422:
433:
466:, (ii) C-terminal sequence that ends exclusively with glutamine or glutamate across bacterial species, (iii) short size (similar to that of ubiquitin) and, (iv) high sequence conservation across tremendous evolutionary distance (a characteristic also in common with ubiquitin). The differences between UBact and Pup are their taxonomic distribution and amino acid sequences. While Pup is predominantly found in the
388:, while UBact ('Ubiquitin bacterial') is very limited in occurrence among the vast number of bacterial species. The terms 'Ubiquitin Bacterial' and 'Prokaryotic ubiquitin-like protein' suggest a molecular similarity between ubiquitin and UBact/Pup which is largely absent. While ubiquitin assumes a highly stable three-dimensional structure in solution, Pup has been shown to belong to the group of
349:
402:) contain both the Pup-proteasome system and a novel ORF-proteasome system that needed to be addressed and therefore was denoted UBact. The conjugation-proteasome components neighboring the UBact and Pup loci in these Nitrospirae bacteria show weak similarity and are probably not entirely redundant. Figure 2 illustrates the differences between the
37:
554:
From evolutionary perspective, the finding of the UBact-proteasome system in gram-negative bacteria suggests that either the Pup/UBact-proteasome systems evolved in bacteria prior to the split into gram positive and negative clades over 3000 million years ago or, that these systems were acquired by
550:
From the knowledge accumulated about the Pup-proteasome system and its importance in bacterial durability and disease causing ability, the homologous UBact-proteasome system is expected to have similar impact on the gram-negative bacteria where it is found. In addition to humans, animals such
395:
The establishment of the term UBact is controversial, since to date there is no experimental evidence presented to justify the distinction of UBact from Pup. The term UBact was denoted because several bacterial species from the phylum
Nitrospirae (where UBact was initially identified; e.g.,
541:
It took almost ten years since the discovery of Pup in 2008, to identify UBact. This is probably due to the difference between Pup and UBact amino acids sequences, and because very few bacteria from the five phyla where UBact is found have been sequenced.
294:(for Ubiquitin Bacterial), although the distinction has not been proven to be phylogenetically supported by a separate evolutionary origin and is without experimental evidence. UBact is a homolog of Pup, and is found in several phyla of
850:
Lehmann G, Udasin RG, Livneh I, Ciechanover A (February 2017). "Identification of UBact, a ubiquitin-like protein, along with other homologous components of a conjugation system and the proteasome in different gram-negative bacteria".
545:
Bacteria from the phyla where UBact is found interact with humans, and are found in the human gut microbiota. In marine systems, the most frequently encountered nitrogen-oxidizing bacteria are related to the UBact encoding
1181:
Dubourg G, Lagier JC, Armougom F, Robert C, Audoly G, Papazian L, Raoult D (February 2013). "High-level colonisation of the human gut by
Verrucomicrobia following broad-spectrum antibiotic treatment".
1447:
559:(s) from a third, yet unknown, organism. In support of the second possibility, two UBact loci were found in the genome of an uncultured anaerobic methanotrophic Archaeon (ANME-1;locus
442:
In spite of the large difference in sequence, UBact is homologous to Pup and shares several characteristics with it: (i) same genomic location within a cluster of genes homologous to
1700:
213:
is different, owing to their distinct evolutionary origins. In contrast to the three-step reaction of ubiquitylation, pupylation requires only two steps, and thus only two
116:
240:. It is then recognized by the protein Mycobacterium proteasomal ATPase (Mpa), in a mechanism that induces folding of Pup. Mpa delivers the substrate protein to the
551:
livestock and fish that eat from the ground or swim in water are expected to be constantly exposed to UBact containing bacteria in the soil and water respectively.
421:
359:. Given the similarity to Pup, the prediction of a structure is doubtful, as Pup is proven to be instrinsically disordered in solution (see text for references).
1440:
1433:
1018:
Liao S, Shang Q, Zhang X, Zhang J, Xu C, Tu X (August 2009). "Pup, a prokaryotic ubiquitin-like protein, is an intrinsically disordered protein".
2045:
366:(UBact) is a protein that is homologous to Prokaryotic ubiquitin-like protein (Pup). UBact was recently described by the group of Professor
730:"Binding-induced folding of prokaryotic ubiquitin-like protein on the Mycobacterium proteasomal ATPase targets substrates for degradation"
426:
Figure 2. Alignment of UBact (EES53751) and Pup (EES52728) from the bacterium
Leptospirillum ferrodiazotrophum to assess their similarity
389:
886:
Delley CL, Müller AU, Ziemski M, Weber-Ban E (November 2017). "Prokaryotic
Ubiquitin-Like Protein and Its Ligase/Deligase Enyzmes
270:
1456:
1656:
274:
538:, UBact does not contain a di-glycine motif at its C-terminus. Rather, it usually ends with the sequence RG (see figure 3).
136:
432:
418:
revealed its existence and extreme conservation across several gram-negative bacterial phyla, as illustrated in figure 3.
1869:
1083:"Numerous uncharacterized and highly divergent microbes which colonize humans are revealed by circulating cell-free DNA"
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1814:
398:
181:
1218:"The Genome of Nitrospina gracilis Illuminates the Metabolism and Evolution of the Major Marine Nitrite Oxidizer"
1754:
556:
124:
2079:
36:
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1134:"Planctomycetes DNA in febrile aplastic patients with leukemia, rash, diarrhea, and micronodular pneumonia"
2120:
1501:
511:
120:
531:
1329:
681:
587:
222:
934:
Vijay-Kumar S, Bugg CE, Cook WJ (April 1987). "Structure of ubiquitin refined at 1.8 A resolution".
515:
1464:
1081:
Kowarsky M, Camunas-Soler J, Kertesz M, De
Vlaminck I, Koh W, Pan W, et al. (September 2017).
527:
519:
291:
245:
194:
507:
2110:
1478:
503:
437:
Figure 3. Demonstration of UBact C-terminus conservation across tremendous evolutionary distance.
1132:
Drancourt M, Prebet T, Aghnatios R, Edouard S, Cayrou C, Henry M, et al. (September 2014).
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1000:
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916:
868:
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367:
355:. Phyre2 server was used to predict the structure of the UBact Pup homolog from the bacterium
277:
determined the X-ray crystal structure of the complex between Pup and its delivery enzyme Mpa
143:
111:
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1999:
1994:
1969:
1959:
1954:
1944:
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1337:
1318:"Posttranslational regulation of coordinated enzyme activities in the Pup-proteasome system"
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607:
263:
2015:
1949:
103:
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1469:
571:
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233:
670:"Ubiquitin-Like Protein Involved in the Proteasome Pathway of Mycobacterium tuberculosis"
1333:
685:
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1244:
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1158:
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303:
283: and found that Pup binding to Mpa induces the folding of a unique alpha-helix.
206:
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299:
295:
287:
41:
Three
Prokaryotic ubiquitin-like proteins (blue) bound to proteasomal ATPase Mpa (red
1068:"Ubiquitin-like protein UBact [Leptospirillum ferriphilum] - Protein - NCBI"
65:
2149:
2131:
969:
Chen X, Solomon WC, Kang Y, Cerda-Maira F, Darwin KH, Walters KJ (September 2009).
798:"Pup, a prokaryotic ubiquitin-like protein, is an intrinsically disordered protein"
665:
636:
523:
491:
479:
286:
In 2017, the presence of Pup homologs in bacterial species outside of the group of
99:
17:
1054:"Ubiquitin-like protein Pup [Leptospirillum ferriphilum] - Protein - NCBI"
1425:
1067:
1053:
564:
560:
340:
77:
2225:
2093:
641:
1377:"The Timetree of Prokaryotes: New Insights into Their Evolution and Speciation"
1322:
Proceedings of the
National Academy of Sciences of the United States of America
1087:
Proceedings of the
National Academy of Sciences of the United States of America
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348:
255:, bacteria may use a small-protein modifier to control protein stability.
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335:
72:
1834:
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Elharar Y, Roth Z, Hecht N, Rotkopf R, Khalaila I, Gur E (March 2016).
1031:
813:
782:
259:
186:
745:
534:". With regard to the amino acid sequence, in difference from Pup and
279:
155:
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2010:
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971:"Prokaryotic ubiquitin-like protein pup is intrinsically disordered"
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Update: UBact is also found in the gram-negative bacterial phylum
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Marin J, Battistuzzi FU, Brown AC, Hedges SB (February 2017).
1269:"Bacterial Proteasomes: Mechanistic and Functional Insights"
664:
Pearce, M. J.; Mintseris, J.; Ferreyra, J.; Gygi, S. P.;
1422:, a database of pupylated proteins and pupylation sites.
1216:
Lücker S, Nowka B, Rattei T, Spieck E, Daims H (2013).
614:), and in the phylum Candidatus Bipolaricaulota (e.g.,
414:. Further analyses of the UBact (and not Pup) locus in
796:
Liao S, Shang Q, Zhang X, Zhang J, Xu C, Tu X (2009).
498:. UBact was also identified in the genomes of several
290:
bacteria was reported. The Pup homologs were termed
217:
are involved in pupylation. The enzymes involved in
2199:
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1802:
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59:
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29:
228:Similar to ubiquitin, Pup is attached to specific
410:loci in the representative Nitrospirae bacterium
781:for "Prokaryotic ubiquitin-like protein Pup" at
1441:
1183:International Journal of Antimicrobial Agents
845:
843:
841:
839:
837:
835:
833:
831:
8:
777:Universal protein resource accession number
384:was named for its ubiquitous presence among
353:Figure 1. A diagram of the UBact Pup homolog
298:bacteria (Pup is found predominantly in the
659:
657:
2127:
1474:
1448:
1434:
1426:
1273:Microbiology and Molecular Biology Reviews
610:" (previously candidate phylum FBP; e.g.,
84:
1392:
1351:
1341:
1292:
1243:
1233:
1157:
1108:
1098:
994:
910:
753:
734:Nature Structural & Molecular Biology
728:Wang T, Darwin KH, Li H (November 2010).
701:
251:The discovery of Pup indicates that like
723:
721:
478:bacteria from the following five phyla:
653:
185:. Like ubiquitin, Pup serves to direct
313:
26:
555:different bacterial lineages through
258:The Pup gene encodes a 64–amino acid
7:
502:bacteria, and specifically from the
1267:Becker SH, Darwin KH (March 2017).
262:with a molecular size of about 6.9
2264:Prokaryotic ubiquitin-like protein
590:" (previously known as WS3; e.g.,
317:Prokaryotic ubiquitin-like protein
232:residues of substrate proteins by
170:Prokaryotic ubiquitin-like protein
25:
1195:10.1016/j.ijantimicag.2012.10.012
390:intrinsically disordered proteins
1138:Journal of Clinical Microbiology
416:Leptospirillum ferrodiazotrophum
412:Leptospirillum ferrodiazotrophum
271:intrinsically disordered protein
172:(Pup) is a functional analog of
35:
1381:Molecular Biology and Evolution
474:, UBact was identified only in
248:by coupling of ATP hydrolysis.
1794:Mitochondrial targeting signal
1457:Posttranslational modification
275:Brookhaven National Laboratory
205:). However, the enzymology of
1:
567:). More possibilities exist.
273:. In 2010, scientists at the
88:Available protein structures:
1870:Ubiquitin-conjugating enzyme
975:Journal of Molecular Biology
948:10.1016/0022-2836(87)90679-6
936:Journal of Molecular Biology
891:Journal of Molecular Biology
357:Methylacidiphilum infernorum
2158:E2 SUMO-conjugating enzyme
1815:Ubiquitin-activating enzyme
586:) in the candidate phylum "
2305:
2141:E1 SUMO-activating enzyme
865:10.1016/j.bbrc.2017.01.037
399:Leptospirillum ferriphilum
182:Mycobacterium tuberculosis
1222:Frontiers in Microbiology
987:10.1016/j.jmb.2009.07.018
903:10.1016/j.jmb.2017.04.020
83:
34:
1755:Survival of motor neuron
1235:10.3389/fmicb.2013.00027
557:horizontal gene transfer
2121:Ubiquitin-like proteins
2080:Deubiquitinating enzyme
1343:10.1073/pnas.1525185113
1100:10.1073/pnas.1707009114
1020:The Biochemical Journal
694:10.1126/science.1163885
30:Pup-like protein family
439:
429:
360:
1394:10.1093/molbev/msw245
1285:10.1128/MMBR.00036-16
435:
424:
351:
199:Pup-proteasome system
1150:10.1128/JCM.01207-14
223:glutamine synthetase
1479:Heat shock proteins
1334:2016PNAS..113E1605E
912:20.500.11850/191988
802:Biochemical Journal
686:2008Sci...322.1104P
680:(5904): 1104–1107.
548:Nitrospina gracilis
504:candidate divisions
364:Ubiquitin Bacterial
310:Ubiquitin bacterial
221:are descended from
18:Ubiquitin bacterial
2289:Bacterial proteins
1032:10.1042/BJ20090738
814:10.1042/BJ20090738
606:), in the phylum "
512:Handelsmanbacteria
440:
430:
361:
2276:
2275:
2272:
2271:
1781:Protein targeting
1775:
1774:
1093:(36): 9623–9628.
897:(22): 3486–3499.
746:10.1038/nsmb.1918
484:Verrucomicrobiota
368:Aaron Ciechanover
346:
345:
302:bacterial phylum
236:; this is called
167:
166:
163:
162:
137:structure summary
16:(Redirected from
2296:
2128:
2041:Ubiquitin ligase
1807:(ubiquitylation)
1745:Alpha crystallin
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1436:
1427:
1407:
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1372:
1366:
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1355:
1345:
1328:(12): E1605-14.
1313:
1307:
1306:
1296:
1279:(1): e00036–16.
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532:Yanofskybacteria
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2170:E3 SUMO ligase
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1470:protein folding
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588:Latescibacteria
572:Gemmatimonadota
496:Planctomycetota
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1414:External links
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1387:(2): 437–446.
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859:(3): 946–950.
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740:(11): 1352–7.
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516:Fraserbacteria
488:Armatimonadota
472:Actinomycetota
344:
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304:Actinomycetota
207:ubiquitylation
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2069:
2067:
2064:
2062:
2059:
2057:
2054:
2052:
2049:
2047:
2044:
2043:
2042:
2038:
2037:
2032:
2029:
2027:
2024:
2022:
2019:
2017:
2014:
2012:
2009:
2006:
2003:
2001:
1998:
1996:
1993:
1991:
1988:
1986:
1983:
1981:
1978:
1976:
1973:
1971:
1968:
1966:
1963:
1961:
1958:
1956:
1953:
1951:
1948:
1946:
1943:
1941:
1938:
1936:
1933:
1931:
1928:
1926:
1923:
1921:
1918:
1916:
1913:
1911:
1908:
1906:
1903:
1901:
1898:
1896:
1893:
1891:
1888:
1886:
1883:
1881:
1878:
1876:
1873:
1872:
1871:
1867:
1866:
1861:
1858:
1856:
1853:
1851:
1848:
1846:
1843:
1841:
1838:
1836:
1833:
1831:
1828:
1826:
1823:
1821:
1818:
1817:
1816:
1812:
1811:
1809:
1805:
1801:
1795:
1792:
1790:
1787:
1786:
1784:
1782:
1778:
1766:
1763:
1761:
1758:
1757:
1756:
1753:
1751:
1748:
1746:
1743:
1742:
1740:
1736:
1728:
1725:
1723:
1720:
1718:
1715:
1713:
1707:
1705:
1699:
1698:
1697:
1694:
1693:
1688:
1685:
1683:
1680:
1678:
1675:
1673:
1670:
1668:
1665:
1663:
1660:
1658:
1655:
1653:
1650:
1648:
1645:
1643:
1640:
1638:
1635:
1633:
1630:
1628:
1625:
1624:
1623:
1620:
1619:
1614:
1611:
1609:
1606:
1604:
1601:
1599:
1596:
1594:
1591:
1589:
1586:
1584:
1581:
1579:
1576:
1574:
1571:
1569:
1566:
1564:
1561:
1559:
1556:
1554:
1551:
1549:
1546:
1544:
1541:
1539:
1536:
1534:
1531:
1529:
1526:
1524:
1521:
1520:
1519:
1516:
1515:
1512:
1508:
1505:
1503:
1500:
1498:
1495:
1493:
1490:
1489:
1487:
1485:
1480:
1476:
1473:
1471:
1466:
1462:
1458:
1451:
1446:
1444:
1439:
1437:
1432:
1431:
1428:
1421:
1418:
1417:
1413:
1404:
1400:
1395:
1390:
1386:
1382:
1378:
1371:
1368:
1363:
1359:
1354:
1349:
1344:
1339:
1335:
1331:
1327:
1323:
1319:
1312:
1309:
1304:
1300:
1295:
1290:
1286:
1282:
1278:
1274:
1270:
1263:
1260:
1255:
1251:
1246:
1241:
1236:
1231:
1227:
1223:
1219:
1212:
1209:
1204:
1200:
1196:
1192:
1189:(2): 149–55.
1188:
1184:
1177:
1174:
1169:
1165:
1160:
1155:
1151:
1147:
1144:(9): 3453–5.
1143:
1139:
1135:
1128:
1125:
1120:
1116:
1111:
1106:
1101:
1096:
1092:
1088:
1084:
1077:
1074:
1069:
1063:
1060:
1055:
1049:
1046:
1041:
1037:
1033:
1029:
1026:(2): 207–15.
1025:
1021:
1014:
1011:
1006:
1002:
997:
992:
988:
984:
981:(1): 208–17.
980:
976:
972:
965:
962:
957:
953:
949:
945:
942:(3): 531–44.
941:
937:
930:
927:
922:
918:
913:
908:
904:
900:
896:
892:
888:
882:
879:
874:
870:
866:
862:
858:
854:
846:
844:
842:
840:
838:
836:
834:
832:
828:
823:
819:
815:
811:
807:
803:
799:
792:
789:
784:
780:
773:
770:
765:
761:
756:
751:
747:
743:
739:
735:
731:
724:
722:
718:
713:
709:
704:
699:
695:
691:
687:
683:
679:
675:
671:
667:
666:Darwin, K. H.
660:
658:
654:
647:
643:
640:
638:
635:
633:
630:
629:
625:
623:
621:
617:
613:
609:
605:
601:
597:
593:
589:
585:
581:
577:
573:
568:
566:
562:
558:
552:
549:
543:
539:
537:
533:
529:
528:Parcubacteria
525:
521:
520:Terrybacteria
517:
513:
509:
505:
501:
497:
493:
489:
485:
481:
477:
476:gram-negative
473:
469:
468:gram-positive
465:
461:
457:
453:
449:
445:
438:
434:
427:
423:
419:
417:
413:
409:
405:
401:
400:
393:
391:
387:
383:
379:
377:
373:
369:
365:
358:
354:
350:
342:
339:
337:
333:
329:
325:
320:
315:
309:
307:
305:
301:
300:gram-positive
297:
296:gram-negative
293:
289:
288:gram-positive
284:
281:
276:
272:
267:
265:
261:
256:
254:
249:
247:
243:
239:
235:
231:
226:
224:
220:
216:
212:
208:
204:
200:
196:
192:
188:
184:
183:
179:
176:found in the
175:
171:
157:
151:
147:
145:
141:
138:
135:
133:
129:
126:
122:
118:
115:
113:
109:
105:
101:
98:
95:
91:
86:
82:
79:
76:
74:
70:
67:
64:
62:
58:
54:
50:
45:
38:
33:
28:
19:
2263:
2132:SUMO protein
1384:
1380:
1370:
1325:
1321:
1311:
1276:
1272:
1262:
1225:
1221:
1211:
1186:
1182:
1176:
1141:
1137:
1127:
1090:
1086:
1076:
1062:
1048:
1023:
1019:
1013:
978:
974:
964:
939:
935:
929:
894:
890:
887:
881:
856:
852:
805:
801:
791:
772:
737:
733:
677:
673:
637:SUMO protein
569:
553:
547:
544:
540:
524:Poribacteria
508:Acetothermia
492:Nitrospinota
480:Nitrospirota
463:
459:
455:
451:
447:
443:
441:
436:
425:
415:
411:
407:
403:
397:
394:
380:
363:
362:
356:
352:
285:
268:
257:
250:
237:
227:
202:
198:
180:
169:
168:
2226:neddylation
1492:Hsp10/GroES
1484:Chaperonins
642:Neddylation
322:Identifiers
246:degradation
195:degradation
47:Identifiers
1518:Hsp40/DnaJ
1465:Chaperones
1228:(27): 27.
648:References
612:A0A2S8SU03
604:A0A3B8MMW3
600:A0A3D4H075
596:A0A3D5FTR6
592:A0A3D2RHP4
584:A0A2E7JSE3
580:A0A2E3J6F7
576:A0A2E8WA32
565:CBH39258.1
563:and locus
561:CBH38808.1
500:candidatus
386:eukaryotes
269:Pup is an
253:eukaryotes
242:proteasome
238:pupylation
219:pupylation
211:pupylation
191:proteasome
178:prokaryote
100:structures
1804:Ubiquitin
1750:Clusterin
632:Ubiquitin
536:Ubiquitin
452:Pup/UBact
382:Ubiquitin
341:IPR037543
174:ubiquitin
153:,
78:IPR008515
2283:Category
2084:Ataxin 3
1403:27965376
1362:26951665
1303:27974513
1254:23439773
1203:23294932
1168:24920769
1119:28830999
1040:19580545
1005:19607839
921:28478282
873:28087277
822:19580545
764:20953180
712:18832610
668:(2008).
626:See also
530:", and "
372:Technion
336:InterPro
187:proteins
117:RCSB PDB
73:InterPro
2007:(CDC34)
1353:4812726
1330:Bibcode
1294:5312241
1245:3578206
1159:4313204
1110:5594678
996:2734869
956:3041007
783:UniProt
755:2988878
703:2698935
682:Bibcode
674:Science
574:(e.g.,
470:phylum
370:at the
260:protein
215:enzymes
197:in the
189:to the
159:
66:PF05639
2051:Cullin
1401:
1360:
1350:
1301:
1291:
1252:
1242:
1201:
1166:
1156:
1117:
1107:
1038:
1003:
993:
954:
919:
871:
820:
779:P9WHN4
762:
752:
710:
700:
620:H5SQ95
616:H5SEU7
602:, and
494:, and
462:->
458:->
454:->
450:->
446:->
376:Israel
327:Symbol
230:lysine
132:PDBsum
106:
96:
52:Symbol
2242:ATG12
2232:FAT10
2222:NEDD8
2207:ISG15
2200:Other
2189:PIAS4
2184:PIAS3
2179:PIAS2
2174:PIAS1
2124:(UBL)
2106:BIRC6
2066:FANCL
1738:Other
1727:TRAP1
1696:Hsp90
1622:Hsp70
1511:GroEL
1507:HSP60
1502:Hsp47
1497:Hsp27
1420:PupDB
404:UBact
330:UBact
292:UBact
2257:UBL5
2247:FUB1
2237:ATG8
2217:UFM1
2212:URM1
2162:UBC9
2150:SAE2
2145:SAE1
2111:UFC1
2101:ATG3
2094:CYLD
2089:USP6
2071:UBR1
2061:MDM2
1860:SAE1
1855:NAE1
1850:ATG7
1845:UBA7
1840:UBA6
1835:UBA5
1830:UBA3
1825:UBA2
1820:UBA1
1765:SMN2
1760:SMN1
1399:PMID
1358:PMID
1299:PMID
1250:PMID
1199:PMID
1164:PMID
1115:PMID
1036:PMID
1001:PMID
952:PMID
917:PMID
869:PMID
818:PMID
760:PMID
708:PMID
618:and
526:", "
522:", "
518:", "
514:", "
510:", "
464:PafA
460:PrcA
456:PrcB
406:and
280:3M9D
244:for
209:and
193:for
156:3M9D
150:3M91
125:PDBj
121:PDBe
104:ECOD
94:Pfam
61:Pfam
2252:MUB
2056:CBL
2046:VHL
2039:E3
1868:E2
1813:E1
1682:12A
1613:C19
1608:C14
1603:C13
1598:C11
1593:C10
1548:B11
1389:doi
1348:PMC
1338:doi
1326:113
1289:PMC
1281:doi
1240:PMC
1230:doi
1191:doi
1154:PMC
1146:doi
1105:PMC
1095:doi
1091:114
1028:doi
1024:422
991:PMC
983:doi
979:392
944:doi
940:194
907:hdl
899:doi
895:429
889:".
861:doi
857:483
810:doi
806:422
750:PMC
742:doi
698:PMC
690:doi
678:322
622:).
448:Dop
444:Mpa
408:Pup
306:).
264:kDa
203:PPS
144:PDB
112:PDB
55:Pup
2285::
2082::
2026:V2
2021:V1
2011:R2
2005:R1
2000:Q2
1995:Q1
1975:L6
1970:L4
1965:L3
1960:L2
1955:L1
1945:J2
1940:J1
1925:G2
1920:G1
1915:E3
1910:E2
1905:E1
1900:D3
1895:D2
1890:D1
1722:ER
1687:14
1652:4L
1637:1L
1632:1B
1627:1A
1588:C7
1583:C6
1578:C5
1573:C3
1568:C1
1563:B9
1558:B6
1553:B4
1543:B2
1538:B1
1533:A3
1528:A2
1523:A1
1397:.
1385:34
1383:.
1379:.
1356:.
1346:.
1336:.
1324:.
1320:.
1297:.
1287:.
1277:81
1275:.
1271:.
1248:.
1238:.
1224:.
1220:.
1197:.
1187:41
1185:.
1162:.
1152:.
1142:52
1140:.
1136:.
1113:.
1103:.
1089:.
1085:.
1034:.
1022:.
999:.
989:.
977:.
973:.
950:.
938:.
915:.
905:.
893:.
867:.
855:.
830:^
816:.
804:.
800:.
758:.
748:.
738:17
736:.
732:.
720:^
706:.
696:.
688:.
676:.
672:.
656:^
598:,
594:,
582:,
578:,
490:,
486:,
482:,
392:.
378:.
374:,
266:.
225:.
123:;
119:;
102:/
2228:)
2224:(
2031:Z
2016:S
1990:O
1985:N
1980:M
1950:K
1935:I
1930:H
1885:C
1880:B
1875:A
1717:β
1711:2
1709:α
1703:1
1701:α
1677:9
1672:8
1667:7
1662:6
1657:5
1647:4
1642:2
1509:/
1481:/
1467:/
1449:e
1442:t
1435:v
1405:.
1391::
1364:.
1340::
1332::
1305:.
1283::
1256:.
1232::
1226:4
1205:.
1193::
1170:.
1148::
1121:.
1097::
1070:.
1056:.
1042:.
1030::
1007:.
985::
958:.
946::
923:.
909::
901::
875:.
863::
824:.
812::
785:.
766:.
744::
714:.
692::
684::
506:"
428:.
201:(
20:)
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
