242:
219:
116:
141:
248:
147:
1202:, characterized by bone marrow failure, premature ageing, predisposition for cancer and a triad of mucocutaneous features including oral leukoplakia, abnormal skin pigmentation and nail dystrophy. Mutations in WRAP53β are inherited in an autosomal recessive fashion, reside in highly conserved regions of its WD40 domain and result in a more severe form of this disease.
1105:
1171:
1097:
1155:), which belongs to the WD40 protein family (Figure 1). WRAP53β facilitates protein-protein and protein-RNA interactions and directs factors to nuclear organelles Cajal bodies, to telomeres and to DNA double-strand breaks. Factors localized to Cajal bodies with the help of WRAP53β includes the SMN protein, small Cajal body-specific
1222:), a neurodegenerative disorder characterized by progressive degeneration of spinal cord anterior horn α-motor neurons and the leading genetic cause of infant mortality with an incidence of approximately 1:6000 live births. Mutations in the SMN1 gene are the underlying cause to spinal muscular atrophy (SMA).
1230:
WRAP53β is overexpressed in a variety of cancer cell lines of different origins and such overexpression promotes carcinogenic transformation indicating that this protein possesses oncogenic properties. WRAP53β is overexpressed in primary nasopharyngeal carcinoma, esophageal squamous cell carcinoma
1234:
In contrary, inactivating mutations in both alleles of WRAP53β causes dyskeratosis congenita, indicating that this protein acts as tumor suppressor, rather than an oncogene. Loss of nuclear WRAP53β is also correlated with shortened survival and resistance to radiotherapy in patients with head and
1247:
The WRAP53 gene was first identified by
Marianne Farnebo (maiden name Hammarsund) at Karolinska Institutet in 2006. In 2009 the WRAP53β (alias TCAB1, WDR79 and WRAP53) protein was described by Steven Artandi (Stanford University), Joan Steitz (Yale University) and Marianne Farnebo (Karolinska
1205:
These mutations reduce the nuclear level of WRAP53β, impair its trafficking of telomerase to telomeres, and subsequently lead to progressive shortening of telomeres in these patients The chaperonin CCT/TRiC is crucial for proper folding of WRAP53β and this folding is impaired in dyskeratosis
1182:) in vertebrates, invertebrates, plants and yeast. WRAP53β consists of a proline-rich N-terminus, a central WD40 domain and a glycine-rich C-terminus (Figure 3). The WD40 domain of WRAP53β serves as a scaffold for multiple interactions between a wide variety of molecules.
2291:
Garvin S, Tiefenböck K, Farnebo L, Thunell LK, Farnebo M, Roberg K (January 2015). "Nuclear expression of WRAP53β is associated with a positive response to radiotherapy and improved overall survival in patients with head and neck squamous cell carcinoma".
1238:
Single nucleotide polymorphisms (SNPs) in the WRAP53 gene have been linked to an increased risk for breast and ovarian cancer. One of these SNPs also associate with defective DNA repair and hematotoxicity in workers exposed to benzene.
1988:
Tapia O, Bengoechea R, Palanca A, Arteaga R, Val-Bernal JF, Tizzano EF, et al. (May 2012). "Reorganization of Cajal bodies and nucleolar targeting of coilin in motor neurons of type I spinal muscular atrophy".
1231:
and rectal cancer. Moreover, knockdown of WRAP53β in cancer cells reduced the size of the tumors formed when these are grafted into mice and triggers mitochondrial-dependent apoptosis in cancer cells.
1162:
In addition to localizing factors to correct cellular sites, WRAP53β maintain structural integrity of Cajal bodies and without WRAP53β these organelles collapse and cannot re-form (Figure 2).
1945:
Lefebvre S, Burlet P, Liu Q, Bertrandy S, Clermont O, Munnich A, et al. (July 1997). "Correlation between severity and SMN protein level in spinal muscular atrophy".
1235:
neck cancer. With its complex roles in a number of cellular processes, WRAP53β may act as a tumor suppressor under certain conditions and as an oncogene in under others.
255:
154:
1100:
Figure1.The WRAP53 and p53 genes and their gene products. The arrows indicate the direction of transcription and the dotted lines the exon overlap between these genes.
1080:
gene is localized on chromosome 17p13.1 and contains 13 exons, including three alternative starting exons (1α, 1β and 1γ) generating at least three gene products. The
732:
2476:"Large-scale evaluation of candidate genes identifies associations between DNA repair and genomic maintenance and development of benzene hematotoxicity"
713:
2242:"WRAP53 is an independent prognostic factor in rectal cancer- a study of Swedish clinical trial of preoperative radiotherapy in rectal cancer patients"
77:
1325:
2529:
1307:
1120:(referred to as WRAP53α transcripts) regulate the levels of p53 mRNA and protein (Figure 1). WRAP53γ transcripts overlap the first intron of
2425:
Schildkraut JM, Goode EL, Clyde MA, Iversen ES, Moorman PG, Berchuck A, et al. (Australian
Ovarian Cancer Study Group) (March 2009).
241:
971:
218:
964:
1294:
1273:
140:
115:
1290:
1269:
57:
254:
153:
1610:"WRAP53 is essential for Cajal body formation and for targeting the survival of motor neuron complex to Cajal bodies"
1554:
Saldaña-Meyer R, González-Buendía E, Guerrero G, Narendra V, Bonasio R, Recillas-Targa F, Reinberg D (April 2014).
247:
146:
1159:
and the enzyme telomerase. WRAP53β also targets the ubiquitin ligase RNF8 to DNA double-strand breaks(Figure 2).
2329:"Common genetic variation in TP53 and its flanking genes, WDR79 and ATP1B2, and susceptibility to breast cancer"
2534:
2183:"Overexpression of WRAP53 is associated with development and progression of esophageal squamous cell carcinoma"
1219:
777:
65:
1664:"The scaffold protein WRAP53β orchestrates the ubiquitin response critical for DNA double-strand break repair"
2427:"Single nucleotide polymorphisms in the TP53 region and susceptibility to invasive epithelial ovarian cancer"
1662:
Henriksson S, Rassoolzadeh H, Hedström E, Coucoravas C, Julner A, Goldstein M, et al. (December 2014).
1556:"CTCF regulates the human p53 gene through direct interaction with its natural antisense transcript, Wrap53"
1215:
758:
1214:
Defective WRAP53β-mediated trafficking of SMN is observed in patients afflicted by the most severe form of
1199:
1851:"Updates on the biology and management of dyskeratosis congenita and related telomere biology disorders"
129:
1608:
Mahmoudi S, Henriksson S, Weibrecht I, Smith S, Söderberg O, Strömblad S, et al. (November 2010).
2194:
1456:
44:
2327:
Garcia-Closas M, Kristensen V, Langerød A, Qi Y, Yeager M, Burdett L, et al. (December 2007).
1445:"A human telomerase holoenzyme protein required for Cajal body localization and telomere synthesis"
947:
918:
914:
910:
884:
855:
851:
847:
2376:
Mędrek K, Magnowski P, Masojć B, Chudecka-Głaz A, Torbe B, Menkiszak J, et al. (March 2013).
1038:
development. The name was coined in 2009 to describe the dual role of this gene, encoding both an
2358:
2057:
2014:
1970:
1878:
1420:
89:
943:
922:
880:
859:
1896:
Freund A, Zhong FL, Venteicher AS, Meng Z, Veenstra TD, Frydman J, Artandi SE (December 2014).
1443:
Venteicher AS, Abreu EB, Meng Z, McCann KE, Terns RM, Veenstra TD, et al. (January 2009).
2505:
2456:
2407:
2350:
2309:
2273:
2222:
2163:
2109:
2049:
2006:
1962:
1927:
1870:
1831:
1796:
1742:
1693:
1641:
1585:
1536:
1482:
1412:
1371:
1195:
37:
2495:
2487:
2446:
2438:
2397:
2389:
2340:
2301:
2263:
2253:
2212:
2202:
2153:
2143:
2099:
2091:
2041:
1998:
1954:
1917:
1909:
1862:
1823:
1786:
1778:
1732:
1724:
1683:
1675:
1631:
1621:
1575:
1567:
1526:
1518:
1472:
1464:
1402:
1361:
1046:
334:
265:
209:
164:
85:
1348:
Mahmoudi S, Henriksson S, Corcoran M, Méndez-Vidal C, Wiman KG, Farnebo M (February 2009).
309:
2378:"Association of common WRAP 53 variant with ovarian cancer risk in the Polish population"
2198:
1460:
2500:
2475:
2451:
2426:
2402:
2377:
2305:
2268:
2241:
2217:
2182:
2158:
2131:
2104:
2079:
1922:
1897:
1791:
1766:
1737:
1713:"A conserved WD40 protein binds the Cajal body localization signal of scaRNP particles"
1712:
1688:
1663:
1636:
1609:
1580:
1555:
1531:
1507:"A conserved WD40 protein binds the Cajal body localization signal of scaRNP particles"
1506:
1477:
1444:
1350:"Wrap53, a natural p53 antisense transcript required for p53 induction upon DNA damage"
1767:"Disruption of telomerase trafficking by TCAB1 mutation causes dyskeratosis congenita"
1765:
Zhong F, Savage SA, Shkreli M, Giri N, Jessop L, Myers T, et al. (January 2011).
647:
642:
637:
632:
627:
622:
617:
612:
596:
591:
586:
581:
576:
571:
566:
550:
545:
540:
535:
530:
525:
2523:
1178:
The WRAP53β protein is highly evolutionary conserved, with homologs (confined to its
1039:
512:
2362:
2061:
2018:
1974:
1898:"Proteostatic control of telomerase function through TRiC-mediated folding of TCAB1"
1882:
1827:
1424:
69:
2080:"WRAP53 promotes cancer cell survival and is a potential target for cancer therapy"
2032:
Coady TH, Lorson CL (2011). "SMN in spinal muscular atrophy and snRNP biogenesis".
327:
106:
2442:
2207:
2132:"TCAB1: a potential target for diagnosis and therapy of head and neck carcinomas"
1728:
1626:
1522:
1366:
1349:
93:
1330:
National Center for
Biotechnology Information, U.S. National Library of Medicine
1312:
National Center for
Biotechnology Information, U.S. National Library of Medicine
1179:
1913:
410:
2474:
Lan Q, Zhang L, Shen M, Jo WJ, Vermeulen R, Li G, et al. (January 2009).
2393:
2002:
1065:
1062:
226:
123:
73:
2258:
2148:
1174:
Figure 3. The structure of WRAP53β. The numbers indicate amino acid residues.
2491:
1468:
677:
470:
348:
293:
280:
192:
179:
81:
2509:
2460:
2411:
2354:
2313:
2277:
2226:
2167:
2113:
2053:
2010:
1931:
1874:
1835:
1800:
1746:
1697:
1679:
1645:
1589:
1571:
1540:
1486:
1416:
1375:
2095:
1966:
1011:
1006:
2078:
Mahmoudi S, Henriksson S, Farnebo L, Roberg K, Farnebo M (January 2011).
1407:
1390:
1125:
1058:
995:
822:
803:
1782:
618:
positive regulation of establishment of protein localization to telomere
1958:
1050:
789:
744:
2345:
2328:
1866:
1170:
1104:
2130:
Sun CK, Luo XB, Gou YP, Hu L, Wang K, Li C, et al. (July 2014).
1156:
1035:
979:
699:
2045:
1850:
1096:
1169:
1103:
1095:
662:
658:
1031:
61:
1054:
2181:
Rao X, Huang D, Sui X, Liu G, Song X, Xie J, Huang D (2014).
1816:
Hematology. American
Society of Hematology. Education Program
48:, DKCB3, TCAB1, WDR79, WD repeat containing antisense to TP53
1124:
and are antisense to the previously identified transcript
317:
1711:
Tycowski KT, Shu MD, Kukoyi A, Steitz JA (April 2009).
1505:
Tycowski KT, Shu MD, Kukoyi A, Steitz JA (April 2009).
1128:
localized within this intron. However, the function of
1088:
tumor suppressor gene in a head-to-head orientation.
482:
936:
903:
873:
840:
1657:
1655:
1286:
1284:
1282:
1265:
1263:
1261:
1438:
1436:
1434:
638:establishment of protein localization to telomere
264:
163:
2073:
2071:
1603:
1601:
1599:
2240:Zhang H, Wang DW, Adell G, Sun XF (July 2012).
1500:
1498:
1496:
1343:
1341:
1339:
16:Protein-coding gene in the species Homo sapiens
1291:GRCm38: Ensembl release 89: ENSMUSG00000041346
1026:(also known as WD40-encoding RNA antisense to
1760:
1758:
1756:
8:
2125:
2123:
1270:GRCh38: Ensembl release 89: ENSG00000141499
1814:Dokal I (2011). "Dyskeratosis congenita".
673:
633:positive regulation of telomerase activity
508:
305:
204:
101:
2499:
2450:
2401:
2344:
2267:
2257:
2216:
2206:
2157:
2147:
2103:
1921:
1790:
1736:
1687:
1635:
1625:
1579:
1530:
1476:
1406:
1365:
613:telomerase RNA localization to Cajal body
1257:
1198:in WRAP53β result in disorder known as
1061:elongation and maintenance of nuclear
18:
269:
230:
225:
168:
127:
122:
7:
2034:Wiley Interdisciplinary Reviews. RNA
1139:gene also encodes a protein termed
648:telomere maintenance via telomerase
2306:10.1016/j.oraloncology.2014.10.003
1849:Ballew BJ, Savage SA (June 2013).
1391:"Wrap53, a novel regulator of p53"
933:
900:
870:
837:
813:
794:
768:
749:
723:
704:
628:protein localization to Cajal body
551:protein-containing complex binding
487:
405:
343:
322:
14:
643:telomere formation via telomerase
623:scaRNA localization to Cajal body
1401:(15). Georgetown, Tex.: 2343–6.
253:
246:
240:
217:
152:
145:
139:
114:
2333:International Journal of Cancer
1991:Histochemistry and Cell Biology
1828:10.1182/asheducation-2011.1.480
1455:(5914). New York, N.Y.: 644–8.
1116:that overlap the first exon of
1108:Figure 2. Functions of WRAP53β
471:More reference expression data
379:olfactory zone of nasal mucosa
1:
2443:10.1158/0008-5472.CAN-08-2902
572:telomerase holoenzyme complex
238:
137:
2530:Genes on human chromosome 17
2208:10.1371/journal.pone.0091670
1729:10.1016/j.molcel.2009.02.020
1627:10.1371/journal.pbio.1000521
1523:10.1016/j.molcel.2009.02.020
1367:10.1016/j.molcel.2009.01.028
1084:gene partially overlaps the
1855:Expert Review of Hematology
1248:Institutet) independently.
2551:
1914:10.1016/j.cell.2014.10.059
395:mucosa of transverse colon
2394:10.1007/s11033-012-2273-9
2382:Molecular Biology Reports
2003:10.1007/s00418-012-0921-8
1389:Farnebo M (August 2009).
1326:"Mouse PubMed Reference:"
1308:"Human PubMed Reference:"
1010:
1005:
1001:
994:
978:
959:
940:
907:
896:
877:
844:
833:
820:
816:
801:
797:
788:
775:
771:
756:
752:
743:
730:
726:
711:
707:
698:
683:
676:
672:
656:
546:identical protein binding
511:
507:
495:
490:
481:
468:
417:
408:
367:bronchial epithelial cell
355:
346:
316:
308:
304:
287:
274:
237:
216:
207:
203:
186:
173:
136:
113:
104:
100:
55:
52:
42:
35:
30:
26:
21:
2259:10.1186/1471-2407-12-294
2149:10.1186/1476-4598-13-180
2084:Cell Death & Disease
1220:Werdnig-Hoffmann disease
972:Chr 11: 69.45 – 69.47 Mb
391:tibialis anterior muscle
1771:Genes & Development
1668:Genes & Development
1560:Genes & Development
1469:10.1126/science.1165357
1216:spinal muscular atrophy
1210:Spinal muscular atrophy
1680:10.1101/gad.246546.114
1572:10.1101/gad.236869.113
1200:dyskeratosis congenita
1191:Dyskeratosis congenita
1175:
1109:
1101:
536:telomerase RNA binding
375:pancreatic ductal cell
363:epithelium of bronchus
2492:10.1093/carcin/bgn249
2096:10.1038/cddis.2010.90
1186:Clinical significance
1173:
1107:
1099:
965:Chr 17: 7.69 – 7.7 Mb
232:Chromosome 11 (mouse)
130:Chromosome 17 (human)
1408:10.4161/cc.8.15.9223
2199:2014PLoSO...991670R
1783:10.1101/gad.2006411
1461:2009Sci...323..644V
1042:that regulates the
383:ganglionic eminence
1959:10.1038/ng0797-265
1196:Germline mutations
1176:
1110:
1102:
778:ENSMUSG00000041346
606:Biological process
560:Cellular component
519:Molecular function
359:right uterine tube
94:WRAP53 - orthologs
2346:10.1002/ijc.22985
1867:10.1586/ehm.13.23
1132:remains elusive.
1021:
1020:
1017:
1016:
990:
989:
955:
954:
930:
929:
892:
891:
867:
866:
829:
828:
810:
809:
784:
783:
765:
764:
739:
738:
720:
719:
668:
667:
526:chaperone binding
503:
502:
499:
498:
477:
476:
464:
463:
402:
401:
300:
299:
199:
198:
2542:
2514:
2513:
2503:
2471:
2465:
2464:
2454:
2422:
2416:
2415:
2405:
2373:
2367:
2366:
2348:
2324:
2318:
2317:
2288:
2282:
2281:
2271:
2261:
2237:
2231:
2230:
2220:
2210:
2178:
2172:
2171:
2161:
2151:
2136:Molecular Cancer
2127:
2118:
2117:
2107:
2075:
2066:
2065:
2029:
2023:
2022:
1985:
1979:
1978:
1942:
1936:
1935:
1925:
1893:
1887:
1886:
1846:
1840:
1839:
1811:
1805:
1804:
1794:
1762:
1751:
1750:
1740:
1708:
1702:
1701:
1691:
1659:
1650:
1649:
1639:
1629:
1620:(11): e1000521.
1605:
1594:
1593:
1583:
1551:
1545:
1544:
1534:
1502:
1491:
1490:
1480:
1440:
1429:
1428:
1410:
1386:
1380:
1379:
1369:
1345:
1334:
1333:
1322:
1316:
1315:
1304:
1298:
1288:
1277:
1267:
1047:tumor suppressor
1003:
1002:
974:
967:
950:
934:
925:
901:
897:RefSeq (protein)
887:
871:
862:
838:
814:
795:
769:
750:
724:
705:
674:
509:
488:
473:
449:ventricular zone
413:
411:Top expressed in
406:
351:
349:Top expressed in
344:
323:
306:
296:
283:
272:
257:
250:
244:
233:
221:
205:
195:
182:
171:
156:
149:
143:
132:
118:
102:
96:
47:
40:
19:
2550:
2549:
2545:
2544:
2543:
2541:
2540:
2539:
2535:Cancer research
2520:
2519:
2518:
2517:
2473:
2472:
2468:
2431:Cancer Research
2424:
2423:
2419:
2375:
2374:
2370:
2326:
2325:
2321:
2290:
2289:
2285:
2239:
2238:
2234:
2180:
2179:
2175:
2129:
2128:
2121:
2077:
2076:
2069:
2046:10.1002/wrna.76
2031:
2030:
2026:
1987:
1986:
1982:
1947:Nature Genetics
1944:
1943:
1939:
1908:(6): 1389–403.
1895:
1894:
1890:
1848:
1847:
1843:
1813:
1812:
1808:
1764:
1763:
1754:
1710:
1709:
1705:
1674:(24): 2726–38.
1661:
1660:
1653:
1607:
1606:
1597:
1553:
1552:
1548:
1504:
1503:
1494:
1442:
1441:
1432:
1388:
1387:
1383:
1347:
1346:
1337:
1324:
1323:
1319:
1306:
1305:
1301:
1289:
1280:
1268:
1259:
1254:
1245:
1228:
1212:
1193:
1188:
1168:
1112:Transcripts of
1094:
1074:
1012:View/Edit Mouse
1007:View/Edit Human
970:
963:
960:Location (UCSC)
946:
942:
921:
917:
913:
909:
883:
879:
858:
854:
850:
846:
759:ENSG00000141499
652:
601:
555:
531:protein binding
469:
460:
455:
451:
447:
443:
439:
435:
431:
427:
423:
409:
398:
393:
389:
385:
381:
377:
373:
369:
365:
361:
347:
291:
278:
270:
260:
259:
258:
251:
231:
208:Gene location (
190:
177:
169:
159:
158:
157:
150:
128:
105:Gene location (
56:
43:
36:
17:
12:
11:
5:
2548:
2546:
2538:
2537:
2532:
2522:
2521:
2516:
2515:
2480:Carcinogenesis
2466:
2437:(6): 2349–57.
2417:
2368:
2339:(11): 2532–8.
2319:
2283:
2232:
2173:
2119:
2067:
2024:
1980:
1937:
1888:
1841:
1806:
1752:
1717:Molecular Cell
1703:
1651:
1595:
1546:
1511:Molecular Cell
1492:
1430:
1381:
1354:Molecular Cell
1335:
1317:
1299:
1278:
1256:
1255:
1253:
1250:
1244:
1241:
1227:
1224:
1211:
1208:
1192:
1189:
1187:
1184:
1167:
1164:
1093:
1090:
1073:
1070:
1034:implicated in
1019:
1018:
1015:
1014:
1009:
999:
998:
992:
991:
988:
987:
985:
983:
976:
975:
968:
961:
957:
956:
953:
952:
938:
937:
931:
928:
927:
905:
904:
898:
894:
893:
890:
889:
875:
874:
868:
865:
864:
842:
841:
835:
831:
830:
827:
826:
818:
817:
811:
808:
807:
799:
798:
792:
786:
785:
782:
781:
773:
772:
766:
763:
762:
754:
753:
747:
741:
740:
737:
736:
728:
727:
721:
718:
717:
709:
708:
702:
696:
695:
690:
685:
681:
680:
670:
669:
666:
665:
654:
653:
651:
650:
645:
640:
635:
630:
625:
620:
615:
609:
607:
603:
602:
600:
599:
594:
589:
584:
579:
574:
569:
563:
561:
557:
556:
554:
553:
548:
543:
538:
533:
528:
522:
520:
516:
515:
505:
504:
501:
500:
497:
496:
493:
492:
485:
479:
478:
475:
474:
466:
465:
462:
461:
459:
458:
454:
450:
446:
442:
438:
434:
430:
426:
422:
418:
415:
414:
403:
400:
399:
397:
396:
392:
388:
384:
380:
376:
372:
368:
364:
360:
356:
353:
352:
340:
339:
331:
320:
314:
313:
310:RNA expression
302:
301:
298:
297:
289:
285:
284:
276:
273:
268:
262:
261:
252:
245:
239:
235:
234:
229:
223:
222:
214:
213:
201:
200:
197:
196:
188:
184:
183:
175:
172:
167:
161:
160:
151:
144:
138:
134:
133:
126:
120:
119:
111:
110:
98:
97:
54:
50:
49:
41:
33:
32:
28:
27:
24:
23:
15:
13:
10:
9:
6:
4:
3:
2:
2547:
2536:
2533:
2531:
2528:
2527:
2525:
2511:
2507:
2502:
2497:
2493:
2489:
2485:
2481:
2477:
2470:
2467:
2462:
2458:
2453:
2448:
2444:
2440:
2436:
2432:
2428:
2421:
2418:
2413:
2409:
2404:
2399:
2395:
2391:
2388:(3): 2145–7.
2387:
2383:
2379:
2372:
2369:
2364:
2360:
2356:
2352:
2347:
2342:
2338:
2334:
2330:
2323:
2320:
2315:
2311:
2307:
2303:
2299:
2295:
2294:Oral Oncology
2287:
2284:
2279:
2275:
2270:
2265:
2260:
2255:
2251:
2247:
2243:
2236:
2233:
2228:
2224:
2219:
2214:
2209:
2204:
2200:
2196:
2193:(3): e91670.
2192:
2188:
2184:
2177:
2174:
2169:
2165:
2160:
2155:
2150:
2145:
2141:
2137:
2133:
2126:
2124:
2120:
2115:
2111:
2106:
2101:
2097:
2093:
2089:
2085:
2081:
2074:
2072:
2068:
2063:
2059:
2055:
2051:
2047:
2043:
2040:(4): 546–64.
2039:
2035:
2028:
2025:
2020:
2016:
2012:
2008:
2004:
2000:
1997:(5): 657–67.
1996:
1992:
1984:
1981:
1976:
1972:
1968:
1964:
1960:
1956:
1952:
1948:
1941:
1938:
1933:
1929:
1924:
1919:
1915:
1911:
1907:
1903:
1899:
1892:
1889:
1884:
1880:
1876:
1872:
1868:
1864:
1861:(3): 327–37.
1860:
1856:
1852:
1845:
1842:
1837:
1833:
1829:
1825:
1821:
1817:
1810:
1807:
1802:
1798:
1793:
1788:
1784:
1780:
1776:
1772:
1768:
1761:
1759:
1757:
1753:
1748:
1744:
1739:
1734:
1730:
1726:
1722:
1718:
1714:
1707:
1704:
1699:
1695:
1690:
1685:
1681:
1677:
1673:
1669:
1665:
1658:
1656:
1652:
1647:
1643:
1638:
1633:
1628:
1623:
1619:
1615:
1611:
1604:
1602:
1600:
1596:
1591:
1587:
1582:
1577:
1573:
1569:
1566:(7): 723–34.
1565:
1561:
1557:
1550:
1547:
1542:
1538:
1533:
1528:
1524:
1520:
1516:
1512:
1508:
1501:
1499:
1497:
1493:
1488:
1484:
1479:
1474:
1470:
1466:
1462:
1458:
1454:
1450:
1446:
1439:
1437:
1435:
1431:
1426:
1422:
1418:
1414:
1409:
1404:
1400:
1396:
1392:
1385:
1382:
1377:
1373:
1368:
1363:
1360:(4): 462–71.
1359:
1355:
1351:
1344:
1342:
1340:
1336:
1331:
1327:
1321:
1318:
1313:
1309:
1303:
1300:
1296:
1292:
1287:
1285:
1283:
1279:
1275:
1271:
1266:
1264:
1262:
1258:
1251:
1249:
1242:
1240:
1236:
1232:
1225:
1223:
1221:
1217:
1209:
1207:
1203:
1201:
1197:
1190:
1185:
1183:
1181:
1172:
1165:
1163:
1160:
1158:
1154:
1150:
1146:
1142:
1138:
1133:
1131:
1127:
1123:
1119:
1115:
1106:
1098:
1091:
1089:
1087:
1083:
1079:
1071:
1069:
1067:
1064:
1060:
1056:
1052:
1048:
1045:
1041:
1040:antisense RNA
1037:
1033:
1029:
1025:
1013:
1008:
1004:
1000:
997:
993:
986:
984:
981:
977:
973:
969:
966:
962:
958:
951:
949:
945:
939:
935:
932:
926:
924:
920:
916:
912:
906:
902:
899:
895:
888:
886:
882:
876:
872:
869:
863:
861:
857:
853:
849:
843:
839:
836:
834:RefSeq (mRNA)
832:
825:
824:
819:
815:
812:
806:
805:
800:
796:
793:
791:
787:
780:
779:
774:
770:
767:
761:
760:
755:
751:
748:
746:
742:
735:
734:
729:
725:
722:
716:
715:
710:
706:
703:
701:
697:
694:
691:
689:
686:
682:
679:
675:
671:
664:
660:
655:
649:
646:
644:
641:
639:
636:
634:
631:
629:
626:
624:
621:
619:
616:
614:
611:
610:
608:
605:
604:
598:
595:
593:
590:
588:
585:
583:
580:
578:
575:
573:
570:
568:
565:
564:
562:
559:
558:
552:
549:
547:
544:
542:
539:
537:
534:
532:
529:
527:
524:
523:
521:
518:
517:
514:
513:Gene ontology
510:
506:
494:
489:
486:
484:
480:
472:
467:
456:
452:
448:
444:
440:
436:
432:
428:
424:
420:
419:
416:
412:
407:
404:
394:
390:
386:
382:
378:
374:
370:
366:
362:
358:
357:
354:
350:
345:
342:
341:
338:
336:
332:
330:
329:
325:
324:
321:
319:
315:
311:
307:
303:
295:
290:
286:
282:
277:
271:11|11 B3
267:
263:
256:
249:
243:
236:
228:
224:
220:
215:
211:
206:
202:
194:
189:
185:
181:
176:
166:
162:
155:
148:
142:
135:
131:
125:
121:
117:
112:
108:
103:
99:
95:
91:
87:
83:
79:
75:
71:
67:
63:
59:
51:
46:
39:
34:
29:
25:
20:
2483:
2479:
2469:
2434:
2430:
2420:
2385:
2381:
2371:
2336:
2332:
2322:
2300:(1): 24–30.
2297:
2293:
2286:
2249:
2245:
2235:
2190:
2186:
2176:
2139:
2135:
2087:
2083:
2037:
2033:
2027:
1994:
1990:
1983:
1953:(3): 265–9.
1950:
1946:
1940:
1905:
1901:
1891:
1858:
1854:
1844:
1819:
1815:
1809:
1774:
1770:
1723:(1): 47–57.
1720:
1716:
1706:
1671:
1667:
1617:
1614:PLOS Biology
1613:
1563:
1559:
1549:
1517:(1): 47–57.
1514:
1510:
1452:
1448:
1398:
1394:
1384:
1357:
1353:
1329:
1320:
1311:
1302:
1246:
1237:
1233:
1229:
1213:
1204:
1194:
1180:WD40 repeats
1177:
1161:
1152:
1148:
1144:
1140:
1136:
1134:
1129:
1121:
1117:
1113:
1111:
1085:
1081:
1077:
1075:
1068:(Figure 1).
1066:Cajal bodies
1053:involved in
1043:
1027:
1023:
1022:
948:NP_001351698
941:
919:NP_001137464
915:NP_001137463
911:NP_001137462
908:
885:NM_001364769
878:
856:NM_001143992
852:NM_001143991
848:NM_001143990
845:
821:
802:
776:
757:
731:
712:
692:
687:
597:nuclear body
453:right kidney
425:spermatocyte
333:
326:
53:External IDs
2486:(1): 50–8.
2090:(1): e114.
1777:(1): 11–6.
1218:(type I or
582:nucleoplasm
541:RNA binding
441:granulocyte
387:granulocyte
292:69,471,081
279:69,452,584
31:Identifiers
2524:Categories
2246:BMC Cancer
1395:Cell Cycle
1297:, May 2017
1276:, May 2017
1252:References
1206:congenita
1063:organelles
567:Cajal body
421:blastocyst
337:(ortholog)
191:7,703,502
178:7,686,071
74:HomoloGene
1822:: 480–6.
1243:Discovery
944:NP_659073
923:NP_060551
881:NM_144824
860:NM_018081
678:Orthologs
587:cytoplasm
457:spermatid
82:GeneCards
2510:18978339
2461:19276375
2412:23192612
2363:25769110
2355:17683073
2314:25456005
2278:22805008
2227:24626331
2187:PLOS ONE
2168:25070141
2114:21368886
2062:19534375
2054:21957043
2019:15514134
2011:22302308
1975:12101961
1932:25467444
1883:26212504
1875:23782086
1836:22160078
1801:21205863
1747:19285445
1698:25512560
1646:21072240
1590:24696455
1541:19285445
1487:19179534
1425:34067208
1417:19571673
1376:19250907
1293:–
1272:–
1126:Hp53int1
1092:Function
1059:telomere
1057:repair,
996:Wikidata
657:Sources:
437:epiblast
371:testicle
2501:2639030
2452:2666150
2403:3563948
2269:3504514
2252:: 294.
2218:3953598
2195:Bibcode
2159:4118648
2142:: 180.
2105:3077286
1967:9207792
1923:4329143
1792:3012932
1738:2700737
1689:4265676
1637:2970535
1581:4015496
1532:2700737
1478:2728071
1457:Bibcode
1449:Science
1295:Ensembl
1274:Ensembl
1166:Domains
1157:scaRNAs
1143:(alias
1141:WRAP53β
1130:WRAP53γ
1051:protein
1030:) is a
790:UniProt
745:Ensembl
684:Species
663:QuickGO
592:cytosol
577:nucleus
312:pattern
170:17p13.1
70:2384933
38:Aliases
2508:
2498:
2459:
2449:
2410:
2400:
2361:
2353:
2312:
2276:
2266:
2225:
2215:
2166:
2156:
2112:
2102:
2060:
2052:
2017:
2009:
1973:
1965:
1930:
1920:
1881:
1873:
1834:
1799:
1789:
1745:
1735:
1696:
1686:
1644:
1634:
1588:
1578:
1539:
1529:
1485:
1475:
1423:
1415:
1374:
1226:Cancer
1145:WRAP53
1137:WRAP53
1114:WRAP53
1082:WRAP53
1078:WRAP53
1049:and a
1036:cancer
1024:WRAP53
982:search
980:PubMed
823:Q8VC51
804:Q9BUR4
733:216853
700:Entrez
483:BioGPS
445:morula
433:embryo
429:embryo
86:WRAP53
62:612661
45:WRAP53
22:WRAP53
2359:S2CID
2058:S2CID
2015:S2CID
1971:S2CID
1879:S2CID
1421:S2CID
1153:TCAB1
1149:WDR79
714:55135
693:Mouse
688:Human
659:Amigo
335:Mouse
328:Human
275:Start
210:Mouse
174:Start
107:Human
2506:PMID
2457:PMID
2408:PMID
2351:PMID
2310:PMID
2274:PMID
2223:PMID
2164:PMID
2110:PMID
2050:PMID
2007:PMID
1963:PMID
1928:PMID
1902:Cell
1871:PMID
1832:PMID
1820:2011
1797:PMID
1743:PMID
1694:PMID
1642:PMID
1586:PMID
1537:PMID
1483:PMID
1413:PMID
1372:PMID
1135:The
1076:The
1072:Gene
1032:gene
318:Bgee
266:Band
227:Chr.
165:Band
124:Chr.
78:9995
58:OMIM
2496:PMC
2488:doi
2447:PMC
2439:doi
2398:PMC
2390:doi
2341:doi
2337:121
2302:doi
2264:PMC
2254:doi
2213:PMC
2203:doi
2154:PMC
2144:doi
2100:PMC
2092:doi
2042:doi
1999:doi
1995:137
1955:doi
1918:PMC
1910:doi
1906:159
1863:doi
1824:doi
1787:PMC
1779:doi
1733:PMC
1725:doi
1684:PMC
1676:doi
1632:PMC
1622:doi
1576:PMC
1568:doi
1527:PMC
1519:doi
1473:PMC
1465:doi
1453:323
1403:doi
1362:doi
1151:or
1147:or
1122:p53
1118:p53
1086:p53
1055:DNA
1044:p53
1028:p53
491:n/a
288:End
187:End
90:OMA
66:MGI
2526::
2504:.
2494:.
2484:30
2482:.
2478:.
2455:.
2445:.
2435:69
2433:.
2429:.
2406:.
2396:.
2386:40
2384:.
2380:.
2357:.
2349:.
2335:.
2331:.
2308:.
2298:51
2296:.
2272:.
2262:.
2250:12
2248:.
2244:.
2221:.
2211:.
2201:.
2189:.
2185:.
2162:.
2152:.
2140:13
2138:.
2134:.
2122:^
2108:.
2098:.
2086:.
2082:.
2070:^
2056:.
2048:.
2036:.
2013:.
2005:.
1993:.
1969:.
1961:.
1951:16
1949:.
1926:.
1916:.
1904:.
1900:.
1877:.
1869:.
1857:.
1853:.
1830:.
1818:.
1795:.
1785:.
1775:25
1773:.
1769:.
1755:^
1741:.
1731:.
1721:34
1719:.
1715:.
1692:.
1682:.
1672:28
1670:.
1666:.
1654:^
1640:.
1630:.
1616:.
1612:.
1598:^
1584:.
1574:.
1564:28
1562:.
1558:.
1535:.
1525:.
1515:34
1513:.
1509:.
1495:^
1481:.
1471:.
1463:.
1451:.
1447:.
1433:^
1419:.
1411:.
1397:.
1393:.
1370:.
1358:33
1356:.
1352:.
1338:^
1328:.
1310:.
1281:^
1260:^
661:/
294:bp
281:bp
193:bp
180:bp
88:;
84::
80:;
76::
72:;
68::
64:;
60::
2512:.
2490::
2463:.
2441::
2414:.
2392::
2365:.
2343::
2316:.
2304::
2280:.
2256::
2229:.
2205::
2197::
2191:9
2170:.
2146::
2116:.
2094::
2088:2
2064:.
2044::
2038:2
2021:.
2001::
1977:.
1957::
1934:.
1912::
1885:.
1865::
1859:6
1838:.
1826::
1803:.
1781::
1749:.
1727::
1700:.
1678::
1648:.
1624::
1618:8
1592:.
1570::
1543:.
1521::
1489:.
1467::
1459::
1427:.
1405::
1399:8
1378:.
1364::
1332:.
1314:.
212:)
109:)
92::
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