107:, most of which were located in coding regions of the UCEs. A study performed in mice determined that deleting UCEs from the genome did not create obvious deleterious phenotypes, despite deletion of UCEs in proximity to promoters and protein coding genes. Affected mice were fertile and targeted screens of the nearby coding genes showed no altered phenotype. A separate mouse study demonstrated that ultraconserved enhancers were robust to mutagenesis, concluding that perfect conservation of UCE sequences is not required for their function, which would suggest another reason for the sequence consistency besides evolutionary importance. Computational analysis of human
1581:
1523:
99:) has been estimated at less than 10 in 2.9 billion bases. In support of the cold spot hypothesis, UCEs were found to be mutating 20 fold less than expected under conservative models for neutral mutation rates. This fold change difference in mutation rates was consistent between humans, chimpanzees, and chickens. Ultraconserved elements are not exempt from mutations, as exemplified by the presence of 29,983 polymorphisms in the UCE regions of the human genome assembly
70:. Out of 481 identified human UCEs, approximately 97% align with high identity to the chicken genome, though only 4% of the human genome can be reliably aligned to the chicken genome. Similarly, the same sequences in the fugu genome have 68% identity to human UCEs, despite the human genome only reliably aligning to 1.8% of the fugu genome. Despite often being
94:
for 300-400 million years. More recently, this assumption has been replaced by two main hypotheses: that UCEs are created through a reduced negative selection rate, or through reduced mutation rates, also known as a "cold spot" of evolution. Many studies have examined the validity of each hypothesis.
153:
in the brain, though the effect was not lethal. Some UCEs are not transcribed, and are referred to as ultraconserved noncoding elements. However, many UCRs in humans are extensively transcribed. A small number of those which are transcribed, known as transcribed UCEs (T-UCEs), have been connected
178:
Research has demonstrated that T-UCRs have a tissue-specific expression, and a differential expression profile between tumors and other diseases. The tables below highlight transcripts and polymorphisms within UCRs that have been shown to contribute to human diseases. For example, UCRs tend to
3011:
Fodde R, van der Luijt R, Wijnen J, Tops C, van der Klift H, van
Leeuwen-Cornelisse I, et al. (August 1992). "Eight novel inactivating germ line mutations at the APC gene identified by denaturing gradient gel electrophoresis".
39:
and shows little or no variation between those taxa. These regions and regions adjacent to them (flanking DNA) are useful for tracing the evolutionary history of groups of organisms. Another term for ultraconserved element is
4324:
Saugier-Veber P, Munnich A, Bonneau D, Rozet JM, Le Merrer M, Gil R, Boespflug-Tanguy O (March 1994). "X-linked spastic paraplegia and
Pelizaeus-Merzbacher disease are allelic disorders at the proteolipid protein locus".
3142:"GeneMatcher aids in the identification of a new malformation syndrome with intellectual disability, unique facial dysmorphisms, and skeletal and connective tissue abnormalities caused by de novo variants in HNRNPK"
3047:
Curia MC, Esposito DL, Aceto G, Palmirotta R, Crognale S, Valanzano R, et al. (1998). "Transcript dosage effect in familial adenomatous polyposis: model offered by two kindreds with exon 9 APC gene mutations".
3784:
Takahashi Y, Kou I, Takahashi A, Johnson TA, Kono K, Kawakami N, et al. (October 2011). "A genome-wide association study identifies common variants near LBX1 associated with adolescent idiopathic scoliosis".
3544:
Grauers A, Wang J, Einarsdottir E, Simony A, Danielsson A, Γ
kesson K, et al. (October 2015). "Candidate gene analysis and exome sequencing confirm LBX1 as a susceptibility gene for idiopathic scoliosis".
1061:
Pereira
Zambalde E, Mathias C, Rodrigues AC, de Souza Fonseca Ribeiro EM, Fiori Gradia D, Calin GA, Carvalho de Oliveira J (March 2020). "Highlighting transcribed ultraconserved regions in human diseases".
845:
Faircloth, BC; McCormack, JE; Crawford, NG; Harvey, MG; Brumfield, RT; Glenn, TC (October 2012). "Ultraconserved elements anchor thousands of genetic markers spanning multiple evolutionary timescales".
3639:
Londono D, Kou I, Johnson TA, Sharma S, Ogura Y, Tsunoda T, et al. (June 2014). "A meta-analysis identifies adolescent idiopathic scoliosis association with LBX1 locus in multiple ethnic groups".
62:
Since its creation, this term's usage has broadened to include more evolutionarily distant species or shorter segments, for example 100 bp instead of 200 bp. By some definitions, segments need not be
47:
The term "ultraconserved element" was originally defined as a genome segment longer than 200 base pairs (bp) that is absolutely conserved, with no insertions or deletions and 100% identity, between
3741:
Jiang Y, Ben Q, Shen H, Lu W, Zhang Y, Zhu J (November 2011). "Diabetes mellitus and incidence and mortality of colorectal cancer: a systematic review and meta-analysis of cohort studies".
170:
cells. Indeed, UCEs are often affected by copy number variation in cancer cells much more than in healthy contexts, suggesting that altering the copy number of T-UCEs may be deleterious.
108:
4281:
Gorman MP, Golomb MR, Walsh LE, Hobson GM, Garbern JY, Kinkel RP, et al. (April 2007). "Steroid-responsive neurologic relapses in a child with a proteolipid protein-1 mutation".
649:
Chettier et al. (2015); Gao et al. (2013); Grauers et al. (2015); Jiang et al. (2013); Londono et al. (2014); Miyake et al. (2013); Shen et al. (2011); Takahashi et al. (2011)
881:
Zhang, Y. Miles; Williams, Jason L.; Lucky, Andrea (3 September 2019). "Understanding UCEs: A Comprehensive Primer on Using
Ultraconserved Elements for Arthropod Phylogenomics".
119:. This may indicate that there is some change to DNA structure in these regions favoring their precise retention, but this possibility has not been validated through testing.
4515:
Knoblauch H, Geier C, Adams S, Budde B, Rudolph A, Zacharias U, et al. (January 2010). "Contractures and hypertrophic cardiomyopathy in a novel FHL1 mutation".
1724:
Derti A, Roth FP, Church GM, Wu CT (October 2006). "Mammalian ultraconserved elements are strongly depleted among segmental duplications and copy number variants".
4561:
Schoser B, Goebel HH, Janisch I, Quasthoff S, Rother J, Bergmann M, et al. (August 2009). "Consequences of mutations within the C terminus of the FHL1 gene".
2429:"Deregulation of methylation of transcribed-ultra conserved regions in colorectal cancer and their value for detection of adenomas and adenocarcinomas"
1213:"Ultraconserved cDNA segments in the human transcriptome exhibit resistance to folding and implicate function in translation and alternative splicing"
1108:
Katzman S, Kern AD, Bejerano G, Fewell G, Fulton L, Wilson RK, et al. (August 2007). "Human genome ultraconserved elements are ultraselected".
3928:"Identifying gene-gene interactions that are highly associated with Body Mass Index using Quantitative Multifactor Dimensionality Reduction (QMDR)"
1874:"Transcribed ultraconserved region Uc.63+ promotes resistance to docetaxel through regulation of androgen receptor signaling in prostate cancer"
4028:"Genetic variants associated with motion sickness point to roles for inner ear development, neurological processes and glucose homeostasis"
2668:"Ultraconserved elements in the human genome: association and transmission analyses of highly constrained single-nucleotide polymorphisms"
111:(UCNEs) found that the regions are enriched for A-T sequences and are generally GC poor. However, the UNCEs were found to be enriched for
995:
Bejerano G, Pheasant M, Makunin I, Stephen S, Kent WJ, Mattick JS, Haussler D (May 2004). "Ultraconserved elements in the human genome".
4385:
3256:
3240:"Homonucleotide expansion and contraction mutations of PAX2 and inclusion of Chiari 1 malformation as part of renal-coloboma syndrome"
3239:
180:
4370:"Nonsense mutation in exon 3 of the proteolipid protein gene (PLP) in a family with an unusual form of Pelizaeus-Merzbacher disease"
3061:
127:
Often, ultraconserved elements are located near transcriptional regulators or developmental genes performing functions such as gene
4409:"Mutations in ionotropic AMPA receptor 3 alter channel properties and are associated with moderate cognitive impairment in humans"
17:
3830:"A peroxisomal disorder of severe intellectual disability, epilepsy, and cataracts due to fatty acyl-CoA reductase 1 deficiency"
1926:"The transcribed-ultraconserved regions in prostate and gastric cancer: DNA hypermethylation and microRNA-associated regulation"
3338:"Further delineation of renal-coloboma syndrome in patients with extreme variability of phenotype and identical PAX2 mutations"
91:
4609:"An X-linked myopathy with postural muscle atrophy and generalized hypertrophy, termed XMPMA, is caused by mutations in FHL1"
2221:"A transcribed ultraconserved noncoding RNA, Uc.173, is a key molecule for the inhibition of lead-induced neuronal apoptosis"
179:
accumulate less mutations than flanking segments, in both neoplastic and non-neoplastic samples from persons with hereditary
96:
3487:"Association between common variants near LBX1 and adolescent idiopathic scoliosis replicated in the Chinese Han population"
1976:"Pancreatic cancer-secreted miR-155 implicates in the conversion from normal fibroblasts to cancer-associated fibroblasts"
1616:"Expression and functional role of a transcribed noncoding RNA with an ultraconserved element in hepatocellular carcinoma"
2427:
Kottorou AE, Antonacopoulou AG, Dimitrakopoulos FD, Diamantopoulou G, Sirinian C, Kalofonou M, et al. (April 2018).
4658:"Mutation analysis of the MECP2 gene in patients of Slavic origin with Rett syndrome: novel mutations and polymorphisms"
1925:
55:. If ribosomal DNA (rDNA regions) are excluded, these range in size from 200 bp to 781 bp. UCEs are found on all human
66:
between species. Human UCEs also show high conservation with more evolutionarily distant species, such as chicken and
3979:"Gene-centric meta-analyses of 108 912 individuals confirm known body mass index loci and reveal three novel signals"
3592:"Association of rs11190870 near LBX1 with adolescent idiopathic scoliosis susceptibility in a Han Chinese population"
4126:"DGGE-based whole-gene mutation scanning of the dystrophin gene in Duchenne and Becker muscular dystrophy patients"
2810:"Genome-wide association analyses identify multiple loci associated with central corneal thickness and keratoconus"
2527:"New Cross-Talk Layer between Ultraconserved Non-Coding RNAs, MicroRNAs and Polycomb Protein YY1 in Bladder Cancer"
2913:"Familial adenomatous polyposis-associated thyroid cancer: a clinical, pathological, and molecular genetics study"
1426:"Nucleotide Composition of Ultra-Conserved Elements Shows Excess of GpC and Depletion of GG and CC Dinucleotides"
3428:"Haplotypes at LBX1 have distinct inheritance patterns with opposite effects in adolescent idiopathic scoliosis"
1816:
De Grassi A, Segala C, Iannelli F, Volorio S, Bertario L, Radice P, et al. (January 2010). Hastie N (ed.).
3189:
Mallery DL, Tanganelli B, Colella S, Steingrimsdottir H, van Gool AJ, Troelstra C, et al. (January 1998).
1818:"Ultradeep sequencing of a human ultraconserved region reveals somatic and constitutional genomic instability"
2123:"Long Noncoding RNA uc.173 Promotes Renewal of the Intestinal Mucosa by Inducing Degradation of MicroRNA 195"
3684:"Identification of a susceptibility locus for severe adolescent idiopathic scoliosis on chromosome 17q24.3"
2066:"Ultraconserved element uc.372 drives hepatic lipid accumulation by suppressing miR-195/miR4668 maturation"
146:
2323:"Transcribed ultraconserved region 339 promotes carcinogenesis by modulating tumor suppressor microRNAs"
3238:
Schimmenti LA, Shim HH, Wirtschafter JD, Panzarino VA, Kashtan CE, Kirkpatrick SJ, et al. (1999).
1675:"Abnormal dosage of ultraconserved elements is highly disfavored in healthy cells but not cancer cells"
1368:
Snetkova V, Ypsilanti AR, Akiyama JA, Mannion BJ, Plajzer-Frick I, Novak CS, et al. (April 2021).
4607:
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2862:
Bosch DG, Boonstra FN, Gonzaga-Jauregui C, Xu M, de Ligt J, Jhangiani S, et al. (February 2014).
2380:"Long non-coding RNA containing ultraconserved genomic region 8 promotes bladder cancer tumorigenesis"
2025:"Regulation of pri-miRNA processing by a long noncoding RNA transcribed from an ultraconserved region"
4124:
Hofstra RM, Mulder IM, Vossen R, de Koning-Gans PA, Kraak M, Ginjaar IB, et al. (January 2004).
826:
2321:
Vannini I, Wise PM, Challagundla KB, Plousiou M, Raffini M, Bandini E, et al. (November 2017).
1580:
1522:
1017:
3140:
Au PY, You J, Caluseriu O, Schwartzentruber J, Majewski J, Bernier FP, et al. (October 2015).
167:
4232:
Tuffery-Giraud S, Saquet C, Thorel D, Disset A, Rivier F, Malcolm S, Claustres M (December 2005).
1480:"Transcriptional profiling of lung cell populations in idiopathic pulmonary arterial hypertension"
135:
regulation. A study comparing ultraconserved elements between humans and the
Japanese puffer fish
4466:
Gueneau L, Bertrand AT, Jais JP, Salih MA, Stojkovic T, Wehnert M, et al. (September 2009).
3877:
O'Donnell PH, Stark AL, Gamazon ER, Wheeler HE, McIlwee BE, Gorsic L, et al. (August 2012).
2525:
Terreri S, Durso M, Colonna V, Romanelli A, Terracciano D, Ferro M, et al. (December 2016).
2619:"A massive parallel sequencing workflow for diagnostic genetic testing of mismatch repair genes"
86:
Researchers originally assumed that perfect conservation of these long stretches of DNA implied
51:
regions of the human, rat, and mouse genomes. 481 of these segments have been identified in the
4075:
Al Turki S, Manickaraj AK, Mercer CL, Gerety SS, Hitz MP, Lindsay S, et al. (April 2014).
2666:
Chiang CW, Liu CT, Lettre G, Lange LA, Jorgensen NW, Keating BJ, et al. (September 2012).
2172:"Non-codingRNA sequence variations in human chronic lymphocytic leukemia and colorectal cancer"
1012:
104:
186:
3336:
Schimmenti LA, Cunliffe HE, McNoe LA, Ward TA, French MC, Shim HH, et al. (April 1997).
2759:
Dobbs MB, Gurnett CA, Pierce B, Exner GU, Robarge J, Morcuende JA, et al. (March 2006).
2717:"Genome-wide association study identifies ZFHX1B as a susceptibility locus for severe myopia"
2576:
Cronin S, Berger S, Ding J, Schymick JC, Washecka N, Hernandez DG, et al. (March 2008).
2378:
Olivieri M, Ferro M, Terreri S, Durso M, Romanelli A, Avitabile C, et al. (April 2016).
2170:
Wojcik SE, Rossi S, Shimizu M, Nicoloso MS, Cimmino A, Alder H, et al. (February 2010).
1872:
Sekino Y, Sakamoto N, Goto K, Honma R, Shigematsu Y, Sentani K, et al. (November 2017).
1170:
Calin GA, Liu CG, Ferracin M, Hyslop T, Spizzo R, Sevignani C, et al. (September 2007).
4407:
Wu Y, Arai AC, Rumbaugh G, Srivastava AK, Turner G, Hayashi T, et al. (November 2007).
3287:
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3191:"Molecular analysis of mutations in the CSB (ERCC6) gene in patients with Cockayne syndrome"
2808:
Lu Y, Vitart V, Burdon KP, Khor CC, Bykhovskaya Y, Mirshahi A, et al. (February 2013).
2478:"Ultraconserved region-containing Transformer 2Ξ²4 controls senescence of colon cancer cells"
4420:
4186:
3828:
Buchert R, Tawamie H, Smith C, Uebe S, Innes AM, Al Hallak B, et al. (November 2014).
3695:
3498:
3439:
3377:
Barua M, Stellacci E, Stella L, Weins A, Genovese G, Muto V, et al. (September 2014).
2962:"Analysing the mutational status of adenomatous polyposis coli (APC) gene in breast cancer"
2334:
2077:
1627:
1614:
Braconi C, Valeri N, Kogure T, Gasparini P, Huang N, Nuovo GJ, et al. (January 2011).
1597:
1117:
1004:
1536:
Woolfe A, Goodson M, Goode DK, Snell P, McEwen GK, Vavouri T, et al. (January 2005).
8:
4656:
Zahorakova D, Rosipal R, Hadac J, Zumrova A, Bzduch V, Misovicova N, et al. (2007).
1924:
Goto K, Ishikawa S, Honma R, Tanimoto K, Sakamoto N, Sentani K, et al. (July 2016).
1478:
Saygin D, Tabib T, Bittar HE, Valenzi E, Sembrat J, Chan SY, et al. (January 2005).
128:
74:, some ultraconserved elements have been found to be transcriptionally active, producing
4424:
4190:
3699:
3502:
3443:
2338:
2081:
1631:
1121:
1060:
1008:
4687:
4633:
4608:
4586:
4540:
4492:
4467:
4443:
4408:
4350:
4306:
4294:
4263:
4155:
4101:
4076:
4052:
4027:
4003:
3978:
3954:
3927:
3903:
3878:
3854:
3829:
3810:
3766:
3718:
3683:
3664:
3616:
3591:
3521:
3486:
3462:
3427:
3403:
3378:
3354:
3337:
3318:
3269:
3215:
3190:
3166:
3141:
3117:
3092:
3091:
Johnson JO, Pioro EP, Boehringer A, Chia R, Feit H, Renton AE, et al. (May 2014).
3073:
2988:
2961:
2937:
2912:
2911:
Soravia C, Sugg SL, Berk T, Mitri A, Cheng H, Gallinger S, et al. (January 1999).
2888:
2863:
2834:
2809:
2790:
2692:
2667:
2643:
2618:
2553:
2526:
2502:
2477:
2453:
2428:
2404:
2379:
2355:
2322:
2298:
2269:
2245:
2220:
2196:
2171:
2147:
2122:
2098:
2065:
2000:
1975:
1956:
1898:
1873:
1844:
1817:
1793:
1768:
1749:
1701:
1674:
1650:
1615:
1452:
1425:
1394:
1369:
1345:
1318:
1291:
1266:
1237:
1212:
1141:
1087:
1038:
947:
920:
3289:"PAX2 mutations in renal-coloboma syndrome: mutational hotspot and germline mosaicism"
2928:
2476:
Kajita K, Kuwano Y, Satake Y, Kano S, Kurokawa K, Akaike Y, et al. (April 2016).
1564:
1537:
1506:
1479:
1172:"Ultraconserved regions encoding ncRNAs are altered in human leukemias and carcinomas"
4679:
4638:
4578:
4532:
4497:
4448:
4389:
4342:
4298:
4255:
4214:
4209:
4174:
4147:
4106:
4057:
4008:
3959:
3926:
De R, Verma SS, Drenos F, Holzinger ER, Holmes MV, Hall MA, et al. (June 2015).
3908:
3859:
3802:
3758:
3723:
3656:
3621:
3572:
3526:
3467:
3408:
3359:
3310:
3261:
3220:
3171:
3122:
3065:
3029:
3025:
2993:
2942:
2893:
2839:
2782:
2738:
2697:
2648:
2599:
2558:
2507:
2458:
2409:
2360:
2303:
2250:
2201:
2152:
2103:
2046:
2005:
1948:
1903:
1849:
1798:
1741:
1706:
1655:
1593:
1569:
1511:
1457:
1399:
1350:
1317:
Ahituv N, Zhu Y, Visel A, Holt A, Afzal V, Pennacchio LA, Rubin EM (September 2007).
1296:
1242:
1193:
1133:
1091:
1079:
1030:
952:
863:
137:
4590:
4544:
4354:
4310:
4267:
4159:
3682:
Miyake A, Kou I, Takahashi Y, Johnson TA, Ogura Y, Dai J, et al. (2013-09-04).
3668:
3322:
3273:
2794:
2715:
Khor CC, Miyake M, Chen LJ, Shi Y, Barathi VA, Qiao F, et al. (December 2013).
1753:
1145:
393:
4691:
4669:
4628:
4620:
4570:
4524:
4487:
4479:
4438:
4428:
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4245:
4204:
4194:
4137:
4096:
4088:
4047:
4039:
3998:
3990:
3949:
3939:
3898:
3890:
3879:"Identification of novel germline polymorphisms governing capecitabine sensitivity"
3849:
3841:
3814:
3794:
3750:
3713:
3703:
3648:
3611:
3603:
3562:
3554:
3516:
3506:
3457:
3447:
3426:
Chettier R, Nelson L, Ogilvie JW, Albertsen HM, Ward K (2015-02-12). Fang S (ed.).
3398:
3390:
3349:
3300:
3251:
3210:
3202:
3161:
3153:
3112:
3104:
3077:
3057:
3021:
2983:
2973:
2932:
2924:
2883:
2875:
2829:
2821:
2772:
2728:
2687:
2679:
2638:
2630:
2617:
Hansen MF, Neckmann U, Lavik LA, Vold T, Gilde B, Toft RK, Sjursen W (March 2014).
2589:
2548:
2538:
2497:
2489:
2448:
2440:
2426:
2399:
2391:
2350:
2342:
2293:
2285:
2240:
2232:
2191:
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2142:
2134:
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1995:
1987:
1960:
1940:
1893:
1885:
1839:
1829:
1788:
1780:
1733:
1696:
1686:
1645:
1635:
1559:
1549:
1501:
1491:
1447:
1437:
1389:
1381:
1340:
1330:
1286:
1278:
1232:
1224:
1183:
1125:
1071:
1042:
1022:
942:
932:
919:
Reneker J, Lyons E, Conant GC, Pires JC, Freeling M, Shyu CR, Korkin D (May 2012).
890:
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3652:
2578:"A genome-wide association study of sporadic ALS in a homogenous Irish population"
1538:"Highly conserved non-coding sequences are associated with vertebrate development"
4574:
3770:
3708:
3558:
3511:
3452:
2138:
2041:
2024:
1834:
1691:
1554:
1496:
1370:"Ultraconserved enhancer function does not require perfect sequence conservation"
1335:
1211:
Sathirapongsasuti JF, Sathira N, Suzuki Y, Huttenhower C, Sugano S (March 2011).
2683:
1784:
1769:"Ultraconserved elements: analyses of dosage sensitivity, motifs and boundaries"
4624:
4483:
4413:
Proceedings of the
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1171:
4716:
3093:"Mutations in the Matrin 3 gene cause familial amyotrophic lateral sclerosis"
2395:
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Wang JY, Cui YH, Xiao L, Chung HK, Zhang Y, Rao JN, et al. (July 2018).
2236:
71:
4433:
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4234:"Mutation spectrum leading to an attenuated phenotype in dystrophinopathies"
4233:
3305:
3288:
2270:"Regulation of Intestinal Epithelial Barrier Function by Long Noncoding RNA
2219:
Nan A, Zhou X, Chen L, Liu M, Zhang N, Zhang L, et al. (January 2016).
2187:
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859:
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4683:
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4582:
4536:
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4302:
4259:
4199:
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4061:
4012:
3963:
3912:
3863:
3806:
3762:
3727:
3660:
3625:
3576:
3530:
3471:
3412:
3394:
3314:
3265:
3175:
3126:
2997:
2897:
2843:
2786:
2761:"HOXD10 M319K mutation in a family with isolated congenital vertical talus"
2742:
2701:
2652:
2603:
2562:
2511:
2493:
2462:
2413:
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2156:
2107:
2050:
2009:
1952:
1907:
1853:
1802:
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1710:
1659:
1573:
1515:
1461:
1403:
1354:
1300:
1246:
1197:
1137:
1083:
1034:
956:
867:
187:
Regulation mechanisms of disease related ultraconserved element transcripts
132:
52:
4393:
4369:
4346:
4218:
3485:
Gao W, Peng Y, Liang G, Liang A, Ye W, Zhang L, et al. (2013-01-04).
3363:
3224:
3069:
3033:
2946:
1442:
1282:
4043:
4026:
Hromatka BS, Tung JY, Kiefer AK, Do CB, Hinds DA, Eriksson N (May 2015).
3994:
3257:
10.1002/(SICI)1098-1004(199911)14:5<369::AID-HUMU2>3.0.CO;2-E
2733:
2716:
2594:
2577:
2543:
2289:
2064:
Guo J, Fang W, Sun L, Lu Y, Dou L, Huang X, et al. (February 2018).
1974:
Pang W, Su J, Wang Y, Feng H, Dai X, Yuan Y, et al. (October 2015).
1944:
1265:
Habic A, Mattick JS, Calin GA, Krese R, Konc J, Kunej T (November 2019).
1228:
1210:
894:
150:
116:
48:
3188:
359:
DNA methylation/uc.158 + A, uc.160+, uc.241 + A, uc.283 + A, uc.346 + A
4338:
4142:
4125:
3894:
3567:
3157:
3062:
10.1002/(SICI)1098-1004(1998)11:3<197::AID-HUMU3>3.0.CO;2-F
56:
4528:
2777:
2760:
1991:
1075:
4606:
155:
87:
4468:"Mutations of the FHL1 gene cause Emery-Dreifuss muscular dystrophy"
3590:
Jiang H, Qiu X, Dai J, Yan H, Zhu Z, Qian B, Qiu Y (February 2013).
3108:
2825:
2634:
646:
Adolescent idiopathic scoliosis, no association with breast cancer
95:
The probability of finding ultraconserved elements by chance (under
3798:
3237:
3206:
1737:
1267:"Genetic Variations of Ultraconserved Elements in the Human Genome"
159:
142:
112:
2864:"NR2F1 mutations cause optic atrophy with intellectual disability"
921:"Long identical multispecies elements in plant and animal genomes"
4123:
4077:"Rare variants in NR2F2 cause congenital heart defects in humans"
394:
Phenotype-associated polymorphisms within ultraconserved elements
63:
4323:
4231:
3876:
844:
103:. However, affected phenotypes were only caused by 112 of these
2861:
2320:
1367:
163:
100:
90:
importance, as these regions appear to have experienced strong
32:
4074:
3425:
3139:
1815:
994:
4707:
4655:
4560:
3543:
36:
3783:
3046:
3010:
2524:
1613:
3976:
3335:
67:
3090:
2169:
3681:
3376:
2960:
Chang YS, Lin CY, Yang SF, Ho CM, Chang JG (2016-03-28).
2377:
1923:
1424:
Fedorova L, Mulyar OA, Lim J, Fedorov A (November 2022).
1107:
194:
miR/methylation/transcript factor associated with T-UCRs
141:
proposed an importance in vertebrate development. Double-
4514:
4465:
4406:
3286:
2575:
1477:
1423:
1319:"Deletion of ultraconserved elements yields viable mice"
4280:
3827:
3638:
2758:
2616:
1871:
1766:
1535:
1056:
1054:
1052:
918:
4367:
3925:
2475:
1673:
McCole RB, Fonseka CY, Koren A, Wu CT (October 2014).
1264:
1169:
4602:
4600:
682:
De et al. (2015); Guo et al. (2013); Hromatka et al.
4172:
4025:
2910:
2665:
2022:
1672:
1049:
781:
Schoser et al. (2009); Windpassinger et al. (2008)
4225:
3379:"Mutations in PAX2 associate with adult-onset FSGS"
4597:
4317:
3870:
2807:
1316:
880:
3484:
2714:
1723:
1598:Mysterious unchanging DNA finds a purpose in life
657:Peroxisomal fatty acyl-CoA reductase 1 disorder
4714:
2267:
1419:
1417:
1415:
1413:
800:Myopathy X-linked with postural muscle atrophy
789:Myopathy X-linked with postural muscle atrophy
778:Myopathy X-linked with postural muscle atrophy
770:Gueneau et al. (2009); Knoblauch et al. (2010)
4649:
4556:
4554:
4173:Roberts RG, Bobrow M, Bentley DR (March 1992).
3329:
3231:
2120:
4508:
3777:
3740:
3589:
2959:
2218:
1973:
1867:
1865:
1863:
1809:
1473:
1471:
1312:
1310:
767:Emery-Dreifuss muscular dystrophy 6 X-linked
704:Hofstra et al. (2004); Roberts et al. (1992)
503:Bosch-Boonstra-Schaaf optic atrophy syndrome
492:Bosch-Boonstra-Schaaf optic atrophy syndrome
481:Bosch-Boonstra-Schaaf optic atrophy syndrome
462:Dobbs et al. (2006); Shrimpton et al. (2004)
35:that is shared between evolutionarily distant
4068:
4019:
3419:
3383:Journal of the American Society of Nephrology
2857:
2855:
2853:
2754:
2752:
2420:
2371:
1410:
1361:
547:Familial adenomatous polyposis 1 attenuated
514:Familial adenomatous polyposis 1 attenuated
4551:
4459:
4166:
4117:
3821:
3632:
3537:
3182:
3084:
2904:
2518:
2314:
2063:
1919:
1917:
1260:
1258:
1256:
1165:
1163:
1161:
1159:
1157:
1155:
838:
4274:
3675:
2659:
2610:
2569:
2469:
2163:
1860:
1666:
1468:
1307:
690:Congenital heart defects multiple types 4
348:DNA methylation/uc.160, uc.283, and uc.346
4361:
3734:
3583:
3370:
3280:
3040:
3004:
2953:
2850:
2749:
365:Goto et al., 2016; Lujambio et al., 2010
326:miR-339-3p, miR-663b-3p, miR-95-5p/uc.339
4673:
4632:
4491:
4442:
4432:
4249:
4208:
4198:
4141:
4100:
4051:
4002:
3953:
3943:
3902:
3853:
3717:
3707:
3615:
3566:
3520:
3510:
3461:
3451:
3402:
3353:
3304:
3255:
3214:
3165:
3116:
2987:
2977:
2936:
2887:
2833:
2776:
2732:
2708:
2691:
2642:
2623:Molecular Genetics & Genomic Medicine
2593:
2552:
2542:
2501:
2452:
2403:
2354:
2297:
2261:
2244:
2195:
2146:
2114:
2097:
2040:
1999:
1967:
1914:
1897:
1843:
1833:
1792:
1700:
1690:
1649:
1639:
1563:
1553:
1505:
1495:
1451:
1441:
1393:
1344:
1334:
1290:
1253:
1236:
1187:
1152:
1016:
946:
936:
914:
912:
910:
908:
906:
904:
370:Transcription factor SP1/uc.138 (TRA2Ξ²4)
4175:"Point mutations in the dystrophin gene"
3970:
3478:
2212:
2057:
2016:
990:
988:
986:
874:
173:
4400:
3919:
3133:
2801:
1103:
1101:
984:
982:
980:
978:
976:
974:
972:
970:
968:
966:
745:Pelizaeus/Merzbacher disease atypical
233:Goto et al., 2016; Sekino et al., 2017
14:
4715:
2274:through Interaction with MicroRNA 29b"
901:
811:Rett syndrome (nonpathogenic variant)
635:Focal segmental glomerulosclerosis 7
244:Calin et al., 2007; Pang et al., 2018
4374:American Journal of Medical Genetics
1098:
1064:Wiley Interdisciplinary Reviews. RNA
963:
525:No association with breast cancer
24:
4613:American Journal of Human Genetics
4472:American Journal of Human Genetics
4295:10.1212/01.wnl.0000259522.49388.53
4238:European Journal of Human Genetics
4081:American Journal of Human Genetics
3834:American Journal of Human Genetics
3342:American Journal of Human Genetics
3293:European Journal of Human Genetics
3195:American Journal of Human Genetics
2868:American Journal of Human Genetics
569:Amyotrophic lateral sclerosis 21
536:Familial adenomatous polyposis 1
404:Associated phenotype description
25:
4734:
4701:
2917:The American Journal of Pathology
166:is strongly upregulated in human
109:ultraconserved noncoding elements
3743:European Journal of Epidemiology
1579:
1521:
883:Insect Systematics and Diversity
756:Mental retardation X-linked 94
2765:Journal of Orthopaedic Research
1760:
1717:
1607:
1587:
1529:
734:Spastic paraplegia 2 X-linked
723:Spastic paraplegia 2 X-linked
415:Amyotrophic lateral sclerosis
181:non-polyposis colorectal cancer
2278:Molecular and Cellular Biology
1204:
715:Tuffery-Giraud et al. (2005)
381:Transcription factor YY1/uc.8
92:negative (purifying) selection
13:
1:
3653:10.1136/jmedgenet-2013-102067
2929:10.1016/S0002-9440(10)65259-5
832:
803:Windpassinger et al. (2008)
737:Saugier-Veber et al. (1994)
701:Duchenne muscular dystrophy
4575:10.1212/WNL.0b013e3181b2a4b3
3709:10.1371/journal.pone.0072802
3559:10.1016/j.spinee.2015.05.013
3512:10.1371/journal.pone.0053234
3453:10.1371/journal.pone.0117708
3026:10.1016/0888-7543(92)90032-n
2139:10.1053/j.gastro.2017.10.009
2042:10.1016/j.molcel.2014.05.005
1835:10.1371/journal.pbio.1000275
1692:10.1371/journal.pgen.1004646
1555:10.1371/journal.pbio.0030007
1497:10.1371/journal.pbio.0030019
1336:10.1371/journal.pbio.0050234
81:
7:
3641:Journal of Medical Genetics
2684:10.1534/genetics.112.141945
1785:10.1534/genetics.108.096537
820:
712:Becker muscular dystrophy
122:
10:
4739:
4708:http://ultraconserved.org/
4625:10.1016/j.ajhg.2007.09.004
4484:10.1016/j.ajhg.2009.07.015
4093:10.1016/j.ajhg.2014.03.007
3846:10.1016/j.ajhg.2014.10.003
2880:10.1016/j.ajhg.2014.01.002
2347:10.1038/s41467-017-01562-9
2090:10.1038/s41467-018-03072-8
1386:10.1038/s41588-021-00812-3
814:Zahorakova et al. (2007)
668:Capecitabine sensitivity
627:Schimmenti et al. (1997)
605:Schimmenti et al. (1999)
470:Central corneal thickness
459:Vertical talus congenital
149:in mice caused a shrunken
4675:10.1007/s10038-007-0121-x
4662:Journal of Human Genetics
3945:10.1186/s13040-015-0074-0
3755:10.1007/s10654-011-9617-y
3608:10.1007/s00586-012-2532-4
2979:10.1186/s12935-016-0297-2
2966:Cancer Cell International
2445:10.18632/oncotarget.25115
1890:10.18632/oncotarget.21688
1189:10.1016/j.ccr.2007.07.027
827:Human accelerated regions
671:O'Donnell et al. (2012)
613:Renal coloboma syndrome
321:J. Y. Wang et al., 2018
271:miR-195, miR-4668/uc.372
31:(UCE) is a region of the
4032:Human Molecular Genetics
3983:Human Molecular Genetics
2721:Human Molecular Genetics
2582:Human Molecular Genetics
2396:10.18632/oncotarget.7833
2237:10.18632/oncotarget.6590
693:Al Turki et al. (2014)
561:Senderek et al. (2009)
230:Colorectal and renal CA
168:hepatocellular carcinoma
4434:10.1073/pnas.0708699104
4251:10.1038/sj.ejhg.5201478
3306:10.1038/sj.ejhg.5200539
1641:10.1073/pnas.1011098108
1130:10.1126/science.1142430
1027:10.1126/science.1098119
938:10.1073/pnas.1121356109
792:Schoser et al. (2009)
660:Buchert et al. (2014)
624:Papillorenal syndrome
602:Papillorenal syndrome
594:Mallery et al. (1998)
572:Johnson et al. (2014)
517:Soravia et al. (1999)
354:Kottorou et al., 2018
343:Olivieri et al., 2016
318:Gastrointestinal tract
285:Gastrointestinal tract
18:Ultra-conserved element
4200:10.1073/pnas.89.6.2331
3596:European Spine Journal
3395:10.1681/ASN.2013070686
2494:10.1038/oncsis.2016.18
1217:Nucleic Acids Research
726:Gorman et al. (2007)
679:Motion sickness; BMI
440:Chiang et al. (2012)
429:Hansen et al. (2014)
418:Cronin et al. (2008)
387:Terreri et al., 2016
29:ultraconserved element
2327:Nature Communications
2188:10.1093/carcin/bgp209
2070:Nature Communications
1484:Pulmonary Circulation
1443:10.3390/genes13112053
1283:10.1089/omi.2019.0156
860:10.1093/sysbio/sys004
748:Hodes et al. (1997)
638:Barua et al. (2014)
616:Amiel et al. (2000)
591:Cockayne syndrome B
550:Curia et al. (1998)
539:Fodde et al. (1992)
528:Chang et al. (2016)
506:Bosch et al. (2014)
495:Bosch et al. (2014)
484:Bosch et al. (2014)
376:Kajita et al., 2016
332:Vannini et al., 2017
222:Sekino et al., 2017
174:Role in human disease
59:except for 21 and Y.
42:ultraconserved region
2544:10.3390/genes7120127
2290:10.1128/MCB.00010-18
1945:10.1038/onc.2015.445
451:Khor et al. (2013)
299:Wojcik et al., 2010
255:Calin et al., 2007
211:Calin et al., 2007
4517:Annals of Neurology
4425:2007PNAS..10418163W
4419:(46): 18163β18168.
4191:1992PNAS...89.2331R
3700:2013PLoSO...872802M
3503:2013PLoSO...853234G
3444:2015PLoSO..1017708C
3097:Nature Neuroscience
2439:(30): 21411β21428.
2390:(15): 20636β20654.
2339:2017NatCo...8.1801V
2082:2018NatCo...9..612G
1884:(55): 94259β94270.
1632:2011PNAS..108..786B
1122:2007Sci...317..915K
1009:2004Sci...304.1321B
1003:(5675): 1321β1325.
931:(19): E1183βE1191.
580:Au-Kline syndrome
401:Polymorphism name
304:miR-291a-3p/uc.173
4339:10.1038/ng0394-257
4143:10.1002/humu.10283
4044:10.1093/hmg/ddv028
3995:10.1093/hmg/dds396
3895:10.1002/cncr.26737
3158:10.1002/humu.22837
2734:10.1093/hmg/ddt385
2595:10.1093/hmg/ddm361
1229:10.1093/nar/gkq949
895:10.1093/isd/ixz016
848:Systematic Biology
759:Wu et al. (2007)
583:Au et al. (2015)
473:Lu et al. (2013)
310:Nan et al., 2016
288:Xiao et al., 2018
277:Guo et al., 2018
266:Liz et al., 2014
4529:10.1002/ana.21839
4289:(16): 1305β1307.
4244:(12): 1254β1260.
3889:(16): 4063β4073.
3793:(12): 1237β1240.
3553:(10): 2239β2246.
3547:The Spine Journal
3152:(10): 1009β1014.
2778:10.1002/jor.20052
2727:(25): 5288β5294.
2284:(13): e00010β18.
1992:10.1111/cas.12747
1986:(10): 1362β1369.
1939:(27): 3598β3606.
1732:(10): 1216β1220.
1594:Elizabeth Pennisi
1076:10.1002/wrna.1567
818:
817:
558:Distal myopathy
391:
390:
274:Lipid metabolism
145:of UCEs near the
138:Takifugu rubripes
97:neutral evolution
16:(Redirected from
4730:
4696:
4695:
4677:
4653:
4647:
4646:
4636:
4604:
4595:
4594:
4558:
4549:
4548:
4512:
4506:
4505:
4495:
4463:
4457:
4456:
4446:
4436:
4404:
4398:
4397:
4365:
4359:
4358:
4321:
4315:
4314:
4278:
4272:
4271:
4253:
4229:
4223:
4222:
4212:
4202:
4185:(6): 2331β2335.
4170:
4164:
4163:
4145:
4121:
4115:
4114:
4104:
4072:
4066:
4065:
4055:
4038:(9): 2700β2708.
4023:
4017:
4016:
4006:
3974:
3968:
3967:
3957:
3947:
3923:
3917:
3916:
3906:
3874:
3868:
3867:
3857:
3825:
3819:
3818:
3781:
3775:
3774:
3738:
3732:
3731:
3721:
3711:
3679:
3673:
3672:
3636:
3630:
3629:
3619:
3587:
3581:
3580:
3570:
3541:
3535:
3534:
3524:
3514:
3482:
3476:
3475:
3465:
3455:
3423:
3417:
3416:
3406:
3389:(9): 1942β1953.
3374:
3368:
3367:
3357:
3333:
3327:
3326:
3308:
3284:
3278:
3277:
3259:
3235:
3229:
3228:
3218:
3186:
3180:
3179:
3169:
3137:
3131:
3130:
3120:
3088:
3082:
3081:
3044:
3038:
3037:
3020:(4): 1162β1168.
3008:
3002:
3001:
2991:
2981:
2957:
2951:
2950:
2940:
2908:
2902:
2901:
2891:
2859:
2848:
2847:
2837:
2805:
2799:
2798:
2780:
2756:
2747:
2746:
2736:
2712:
2706:
2705:
2695:
2663:
2657:
2656:
2646:
2614:
2608:
2607:
2597:
2573:
2567:
2566:
2556:
2546:
2522:
2516:
2515:
2505:
2473:
2467:
2466:
2456:
2424:
2418:
2417:
2407:
2375:
2369:
2368:
2358:
2318:
2312:
2311:
2301:
2265:
2259:
2258:
2248:
2216:
2210:
2209:
2199:
2167:
2161:
2160:
2150:
2127:Gastroenterology
2118:
2112:
2111:
2101:
2061:
2055:
2054:
2044:
2020:
2014:
2013:
2003:
1971:
1965:
1964:
1930:
1921:
1912:
1911:
1901:
1869:
1858:
1857:
1847:
1837:
1813:
1807:
1806:
1796:
1779:(4): 2277β2293.
1764:
1758:
1757:
1721:
1715:
1714:
1704:
1694:
1685:(10): e1004646.
1670:
1664:
1663:
1653:
1643:
1611:
1605:
1591:
1585:
1584:
1583:
1577:
1567:
1557:
1533:
1527:
1526:
1525:
1519:
1509:
1499:
1475:
1466:
1465:
1455:
1445:
1421:
1408:
1407:
1397:
1365:
1359:
1358:
1348:
1338:
1314:
1305:
1304:
1294:
1262:
1251:
1250:
1240:
1223:(6): 1967β1979.
1208:
1202:
1201:
1191:
1167:
1150:
1149:
1105:
1096:
1095:
1058:
1047:
1046:
1020:
992:
961:
960:
950:
940:
916:
899:
898:
878:
872:
871:
842:
426:Lynch syndrome
398:
397:
205:miR-24-1/uc.160
191:
190:
21:
4738:
4737:
4733:
4732:
4731:
4729:
4728:
4727:
4713:
4712:
4704:
4699:
4654:
4650:
4605:
4598:
4559:
4552:
4513:
4509:
4464:
4460:
4405:
4401:
4366:
4362:
4327:Nature Genetics
4322:
4318:
4279:
4275:
4230:
4226:
4171:
4167:
4122:
4118:
4073:
4069:
4024:
4020:
3975:
3971:
3924:
3920:
3875:
3871:
3826:
3822:
3787:Nature Genetics
3782:
3778:
3749:(11): 863β876.
3739:
3735:
3680:
3676:
3637:
3633:
3588:
3584:
3542:
3538:
3483:
3479:
3438:(2): e0117708.
3424:
3420:
3375:
3371:
3334:
3330:
3299:(11): 820β826.
3285:
3281:
3236:
3232:
3187:
3183:
3138:
3134:
3109:10.1038/nn.3688
3089:
3085:
3045:
3041:
3009:
3005:
2958:
2954:
2909:
2905:
2860:
2851:
2826:10.1038/ng.2506
2814:Nature Genetics
2806:
2802:
2757:
2750:
2713:
2709:
2664:
2660:
2635:10.1002/mgg3.62
2615:
2611:
2574:
2570:
2523:
2519:
2474:
2470:
2425:
2421:
2376:
2372:
2319:
2315:
2266:
2262:
2217:
2213:
2168:
2164:
2119:
2115:
2062:
2058:
2021:
2017:
1972:
1968:
1928:
1922:
1915:
1870:
1861:
1828:(1): e1000275.
1814:
1810:
1765:
1761:
1726:Nature Genetics
1722:
1718:
1671:
1667:
1612:
1608:
1592:
1588:
1578:
1534:
1530:
1520:
1476:
1469:
1422:
1411:
1374:Nature Genetics
1366:
1362:
1315:
1308:
1277:(11): 549β559.
1263:
1254:
1209:
1205:
1168:
1153:
1106:
1099:
1059:
1050:
1018:10.1.1.380.9305
993:
964:
917:
902:
879:
875:
843:
839:
835:
823:
396:
315:miR-29b/uc.173
307:Nervous system
293:miR-214/uc.276
282:mir-195/uc.173
260:mir-195/uc.283
249:miR-155/uc346A
238:miR-155/uc.160
227:miR-153/uc.416
216:miR-130b/uc.63
189:
176:
162:. For example,
125:
84:
23:
22:
15:
12:
11:
5:
4736:
4726:
4725:
4711:
4710:
4703:
4702:External links
4700:
4698:
4697:
4668:(4): 342β348.
4648:
4596:
4569:(7): 543β551.
4550:
4523:(1): 136β140.
4507:
4478:(3): 338β353.
4458:
4399:
4380:(2): 121β125.
4360:
4333:(3): 257β262.
4316:
4273:
4224:
4165:
4130:Human Mutation
4116:
4087:(4): 574β585.
4067:
4018:
3989:(1): 184β201.
3969:
3932:BioData Mining
3918:
3869:
3840:(5): 602β610.
3820:
3799:10.1038/ng.974
3776:
3733:
3674:
3647:(6): 401β406.
3631:
3602:(2): 282β286.
3582:
3536:
3477:
3418:
3369:
3348:(4): 869β878.
3328:
3279:
3250:(5): 369β376.
3244:Human Mutation
3230:
3207:10.1086/301686
3181:
3146:Human Mutation
3132:
3103:(5): 664β666.
3083:
3056:(3): 197β201.
3050:Human Mutation
3039:
3003:
2952:
2923:(1): 127β135.
2903:
2874:(2): 303β309.
2849:
2820:(2): 155β163.
2800:
2771:(3): 448β453.
2748:
2707:
2678:(1): 253β266.
2658:
2629:(2): 186β200.
2609:
2588:(5): 768β774.
2568:
2517:
2468:
2419:
2370:
2313:
2260:
2231:(1): 112β124.
2211:
2182:(2): 208β215.
2176:Carcinogenesis
2162:
2133:(3): 599β611.
2113:
2056:
2035:(1): 138β147.
2029:Molecular Cell
2015:
1980:Cancer Science
1966:
1913:
1859:
1808:
1759:
1738:10.1038/ng1888
1716:
1665:
1626:(2): 786β791.
1606:
1586:
1528:
1467:
1409:
1380:(4): 521β528.
1360:
1306:
1252:
1203:
1182:(3): 215β229.
1151:
1097:
1048:
962:
900:
873:
836:
834:
831:
830:
829:
822:
819:
816:
815:
812:
809:
805:
804:
801:
798:
794:
793:
790:
787:
783:
782:
779:
776:
772:
771:
768:
765:
761:
760:
757:
754:
750:
749:
746:
743:
739:
738:
735:
732:
728:
727:
724:
721:
717:
716:
713:
710:
706:
705:
702:
699:
695:
694:
691:
688:
684:
683:
680:
677:
673:
672:
669:
666:
662:
661:
658:
655:
651:
650:
647:
644:
640:
639:
636:
633:
629:
628:
625:
622:
618:
617:
614:
611:
607:
606:
603:
600:
596:
595:
592:
589:
585:
584:
581:
578:
574:
573:
570:
567:
563:
562:
559:
556:
552:
551:
548:
545:
541:
540:
537:
534:
530:
529:
526:
523:
519:
518:
515:
512:
508:
507:
504:
501:
497:
496:
493:
490:
486:
485:
482:
479:
475:
474:
471:
468:
464:
463:
460:
457:
453:
452:
449:
448:Severe myopia
446:
442:
441:
438:
435:
431:
430:
427:
424:
420:
419:
416:
413:
409:
408:
405:
402:
395:
392:
389:
388:
385:
382:
378:
377:
374:
373:Colorectal CA
371:
367:
366:
363:
360:
356:
355:
352:
351:Colorectal CA
349:
345:
344:
341:
338:
334:
333:
330:
327:
323:
322:
319:
316:
312:
311:
308:
305:
301:
300:
297:
296:Colorectal CA
294:
290:
289:
286:
283:
279:
278:
275:
272:
268:
267:
264:
261:
257:
256:
253:
250:
246:
245:
242:
239:
235:
234:
231:
228:
224:
223:
220:
217:
213:
212:
209:
206:
202:
201:
198:
195:
188:
185:
175:
172:
124:
121:
83:
80:
76:non-coding RNA
9:
6:
4:
3:
2:
4735:
4724:
4721:
4720:
4718:
4709:
4706:
4705:
4693:
4689:
4685:
4681:
4676:
4671:
4667:
4663:
4659:
4652:
4644:
4640:
4635:
4630:
4626:
4622:
4618:
4614:
4610:
4603:
4601:
4592:
4588:
4584:
4580:
4576:
4572:
4568:
4564:
4557:
4555:
4546:
4542:
4538:
4534:
4530:
4526:
4522:
4518:
4511:
4503:
4499:
4494:
4489:
4485:
4481:
4477:
4473:
4469:
4462:
4454:
4450:
4445:
4440:
4435:
4430:
4426:
4422:
4418:
4414:
4410:
4403:
4395:
4391:
4387:
4383:
4379:
4375:
4371:
4364:
4356:
4352:
4348:
4344:
4340:
4336:
4332:
4328:
4320:
4312:
4308:
4304:
4300:
4296:
4292:
4288:
4284:
4277:
4269:
4265:
4261:
4257:
4252:
4247:
4243:
4239:
4235:
4228:
4220:
4216:
4211:
4206:
4201:
4196:
4192:
4188:
4184:
4180:
4176:
4169:
4161:
4157:
4153:
4149:
4144:
4139:
4135:
4131:
4127:
4120:
4112:
4108:
4103:
4098:
4094:
4090:
4086:
4082:
4078:
4071:
4063:
4059:
4054:
4049:
4045:
4041:
4037:
4033:
4029:
4022:
4014:
4010:
4005:
4000:
3996:
3992:
3988:
3984:
3980:
3973:
3965:
3961:
3956:
3951:
3946:
3941:
3937:
3933:
3929:
3922:
3914:
3910:
3905:
3900:
3896:
3892:
3888:
3884:
3880:
3873:
3865:
3861:
3856:
3851:
3847:
3843:
3839:
3835:
3831:
3824:
3816:
3812:
3808:
3804:
3800:
3796:
3792:
3788:
3780:
3772:
3768:
3764:
3760:
3756:
3752:
3748:
3744:
3737:
3729:
3725:
3720:
3715:
3710:
3705:
3701:
3697:
3694:(9): e72802.
3693:
3689:
3685:
3678:
3670:
3666:
3662:
3658:
3654:
3650:
3646:
3642:
3635:
3627:
3623:
3618:
3613:
3609:
3605:
3601:
3597:
3593:
3586:
3578:
3574:
3569:
3564:
3560:
3556:
3552:
3548:
3540:
3532:
3528:
3523:
3518:
3513:
3508:
3504:
3500:
3497:(1): e53234.
3496:
3492:
3488:
3481:
3473:
3469:
3464:
3459:
3454:
3449:
3445:
3441:
3437:
3433:
3429:
3422:
3414:
3410:
3405:
3400:
3396:
3392:
3388:
3384:
3380:
3373:
3365:
3361:
3356:
3351:
3347:
3343:
3339:
3332:
3324:
3320:
3316:
3312:
3307:
3302:
3298:
3294:
3290:
3283:
3275:
3271:
3267:
3263:
3258:
3253:
3249:
3245:
3241:
3234:
3226:
3222:
3217:
3212:
3208:
3204:
3200:
3196:
3192:
3185:
3177:
3173:
3168:
3163:
3159:
3155:
3151:
3147:
3143:
3136:
3128:
3124:
3119:
3114:
3110:
3106:
3102:
3098:
3094:
3087:
3079:
3075:
3071:
3067:
3063:
3059:
3055:
3051:
3043:
3035:
3031:
3027:
3023:
3019:
3015:
3007:
2999:
2995:
2990:
2985:
2980:
2975:
2971:
2967:
2963:
2956:
2948:
2944:
2939:
2934:
2930:
2926:
2922:
2918:
2914:
2907:
2899:
2895:
2890:
2885:
2881:
2877:
2873:
2869:
2865:
2858:
2856:
2854:
2845:
2841:
2836:
2831:
2827:
2823:
2819:
2815:
2811:
2804:
2796:
2792:
2788:
2784:
2779:
2774:
2770:
2766:
2762:
2755:
2753:
2744:
2740:
2735:
2730:
2726:
2722:
2718:
2711:
2703:
2699:
2694:
2689:
2685:
2681:
2677:
2673:
2669:
2662:
2654:
2650:
2645:
2640:
2636:
2632:
2628:
2624:
2620:
2613:
2605:
2601:
2596:
2591:
2587:
2583:
2579:
2572:
2564:
2560:
2555:
2550:
2545:
2540:
2536:
2532:
2528:
2521:
2513:
2509:
2504:
2499:
2495:
2491:
2487:
2483:
2479:
2472:
2464:
2460:
2455:
2450:
2446:
2442:
2438:
2434:
2430:
2423:
2415:
2411:
2406:
2401:
2397:
2393:
2389:
2385:
2381:
2374:
2366:
2362:
2357:
2352:
2348:
2344:
2340:
2336:
2332:
2328:
2324:
2317:
2309:
2305:
2300:
2295:
2291:
2287:
2283:
2279:
2275:
2273:
2264:
2256:
2252:
2247:
2242:
2238:
2234:
2230:
2226:
2222:
2215:
2207:
2203:
2198:
2193:
2189:
2185:
2181:
2177:
2173:
2166:
2158:
2154:
2149:
2144:
2140:
2136:
2132:
2128:
2124:
2117:
2109:
2105:
2100:
2095:
2091:
2087:
2083:
2079:
2075:
2071:
2067:
2060:
2052:
2048:
2043:
2038:
2034:
2030:
2026:
2019:
2011:
2007:
2002:
1997:
1993:
1989:
1985:
1981:
1977:
1970:
1962:
1958:
1954:
1950:
1946:
1942:
1938:
1934:
1927:
1920:
1918:
1909:
1905:
1900:
1895:
1891:
1887:
1883:
1879:
1875:
1868:
1866:
1864:
1855:
1851:
1846:
1841:
1836:
1831:
1827:
1823:
1819:
1812:
1804:
1800:
1795:
1790:
1786:
1782:
1778:
1774:
1770:
1763:
1755:
1751:
1747:
1743:
1739:
1735:
1731:
1727:
1720:
1712:
1708:
1703:
1698:
1693:
1688:
1684:
1680:
1679:PLOS Genetics
1676:
1669:
1661:
1657:
1652:
1647:
1642:
1637:
1633:
1629:
1625:
1621:
1617:
1610:
1603:
1599:
1595:
1590:
1582:
1575:
1571:
1566:
1561:
1556:
1551:
1547:
1543:
1539:
1532:
1524:
1517:
1513:
1508:
1503:
1498:
1493:
1489:
1485:
1481:
1474:
1472:
1463:
1459:
1454:
1449:
1444:
1439:
1435:
1431:
1427:
1420:
1418:
1416:
1414:
1405:
1401:
1396:
1391:
1387:
1383:
1379:
1375:
1371:
1364:
1356:
1352:
1347:
1342:
1337:
1332:
1328:
1324:
1320:
1313:
1311:
1302:
1298:
1293:
1288:
1284:
1280:
1276:
1272:
1268:
1261:
1259:
1257:
1248:
1244:
1239:
1234:
1230:
1226:
1222:
1218:
1214:
1207:
1199:
1195:
1190:
1185:
1181:
1177:
1173:
1166:
1164:
1162:
1160:
1158:
1156:
1147:
1143:
1139:
1135:
1131:
1127:
1123:
1119:
1116:(5840): 915.
1115:
1111:
1104:
1102:
1093:
1089:
1085:
1081:
1077:
1073:
1069:
1065:
1057:
1055:
1053:
1044:
1040:
1036:
1032:
1028:
1024:
1019:
1014:
1010:
1006:
1002:
998:
991:
989:
987:
985:
983:
981:
979:
977:
975:
973:
971:
969:
967:
958:
954:
949:
944:
939:
934:
930:
926:
922:
915:
913:
911:
909:
907:
905:
896:
892:
888:
884:
877:
869:
865:
861:
857:
854:(5): 717β26.
853:
849:
841:
837:
828:
825:
824:
813:
810:
807:
806:
802:
799:
796:
795:
791:
788:
785:
784:
780:
777:
774:
773:
769:
766:
763:
762:
758:
755:
752:
751:
747:
744:
741:
740:
736:
733:
730:
729:
725:
722:
719:
718:
714:
711:
708:
707:
703:
700:
697:
696:
692:
689:
686:
685:
681:
678:
675:
674:
670:
667:
664:
663:
659:
656:
653:
652:
648:
645:
642:
641:
637:
634:
631:
630:
626:
623:
620:
619:
615:
612:
609:
608:
604:
601:
598:
597:
593:
590:
587:
586:
582:
579:
576:
575:
571:
568:
565:
564:
560:
557:
554:
553:
549:
546:
543:
542:
538:
535:
532:
531:
527:
524:
521:
520:
516:
513:
510:
509:
505:
502:
499:
498:
494:
491:
488:
487:
483:
480:
477:
476:
472:
469:
466:
465:
461:
458:
455:
454:
450:
447:
444:
443:
439:
436:
433:
432:
428:
425:
422:
421:
417:
414:
411:
410:
406:
403:
400:
399:
386:
383:
380:
379:
375:
372:
369:
368:
364:
361:
358:
357:
353:
350:
347:
346:
342:
339:
337:miR-596/uc.8
336:
335:
331:
328:
325:
324:
320:
317:
314:
313:
309:
306:
303:
302:
298:
295:
292:
291:
287:
284:
281:
280:
276:
273:
270:
269:
265:
262:
259:
258:
254:
251:
248:
247:
243:
240:
237:
236:
232:
229:
226:
225:
221:
218:
215:
214:
210:
207:
204:
203:
199:
196:
193:
192:
184:
182:
171:
169:
165:
161:
157:
152:
148:
144:
140:
139:
134:
130:
120:
118:
114:
110:
106:
105:polymorphisms
102:
98:
93:
89:
79:
77:
73:
72:noncoding DNA
69:
65:
60:
58:
54:
50:
45:
43:
38:
34:
30:
19:
4665:
4661:
4651:
4619:(1): 88β99.
4616:
4612:
4566:
4562:
4520:
4516:
4510:
4475:
4471:
4461:
4416:
4412:
4402:
4377:
4373:
4363:
4330:
4326:
4319:
4286:
4282:
4276:
4241:
4237:
4227:
4182:
4178:
4168:
4136:(1): 57β66.
4133:
4129:
4119:
4084:
4080:
4070:
4035:
4031:
4021:
3986:
3982:
3972:
3935:
3931:
3921:
3886:
3882:
3872:
3837:
3833:
3823:
3790:
3786:
3779:
3746:
3742:
3736:
3691:
3687:
3677:
3644:
3640:
3634:
3599:
3595:
3585:
3550:
3546:
3539:
3494:
3490:
3480:
3435:
3431:
3421:
3386:
3382:
3372:
3345:
3341:
3331:
3296:
3292:
3282:
3247:
3243:
3233:
3201:(1): 77β85.
3198:
3194:
3184:
3149:
3145:
3135:
3100:
3096:
3086:
3053:
3049:
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