77:
341:. Its effectiveness in transformation is thought to be somewhat epiphenomenal to its role in the typical lytic viral life cycle. The transformation capacity of MTag can be eliminated by mutations that remove the membrane anchor, and reduced or eliminated by mutations in the phosphorylated tyrosines and the proline-rich region.
250:, a function shared with the other tumor antigens; however, MTag preferentially performs other protein-protein interactions that compete with the Hsc70 interaction. MTag plays a role in viral DNA replication and in the transition from early to late gene expression, and its absence can cause defects in
265:
A (PP2A) through the same physical mechanism, interacting with the A subunit in a way that occludes binding of PP2A B subunits and thus inactivates the enzyme. This interaction is required for the formation of other MTag-host cell protein complexes; however, PP2A catalytic activity is not required.
305:
Studies of MTag have often concentrated on its role in cellular transformation more than its natural role in the life cycles of the polyomaviruses in which it occurs. One hypothesis for the evolutionary role of MTag relies on the observation that MPyV LTag lacks an apparent ability to bind the
787:; McBride, Alison A.; Camus, Alvin C.; McDermott, Alexa J.; Dill, Jennifer A.; Delwart, Eric; Ng, Terry F. F.; Farkas, Kata; Austin, Charlotte; Kraberger, Simona; Davison, William; Pastrana, Diana V.; Varsani, Arvind; Galloway, Denise A. (19 April 2016).
278:
residues in the MTag C-terminus. Preference for members of the Src family varies, with mouse and hamster polyomavirus MTags having different distributions. Once phosphorylated, MTag interacts with and activates downstream signaling pathways through
838:
Freund, Robert; Sotnikov, Alexander; Bronson, Roderick T.; Benjamin, Thomas L. (December 1992). "Polyoma virus middle T is essential for virus replication and persistence as well as for tumor induction in mice".
108:(LTag) are encoded in the "early region" of the polyomavirus genome, so named because this region of the genome is expressed early in the infectious process. (The "late region" contains genes encoding the viral
210:, although the sequence C-terminal to the J domain has little homology between the mouse and hamster viruses. Until recently, these were the only two polyomaviruses known to encode MTag, but in 2015 the
1311:
1306:
222:
lineage of the polyomavirus family. However, evidence of MTag encoding and expression has also recently been reported in at least one virus of unrelated lineage, the
1363:
734:
van der
Meijden, Els; Kazem, Siamaque; Dargel, Christina A.; van Vuren, Nick; Hensbergen, Paul J.; Feltkamp, Mariet C. W.; Imperiale, M. J. (15 September 2015).
261:
However, MTag's best-studied functions center on its interaction with host cell proteins to activate cellular signaling pathways. Like STag, MTag can bind
1891:
1886:
1881:
1876:
1838:
1833:
2213:
1871:
1866:
1342:
1337:
1332:
736:"Characterization of T Antigens, Including Middle T and Alternative T, Expressed by the Human Polyomavirus Associated with Trichodysplasia Spinulosa"
246:
MTag is essential for viral proliferation, though some of its functions overlap with those of STag. Through its J domain, MTag can bind and activate
1056:
112:.) In MTag-containing polyomaviruses, the early region contains at least three genes encoding STag, MTag, and LTag, and is transcribed as a single
874:
Courtneidge, Sara A.; Smith, Alan E. (2 June 1983). "Polyoma virus transforming protein associates with the product of the c-src cellular gene".
478:"Global Analysis of Mouse Polyomavirus Infection Reveals Dynamic Regulation of Viral and Host Gene Expression and Promiscuous Viral RNA Editing"
194:
MTag occurs only in a few known polyomaviruses, while STag and LTag both appear in all known members of the family. MTag is best studied in the
2594:
223:
2357:
124:, of which the first overlaps with the genes for STag and MTag. The result of this genetic encoding is three proteins that share a common
1169:
361:. Because most of that efficiency is due to MTag, the protein alone has also been extensively used to induce tumors in animal models.
783:
Buck, Christopher B.; Van
Doorslaer, Koenraad; Peretti, Alberto; Geoghegan, Eileen M.; Tisza, Michael J.; An, Ping; Katz, Joshua P.;
2327:
2261:
1488:
218:
polyomavirus was reported to contain MTag as well. This observation is consistent with expectations that it evolved uniquely in the
80:
A map of the murine polyomavirus genome, indicating the early genes (LTag, MTag, and STag) at right in blue and the late genes (the
1955:
1623:
992:"Induction of mammary tumors by expression of polyomavirus middle T oncogene: a transgenic mouse model for metastatic disease"
2733:
476:
Garren, Seth B.; Kondaveeti, Yuvabharath; Duff, Michael O.; Carmichael, Gordon G.; McBride, Alison Anne (25 September 2015).
2474:
477:
296:
1948:
1943:
1049:
2728:
2576:
2511:
2183:
2159:
1966:
1851:
1757:
1683:
1608:
1503:
1387:
1322:
1242:
1208:
1159:
159:
1252:
1179:
1144:
186:-rich sequence region where mutations disrupt MTag function, though the mechanism of this disruption is not known.
2134:
1938:
1139:
334:
288:
227:
65:
52:. MTag occurs only in a few known polyomaviruses, while STag and LTag are universal - it was first identified in
37:
2633:
2521:
2447:
2385:
2337:
2289:
2241:
2193:
2072:
1921:
1816:
1577:
1483:
1463:
1446:
1296:
1184:
1119:
299:
179:
1042:
271:
1286:
2696:
2532:
2417:
1257:
638:"Genome Sequences of a Rat Polyomavirus Related to Murine Polyomavirus, Rattus norvegicus Polyomavirus 1"
2666:
2557:
2437:
2309:
2169:
262:
175:
89:
295:. The signaling functions of the phosphorylated MTag have been described as behaving like a mimic of a
2468:
2406:
2093:
1537:
1450:
1397:
1218:
1109:
207:
117:
57:
2139:
1533:
1523:
1442:
140:
105:
101:
49:
45:
2661:
2538:
354:
195:
53:
2113:
1861:
1800:
1565:
1021:
972:
891:
856:
820:
765:
716:
667:
618:
558:
509:
458:
231:
226:, which is a normally asymptomatic infection in humans that sometimes causes the rare disease
133:
587:
Cheng, Jingwei; DeCaprio, James A.; Fluck, Michele M.; Schaffhausen, Brian S. (August 2009).
2375:
1981:
1718:
1713:
1708:
1703:
1698:
1693:
1672:
1011:
1003:
962:
952:
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448:
440:
385:
310:
235:
171:
76:
2605:
2062:
2052:
1805:
909:
Dilworth, Stephen M. (January 1995). "Polyoma virus middle T antigen: meddler or mimic?".
211:
167:
41:
529:"Small tumor antigen of polyomaviruses: role in viral life cycle and cell transformation"
321:. The function of MTag is thus hypothesized to indirectly replace this lost interaction.
2679:
2562:
2107:
1932:
1827:
1433:
1353:
815:
788:
760:
735:
662:
637:
613:
588:
553:
528:
504:
453:
428:
284:
182:
places it in association with membranes. Immediately prior to the membrane anchor is a
129:
109:
1016:
991:
922:
2722:
2628:
2279:
1740:
1377:
1094:
1069:
1065:
967:
940:
852:
711:
686:
373:
358:
255:
113:
85:
604:
2348:
2300:
2252:
2204:
2125:
1910:
1746:
1423:
1232:
1099:
1073:
957:
702:
338:
251:
81:
33:
1034:
166:. Particularly in the unique region of its C-terminus, MTag possesses a number of
805:
494:
2688:
2231:
1551:
1514:
784:
589:"Cellular transformation by Simian Virus 40 and Murine Polyoma Virus T antigens"
369:
330:
61:
2570:
2502:
2099:
1905:
1736:
1732:
362:
267:
148:
144:
125:
653:
2616:
2611:
2552:
2546:
1790:
1667:
1657:
1645:
1418:
1082:
366:
350:
307:
199:
163:
155:
976:
824:
769:
720:
671:
622:
562:
513:
462:
429:"Lessons in Signaling and Tumorigenesis from Polyomavirus Middle T Antigen"
1025:
895:
860:
444:
1198:
1007:
751:
317:, which interacts with the LTag proteins of other polyomaviruses such as
275:
44:
early in the infectious cycle along with two other related proteins, the
238:
encoding a protein called ALTO, may be evolutionarily related to MTag.
183:
25:
544:
2458:
2396:
2144:
2083:
1976:
1474:
1130:
887:
636:
Ehlers, B; Richter, D; Matuschka, FR; Ulrich, RG (3 September 2015).
219:
29:
274:
in a PP2A-dependent manner, and in turn is phosphorylated by Src on
198:, which was the first polyomavirus discovered and which is a potent
2029:
2019:
2014:
2004:
1999:
1994:
1989:
1596:
1591:
1586:
292:
247:
75:
84:
proteins) at left in red. Each region is transcribed as a single
2581:
2412:
2024:
2009:
1438:
1174:
318:
280:
137:
121:
93:
1038:
151:. The full-length MTag protein is around 420 amino acids long.
2642:
2638:
2621:
1767:
1618:
314:
234:
individuals. A somewhat more common tumor antigen variant, an
215:
56:(MPyV), the first polyomavirus discovered, and also occurs in
170:
sites. Unlike STag or LTag, the MTag C-terminus contains a
697:(2): 288–318, second and third pages, table of contents.
329:
MTag's most distinctive property is its efficiency as an
337:
in a variety of cell types and can immortalize cells in
162:
sites that mediate interactions with proteins in the
789:"The Ancient Evolutionary History of Polyomaviruses"
427:
Fluck, M. M.; Schaffhausen, B. S. (31 August 2009).
2687:
2678:
2655:
2592:
2519:
2510:
2501:
2457:
2445:
2436:
2395:
2383:
2374:
2347:
2335:
2326:
2299:
2287:
2278:
2251:
2239:
2230:
2203:
2191:
2182:
2157:
2124:
2082:
2070:
2061:
2046:
1964:
1919:
1904:
1849:
1814:
1799:
1784:
1755:
1731:
1681:
1666:
1653:
1644:
1606:
1575:
1564:
1513:
1501:
1473:
1461:
1432:
1414:
1385:
1376:
1351:
1320:
1294:
1285:
1274:
1240:
1231:
1206:
1197:
1157:
1129:
1117:
1108:
1090:
1081:
353:, particularly in newborn or immunodeficient mice,
990:Guy, C T; Cardiff, R D; Muller, W J (March 1992).
941:"Natural Biology of Polyomavirus Middle T Antigen"
939:Gottlieb, K. A.; Villarreal, L. P. (1 June 2001).
687:"Natural biology of polyomavirus middle T antigen"
357:has served as a productive mechanism for modeling
143:proteins. MTag and STag share an additional ~100
934:
932:
527:Khalili, K; Sariyer, IK; Safak, M (May 2008).
1050:
422:
420:
365:expressed MTag is used in the widely studied
206:conditions. MTag is also well known from the
8:
582:
580:
578:
576:
574:
572:
418:
416:
414:
412:
410:
408:
406:
404:
402:
400:
2684:
2675:
2516:
2507:
2454:
2442:
2392:
2380:
2344:
2332:
2296:
2284:
2248:
2236:
2200:
2188:
2079:
2067:
2058:
1916:
1811:
1796:
1752:
1678:
1663:
1650:
1572:
1510:
1470:
1458:
1429:
1382:
1291:
1282:
1237:
1203:
1126:
1114:
1105:
1087:
1057:
1043:
1035:
945:Microbiology and Molecular Biology Reviews
691:Microbiology and Molecular Biology Reviews
685:Gottlieb, KA; Villarreal, LP (June 2001).
433:Microbiology and Molecular Biology Reviews
120:. The LTag gene is usually encoded in two
2214:Parainfluenza hemagglutinin-neuraminidase
1015:
966:
956:
814:
804:
759:
710:
661:
612:
552:
503:
493:
452:
158:activity of its own, but has a number of
104:(STag), middle tumor antigen (MTag), and
396:
266:For example, MTag binds and activates
224:trichodysplasia spinulosa polyomavirus
2358:Respiratory syncytial virus G protein
349:Because of its high efficiency as an
7:
254:assembly. MTag is also required for
92:to express multiple proteins; the
14:
2262:Mumps hemagglutinin-neuraminidase
64:that can be sufficient to induce
24:and abbreviated MTag or MT) is a
1624:Hepatitis B virus DNA polymerase
60:. In MPyV, MTag is an efficient
1455:(non-enveloped circular ds-DNA)
605:10.1016/j.semcancer.2009.03.002
333:. It has the ability to induce
996:Molecular and Cellular Biology
958:10.1128/MMBR.65.2.288-318.2001
703:10.1128/mmbr.65.2.288-318.2001
533:Journal of Cellular Physiology
132:called the J domain which has
1:
2475:Vesiculovirus matrix proteins
923:10.1016/S0966-842X(00)88866-6
96:are shown as thickened lines.
853:10.1016/0042-6822(92)90247-M
806:10.1371/journal.ppat.1005574
495:10.1371/journal.ppat.1005166
2512:Structure and genome of HIV
1569:(circular partially ds-DNA)
160:protein-protein interaction
147:residues and have distinct
38:double-stranded DNA viruses
2750:
1145:Herpesvirus glycoprotein B
593:Seminars in Cancer Biology
1140:HHV capsid portal protein
335:neoplastic transformation
289:phosphoinositide 3-kinase
228:trichodysplasia spinulosa
66:neoplastic transformation
654:10.1128/genomeA.00997-15
300:receptor tyrosine kinase
272:protein tyrosine kinases
180:subcellular localization
172:membrane anchor sequence
72:Structure and expression
2135:Influenza hemagglutinin
325:Cellular transformation
154:Like STag, MTag has no
2697:Gag-onc fusion protein
911:Trends in Microbiology
190:Taxonomic distribution
178:region. The protein's
97:
2734:Polyomavirus proteins
2667:murine leukemia virus
2558:Reverse transcriptase
2438:Indiana vesiculovirus
2310:Measles hemagglutinin
445:10.1128/MMBR.00009-09
297:constitutively active
263:protein phosphatase 2
90:alternatively spliced
79:
1398:Early 35 kDa protein
1008:10.1128/MCB.12.3.954
752:10.1128/JVI.00911-15
642:Genome Announcements
208:hamster polyomavirus
174:that likely forms a
128:sequence, forming a
118:alternative splicing
58:hamster polyomavirus
18:middle tumor antigen
740:Journal of Virology
141:molecular chaperone
106:large tumor antigen
102:small tumor antigen
50:large tumor antigen
46:small tumor antigen
2729:Viral oncoproteins
2662:Rous sarcoma virus
355:mouse polyomavirus
196:mouse polyomavirus
100:The genes for the
98:
54:mouse polyomavirus
36:, which are small
2716:
2715:
2712:
2711:
2708:
2707:
2704:
2703:
2651:
2650:
2497:
2496:
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2492:
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2488:
2485:
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2432:
2431:
2428:
2427:
2370:
2369:
2366:
2365:
2322:
2321:
2318:
2317:
2274:
2273:
2270:
2269:
2226:
2225:
2222:
2221:
2178:
2177:
2153:
2152:
2042:
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2038:
2037:
1900:
1899:
1780:
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1635:
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1631:
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1546:
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1410:
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1406:
1405:
1372:
1371:
1270:
1269:
1266:
1265:
1227:
1226:
1193:
1192:
1153:
1152:
882:(5916): 435–439.
746:(18): 9427–9439.
545:10.1002/jcp.21326
293:phospholipase CÎł1
256:viral persistence
232:immunocompromised
134:sequence homology
20:(also called the
2741:
2685:
2676:
2517:
2508:
2455:
2443:
2393:
2381:
2376:Zaire ebolavirus
2345:
2333:
2297:
2285:
2249:
2237:
2201:
2189:
2080:
2068:
2059:
1982:3C-like protease
1917:
1812:
1797:
1753:
1679:
1673:Duplornaviricota
1664:
1651:
1573:
1511:
1471:
1459:
1430:
1383:
1292:
1283:
1238:
1204:
1127:
1115:
1106:
1088:
1059:
1052:
1045:
1036:
1030:
1029:
1019:
987:
981:
980:
970:
960:
936:
927:
926:
906:
900:
899:
888:10.1038/303435a0
871:
865:
864:
835:
829:
828:
818:
808:
780:
774:
773:
763:
731:
725:
724:
714:
682:
676:
675:
665:
648:(5): e00997-15.
633:
627:
626:
616:
584:
567:
566:
556:
524:
518:
517:
507:
497:
473:
467:
466:
456:
424:
386:List of proteins
311:tumor suppressor
236:overprinted gene
22:middle T-antigen
2749:
2748:
2744:
2743:
2742:
2740:
2739:
2738:
2719:
2718:
2717:
2700:
2647:
2606:transactivators
2588:
2481:
2424:
2362:
2314:
2266:
2218:
2174:
2149:
2120:
2063:Influenza virus
2053:Negarnaviricota
2050:
2048:
2034:
1960:
1908:
1896:
1845:
1806:Kitrinoviricota
1803:
1788:
1786:
1772:
1744:
1723:
1670:
1655:
1628:
1602:
1568:
1556:
1543:
1493:
1454:
1436:
1422:
1416:
1402:
1368:
1347:
1316:
1278:
1277:(Duplodnaviria,
1276:
1275:circular ds-DNA
1262:
1223:
1189:
1149:
1098:
1092:
1077:
1063:
1033:
989:
988:
984:
938:
937:
930:
908:
907:
903:
873:
872:
868:
837:
836:
832:
799:(4): e1005574.
785:Pipas, James M.
782:
781:
777:
733:
732:
728:
684:
683:
679:
635:
634:
630:
586:
585:
570:
526:
525:
521:
488:(9): e1005166.
475:
474:
470:
426:
425:
398:
394:
382:
347:
345:Use in research
327:
285:14-3-3 proteins
244:
212:genome sequence
192:
168:phosphorylation
110:capsid proteins
74:
68:in some cells.
28:encoded in the
12:
11:
5:
2747:
2745:
2737:
2736:
2731:
2721:
2720:
2714:
2713:
2710:
2709:
2706:
2705:
2702:
2701:
2699:
2694:
2692:
2682:
2680:Fusion protein
2673:
2672:
2671:
2670:
2669:
2664:
2653:
2652:
2649:
2648:
2646:
2645:
2636:
2631:
2626:
2625:
2624:
2619:
2614:
2601:
2599:
2590:
2589:
2587:
2586:
2585:
2584:
2579:
2567:
2566:
2565:
2563:HIV-1 protease
2560:
2555:
2543:
2542:
2541:
2528:
2526:
2514:
2505:
2499:
2498:
2495:
2494:
2491:
2490:
2487:
2486:
2483:
2482:
2480:
2479:
2478:
2477:
2469:matrix protein
2464:
2462:
2452:
2440:
2434:
2433:
2430:
2429:
2426:
2425:
2423:
2422:
2421:
2420:
2415:
2407:matrix protein
2402:
2400:
2390:
2378:
2372:
2371:
2368:
2367:
2364:
2363:
2361:
2360:
2354:
2352:
2342:
2330:
2324:
2323:
2320:
2319:
2316:
2315:
2313:
2312:
2306:
2304:
2294:
2282:
2276:
2275:
2272:
2271:
2268:
2267:
2265:
2264:
2258:
2256:
2246:
2234:
2228:
2227:
2224:
2223:
2220:
2219:
2217:
2216:
2210:
2208:
2198:
2186:
2180:
2179:
2176:
2175:
2173:
2172:
2166:
2164:
2155:
2154:
2151:
2150:
2148:
2147:
2142:
2137:
2131:
2129:
2122:
2121:
2119:
2118:
2117:
2116:
2108:viral envelope
2104:
2103:
2102:
2094:matrix protein
2089:
2087:
2077:
2065:
2056:
2049:negative-sense
2044:
2043:
2040:
2039:
2036:
2035:
2033:
2032:
2027:
2022:
2017:
2012:
2007:
2002:
1997:
1992:
1987:
1986:
1985:
1973:
1971:
1962:
1961:
1959:
1958:
1953:
1952:
1951:
1946:
1941:
1933:viral envelope
1928:
1926:
1914:
1902:
1901:
1898:
1897:
1895:
1894:
1889:
1884:
1879:
1874:
1869:
1864:
1858:
1856:
1847:
1846:
1844:
1843:
1842:
1841:
1836:
1828:viral envelope
1823:
1821:
1809:
1794:
1787:positive-sense
1782:
1781:
1778:
1777:
1774:
1773:
1771:
1770:
1764:
1762:
1750:
1729:
1728:
1725:
1724:
1722:
1721:
1716:
1711:
1706:
1701:
1696:
1690:
1688:
1676:
1661:
1648:
1642:
1641:
1638:
1637:
1634:
1633:
1630:
1629:
1627:
1626:
1621:
1615:
1613:
1604:
1603:
1601:
1600:
1599:
1589:
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1582:
1570:
1562:
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1557:
1555:
1554:
1548:
1545:
1544:
1542:
1541:
1531:
1526:
1520:
1518:
1508:
1499:
1498:
1495:
1494:
1492:
1491:
1486:
1480:
1478:
1468:
1456:
1434:Polyomaviridae
1427:
1412:
1411:
1408:
1407:
1404:
1403:
1401:
1400:
1394:
1392:
1380:
1374:
1373:
1370:
1369:
1367:
1366:
1360:
1358:
1349:
1348:
1346:
1345:
1340:
1335:
1329:
1327:
1318:
1317:
1315:
1314:
1309:
1303:
1301:
1289:
1280:
1279:Varidnaviria?)
1272:
1271:
1268:
1267:
1264:
1263:
1261:
1260:
1255:
1249:
1247:
1235:
1229:
1228:
1225:
1224:
1222:
1221:
1215:
1213:
1201:
1195:
1194:
1191:
1190:
1188:
1187:
1182:
1177:
1172:
1166:
1164:
1155:
1154:
1151:
1150:
1148:
1147:
1142:
1136:
1134:
1124:
1112:
1110:Herpes simplex
1103:
1085:
1079:
1078:
1066:Viral proteins
1064:
1062:
1061:
1054:
1047:
1039:
1032:
1031:
1002:(3): 954–961.
982:
951:(2): 288–318.
928:
901:
866:
847:(2): 716–723.
830:
793:PLOS Pathogens
775:
726:
677:
628:
599:(4): 218–228.
568:
519:
482:PLOS Pathogens
468:
439:(3): 542–563.
395:
393:
390:
389:
388:
381:
378:
363:Transgenically
346:
343:
326:
323:
243:
240:
202:under certain
191:
188:
130:protein domain
73:
70:
34:polyomaviruses
13:
10:
9:
6:
4:
3:
2:
2746:
2735:
2732:
2730:
2727:
2726:
2724:
2698:
2695:
2693:
2690:
2686:
2683:
2681:
2677:
2674:
2668:
2665:
2663:
2660:
2659:
2658:
2657:
2654:
2644:
2640:
2637:
2635:
2632:
2630:
2627:
2623:
2620:
2618:
2615:
2613:
2610:
2609:
2608:
2607:
2603:
2602:
2600:
2598:
2596:
2591:
2583:
2580:
2578:
2575:
2574:
2573:
2572:
2568:
2564:
2561:
2559:
2556:
2554:
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2550:
2549:
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2540:
2537:
2536:
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2534:
2530:
2529:
2527:
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2523:
2518:
2515:
2513:
2509:
2506:
2504:
2500:
2476:
2473:
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2471:
2470:
2466:
2465:
2463:
2460:
2456:
2453:
2451:
2449:
2444:
2441:
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2435:
2419:
2416:
2414:
2411:
2410:
2409:
2408:
2404:
2403:
2401:
2398:
2394:
2391:
2389:
2387:
2382:
2379:
2377:
2373:
2359:
2356:
2355:
2353:
2350:
2346:
2343:
2341:
2339:
2334:
2331:
2329:
2325:
2311:
2308:
2307:
2305:
2302:
2298:
2295:
2293:
2291:
2286:
2283:
2281:
2277:
2263:
2260:
2259:
2257:
2254:
2250:
2247:
2245:
2243:
2238:
2235:
2233:
2229:
2215:
2212:
2211:
2209:
2206:
2202:
2199:
2197:
2195:
2190:
2187:
2185:
2184:Parainfluenza
2181:
2171:
2168:
2167:
2165:
2163:
2161:
2156:
2146:
2143:
2141:
2140:Neuraminidase
2138:
2136:
2133:
2132:
2130:
2127:
2123:
2115:
2112:
2111:
2110:
2109:
2105:
2101:
2098:
2097:
2096:
2095:
2091:
2090:
2088:
2085:
2081:
2078:
2076:
2074:
2069:
2066:
2064:
2060:
2057:
2054:
2045:
2031:
2028:
2026:
2023:
2021:
2018:
2016:
2013:
2011:
2008:
2006:
2003:
2001:
1998:
1996:
1993:
1991:
1988:
1983:
1980:
1979:
1978:
1975:
1974:
1972:
1970:
1968:
1963:
1957:
1954:
1950:
1947:
1945:
1942:
1940:
1937:
1936:
1935:
1934:
1930:
1929:
1927:
1925:
1923:
1918:
1915:
1912:
1907:
1903:
1893:
1890:
1888:
1885:
1883:
1880:
1878:
1875:
1873:
1870:
1868:
1865:
1863:
1860:
1859:
1857:
1855:
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1848:
1840:
1837:
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1832:
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1830:
1829:
1825:
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1802:
1798:
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1783:
1769:
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1742:
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1712:
1710:
1707:
1705:
1702:
1700:
1697:
1695:
1692:
1691:
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1687:
1685:
1680:
1677:
1674:
1669:
1665:
1662:
1659:
1652:
1649:
1647:
1643:
1625:
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1620:
1617:
1616:
1614:
1612:
1610:
1605:
1598:
1595:
1594:
1593:
1590:
1588:
1585:
1583:
1581:
1579:
1574:
1571:
1567:
1563:
1553:
1550:
1549:
1539:
1535:
1532:
1530:
1527:
1525:
1522:
1521:
1519:
1516:
1512:
1509:
1507:
1505:
1500:
1490:
1487:
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1479:
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1469:
1467:
1465:
1460:
1457:
1452:
1448:
1444:
1440:
1435:
1431:
1428:
1425:
1420:
1413:
1399:
1396:
1395:
1393:
1391:
1389:
1384:
1381:
1379:
1378:Baculoviridae
1375:
1365:
1362:
1361:
1359:
1357:
1355:
1350:
1344:
1341:
1339:
1336:
1334:
1331:
1330:
1328:
1326:
1324:
1319:
1313:
1310:
1308:
1305:
1304:
1302:
1300:
1298:
1293:
1290:
1288:
1284:
1281:
1273:
1259:
1256:
1254:
1251:
1250:
1248:
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1244:
1239:
1236:
1234:
1230:
1220:
1217:
1216:
1214:
1212:
1210:
1205:
1202:
1200:
1196:
1186:
1183:
1181:
1178:
1176:
1173:
1171:
1168:
1167:
1165:
1163:
1161:
1156:
1146:
1143:
1141:
1138:
1137:
1135:
1132:
1128:
1125:
1123:
1121:
1116:
1113:
1111:
1107:
1104:
1101:
1096:
1095:Duplodnaviria
1091:linear ds-DNA
1089:
1086:
1084:
1080:
1075:
1071:
1067:
1060:
1055:
1053:
1048:
1046:
1041:
1040:
1037:
1027:
1023:
1018:
1013:
1009:
1005:
1001:
997:
993:
986:
983:
978:
974:
969:
964:
959:
954:
950:
946:
942:
935:
933:
929:
924:
920:
916:
912:
905:
902:
897:
893:
889:
885:
881:
877:
870:
867:
862:
858:
854:
850:
846:
842:
834:
831:
826:
822:
817:
812:
807:
802:
798:
794:
790:
786:
779:
776:
771:
767:
762:
757:
753:
749:
745:
741:
737:
730:
727:
722:
718:
713:
708:
704:
700:
696:
692:
688:
681:
678:
673:
669:
664:
659:
655:
651:
647:
643:
639:
632:
629:
624:
620:
615:
610:
606:
602:
598:
594:
590:
583:
581:
579:
577:
575:
573:
569:
564:
560:
555:
550:
546:
542:
539:(2): 309–19.
538:
534:
530:
523:
520:
515:
511:
506:
501:
496:
491:
487:
483:
479:
472:
469:
464:
460:
455:
450:
446:
442:
438:
434:
430:
423:
421:
419:
417:
415:
413:
411:
409:
407:
405:
403:
401:
397:
391:
387:
384:
383:
379:
377:
375:
374:breast cancer
371:
368:
364:
360:
359:tumorigenesis
356:
352:
344:
342:
340:
336:
332:
324:
322:
320:
316:
312:
309:
303:
301:
298:
294:
290:
286:
282:
277:
273:
269:
264:
259:
257:
253:
249:
241:
239:
237:
233:
229:
225:
221:
217:
213:
209:
205:
201:
197:
189:
187:
185:
181:
177:
176:transmembrane
173:
169:
165:
161:
157:
152:
150:
146:
142:
139:
135:
131:
127:
123:
119:
116:processed by
115:
114:messenger RNA
111:
107:
103:
95:
91:
87:
86:messenger RNA
83:
78:
71:
69:
67:
63:
59:
55:
51:
47:
43:
39:
35:
31:
27:
23:
19:
2604:
2593:
2569:
2545:
2531:
2520:
2467:
2446:
2405:
2384:
2349:glycoprotein
2336:
2301:glycoprotein
2288:
2253:glycoprotein
2240:
2205:glycoprotein
2192:
2158:
2126:glycoprotein
2106:
2092:
2071:
1965:
1956:Nucleocapsid
1931:
1920:
1911:Pisuviricota
1850:
1826:
1815:
1756:
1747:Pisuviricota
1682:
1607:
1576:
1528:
1502:
1462:
1424:Monodnaviria
1386:
1352:
1321:
1295:
1287:Epstein–Barr
1241:
1233:Adenoviridae
1207:
1158:
1118:
1100:Varidnaviria
999:
995:
985:
948:
944:
917:(1): 31–35.
914:
910:
904:
879:
875:
869:
844:
840:
833:
796:
792:
778:
743:
739:
729:
694:
690:
680:
645:
641:
631:
596:
592:
536:
532:
522:
485:
481:
471:
436:
432:
348:
328:
304:
260:
252:viral capsid
245:
203:
193:
153:
99:
82:viral capsid
21:
17:
15:
2689:oncoprotein
1801:Hepatitis C
1566:Hepatitis B
1552:Agnoprotein
1515:oncoprotein
1489:VP2 and VP3
370:mouse model
331:oncoprotein
62:oncoprotein
2723:Categories
2114:M2 protein
2100:M1 protein
1906:SARS-CoV-2
392:References
145:amino acid
126:N-terminal
40:. MTag is
2656:Multiple
2553:Integrase
1791:Riboviria
1668:Rotavirus
1658:Riboviria
1419:Riboviria
367:MMTV-PyMT
351:oncovirus
308:host cell
200:oncovirus
164:host cell
156:enzymatic
149:C-termini
42:expressed
1949:Membrane
1944:Envelope
1538:SV40 Tag
1199:Vaccinia
977:11381103
841:Virology
825:27093155
770:26136575
721:11381103
672:26337891
623:19505649
563:18022798
514:26407100
463:19721090
380:See also
313:protein
276:tyrosine
270:-family
242:Function
32:of some
2280:Measles
1733:Rhinov.
1180:ICP34.5
1026:1312220
896:6304524
861:1333120
816:4836724
761:4542345
663:4559740
614:2694755
554:2716072
505:4583464
454:2738132
339:culture
204:in vivo
184:proline
30:genomes
26:protein
2459:capsid
2397:capsid
2145:HA-tag
2084:capsid
2047:ss-RNA
1977:ORF1ab
1785:ss-RNA
1745:etc. (
1654:ds-RNA
1475:capsid
1343:EBNA-3
1338:EBNA-2
1333:EBNA-1
1131:capsid
1024:
1017:369527
1014:
975:
965:
894:
876:Nature
859:
823:
813:
768:
758:
719:
709:
670:
660:
621:
611:
561:
551:
512:
502:
461:
451:
291:, and
220:rodent
2595:VRAPs
2577:gp120
2232:Mumps
2030:ORF9b
2020:ORF7b
2015:ORF7a
2005:ORF3d
2000:ORF3c
1995:ORF3b
1990:ORF3a
1984:(NS5)
1939:Spike
1741:Hep A
1737:Polio
1597:HBeAg
1592:HBcAg
1587:HBsAg
1451:HaPyV
1447:MCPyV
1415:other
1354:ncRNA
1312:LMP-2
1307:LMP-1
1185:ICP47
1170:vmw65
1070:early
968:99028
712:99028
248:Hsc70
214:of a
122:exons
94:exons
2582:gp41
2522:VSPs
2448:VSPs
2418:VP24
2413:VP40
2386:VSPs
2338:VSPs
2290:VSPs
2242:VSPs
2194:VSPs
2160:VNPs
2073:VSPs
2025:ORF8
2010:ORF6
1967:VNPs
1922:VSPs
1892:NS5B
1887:NS5A
1882:NS4B
1877:NS4A
1852:VNPs
1817:VSPs
1758:VNPs
1719:NSP6
1714:NSP5
1709:NSP4
1704:NSP3
1699:NSP2
1694:NSP1
1684:VNPs
1609:VNPs
1578:VSPs
1534:LTag
1529:MTag
1524:STag
1504:VNPs
1464:VSPs
1443:MPyV
1439:SV40
1388:VNPs
1364:EBER
1323:VNPs
1297:VSPs
1243:VNPs
1219:B13R
1209:VNPs
1175:ICP8
1160:VNPs
1120:VSPs
1074:late
1072:and
1022:PMID
973:PMID
892:PMID
857:PMID
821:PMID
766:PMID
717:PMID
668:PMID
619:PMID
559:PMID
510:PMID
459:PMID
319:SV40
138:DnaJ
88:and
48:and
16:The
2643:Vpx
2641:or
2639:Vpu
2634:Vif
2629:Nef
2622:Vpr
2617:Rev
2612:Tat
2571:env
2547:pol
2539:p24
2533:gag
2328:RSV
2170:NS1
1872:NS3
1867:NS2
1768:VPg
1646:RNA
1619:HBx
1484:VP1
1258:E1B
1253:E1A
1083:DNA
1012:PMC
1004:doi
963:PMC
953:doi
919:doi
884:doi
880:303
849:doi
845:191
811:PMC
801:doi
756:PMC
748:doi
707:PMC
699:doi
658:PMC
650:doi
609:PMC
601:doi
549:PMC
541:doi
537:215
500:PMC
490:doi
449:PMC
441:doi
372:of
315:p53
281:Shc
268:Src
230:in
216:rat
136:to
2725::
2503:RT
1862:P7
1839:E2
1834:E1
1739:,
1735:,
1449:,
1445:,
1441:,
1020:.
1010:.
1000:12
998:.
994:.
971:.
961:.
949:65
947:.
943:.
931:^
913:.
890:.
878:.
855:.
843:.
819:.
809:.
797:12
795:.
791:.
764:.
754:.
744:89
742:.
738:.
715:.
705:.
695:65
693:.
689:.
666:.
656:.
644:.
640:.
617:.
607:.
597:19
595:.
591:.
571:^
557:.
547:.
535:.
531:.
508:.
498:.
486:11
484:.
480:.
457:.
447:.
437:73
435:.
431:.
399:^
376:.
302:.
287:,
283:,
258:.
2691::
2597::
2524::
2461::
2450::
2399::
2388::
2351::
2340::
2303::
2292::
2255::
2244::
2207::
2196::
2162::
2128::
2086::
2075::
2055:)
2051:(
1969::
1924::
1913:)
1909:(
1854::
1819::
1808:)
1804:(
1793:)
1789:(
1760::
1749:)
1743:,
1686::
1675:)
1671:(
1660:)
1656:(
1611::
1580::
1540:)
1536:(
1517::
1506::
1477::
1466::
1453:)
1437:(
1426:)
1421:,
1417:(
1390::
1356::
1325::
1299::
1245::
1211::
1162::
1133::
1122::
1102:)
1097:,
1093:(
1076:)
1068:(
1058:e
1051:t
1044:v
1028:.
1006::
979:.
955::
925:.
921::
915:3
898:.
886::
863:.
851::
827:.
803::
772:.
750::
723:.
701::
674:.
652::
646:3
625:.
603::
565:.
543::
516:.
492::
465:.
443::
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