:Domains
274:Chr. 22
252:UniProt
198:L1ORF1p
55:in the
3364:CRISPR
3230:Merlin
3215:ISL2EU
3165:Academ
2998:repeat
2874:
2864:
2831:
2796:
2786:
2751:
2743:
2714:
2706:
2676:
2666:
2631:
2621:
2588:
2578:
2516:
2506:
2467:
2457:
2418:
2408:
2359:
2349:
2310:
2300:
2251:
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2205:
2179:Nature
2158:
2148:
2107:
2066:
2056:
2030:Nature
2009:
1999:
1957:
1947:
1900:
1890:
1864:Nature
1843:
1833:
1794:
1786:
1743:
1733:
1684:
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1625:
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1550:
1540:
1499:
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1448:
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1392:506791
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1350:
1343:102531
1340:
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1213:
1203:
1164:
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838:Nature
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711:L1Base
680:gene.
568:Cancer
443:Chr. 1
426:O00370
368:Symbol
257:Q9UN81
233:151626
187:Symbol
145:5' UTR
83:, and
49:LINE-1
3270:Zator
3210:IS3EU
3115:LINE2
3110:LINE1
3102:LINEs
3079:SINEs
3035:Other
2749:S2CID
2712:S2CID
2646:Genes
2544:LINE1
2538:"The
1792:S2CID
678:Nr2e3
623:MeCP2
610:brain
438:Locus
371:L1RE2
278:q12.1
269:Locus
190:L1RE1
85:piRNA
37:LINE1
3260:Sola
3235:MuDR
3175:Dada
3138:MER4
3133:HERV
3125:LTRs
2872:PMID
2829:PMID
2794:PMID
2741:PMID
2704:PMID
2674:PMID
2629:PMID
2586:PMID
2540:Rhox
2514:PMID
2465:PMID
2416:PMID
2357:PMID
2308:PMID
2249:PMID
2213:PMID
2156:PMID
2105:PMID
2064:PMID
2007:PMID
1955:PMID
1898:PMID
1841:PMID
1784:PMID
1764:1137
1741:PMID
1682:PMID
1633:PMID
1584:PMID
1548:PMID
1524:2006
1497:PMID
1473:2006
1446:PMID
1397:PMID
1348:PMID
1299:PMID
1260:PMID
1211:PMID
1162:PMID
1136:mBio
1087:PMID
1036:PMID
987:PMID
928:PMID
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815:PMID
766:PMID
698:Huh7
671:eyes
619:Sox2
593:qPCR
582:PTEN
580:and
523:and
519:The
508:and
414:1VYB
402:6687
397:HGNC
390:4030
351:ORF2
245:2LDY
228:OMIM
221:6686
216:HGNC
209:4029
170:ORF1
100:SINE
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3092:MIR
2862:PMC
2854:doi
2819:doi
2784:PMC
2774:doi
2733:doi
2696:doi
2664:PMC
2654:doi
2619:PMC
2611:doi
2576:PMC
2566:doi
2554:118
2504:PMC
2496:doi
2455:PMC
2447:doi
2406:PMC
2396:doi
2384:118
2347:PMC
2339:doi
2298:PMC
2288:doi
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2241:doi
2203:PMC
2195:doi
2183:471
2146:PMC
2136:doi
2095:doi
2054:PMC
2046:doi
2034:468
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