373:
324:. The stages that a cell passes through that culminate in its division and replication are collectively known as the cell cycle Since the successful division and replication of a cell is essential for its survival, the cell cycle is tightly regulated by several components to ensure the efficient and error-free progression through the cell cycle. One such regulatory component is cyclin A which plays a role in the regulation of two different cell cycle stages.
608:(pRb) is involved in the regulation of cyclin A through its interaction with E2F. It exists in two states: hypophosphorylated pRb and hyperphosphorylated pRb. Hypophosphorylated pRb binds E2F, which prevents transcription of cyclin A. The absence of cyclin A prior to the R point is due to the inhibition of E2F by hypophosphorylated pRb. After the cell passes through the R point,
525:. Cyclin A binds to the mitotic spindle poles in the centrosome however, the mechanism by which the complex is shuttled to the centrosome is not well understood. It is suspected that the presence of cyclin A/CDK2 at the centrosomes may confer a means of regulating the movement of cyclin B/CDK1 to the centrosome and thus the timing of mitotic events.
655:, and blocks the kinase activity of CDKs. Activated p21 can bind cyclin D/CDK4 and render it incapable of phosphorylating pRb. PRb remains hypophosphorylated and binds E2F. E2F is unable to activate the transcription of cyclins involved in cell cycle progression, such as cyclin A and the cell cycle is arrested at G
623:
molecule called the cell-cycle-responsive element (CCRE). E2F binds to an E2F binding site on the CCRE, releasing the repressor from the promoter and allowing the transcription of cyclin A. Cyclin A/CDK2 will eventually phosphorylate E2F when cyclin A reaches a certain level, completing the negative
407:, confers the foundation of the cyclin-CDK complex. Cyclin A is the only cyclin that regulates multiple steps of the cell cycle. Cyclin A can regulate multiple cell cycle steps because it associates with, and thereby activates, two distinct CDKs –
472:
Cyclin A has a second function in S phase. In addition to initiating DNA synthesis, Cyclin A ensures that DNA is replicated once per cell cycle by preventing the assembly of additional replication complexes. This is thought to occur through the
481:, by the cyclin A/CDK2 complex. Since the action of cyclin A/CDK2 inhibits that of cyclin E/CDK2, the sequential activation of cyclin E followed by the activation of cyclin A is important and tightly regulated in S phase.
516:
Cyclin A/CDK2 complex was thought to be restricted to the nucleus and thus exclusively involved in S phase progression. New research has since debunked this assumption, shedding light on cyclin A/CDK2 migration to the
505:. Cyclin A/CDK1 is thought to be involved in the activation and stabilization of cyclin B/CDK1 complex. Once cyclin B is activated, cyclin A is no longer needed and is subsequently degraded through the
1209:
Jeffrey PD, Russo AA, Polyak K, Gibbs E, Hurwitz J, Massagué J, Pavletich NP (July 1995). "Mechanism of CDK activation revealed by the structure of a cyclinA-CDK2 complex".
612:/E- complexes phosphorylate pRb. Hyperphosphorylated pRb can no longer bind E2F, E2F is released and cyclin A genes, and other crucial genes for S phase, are transcribed.
643:
and turns on several downstream pathways, including cell cycle arrest. Cell cycle arrest is carried out by the p53-pRb pathway. Activated p53 turns on genes for
465:
capable of replication. When the amount of cyclin A/CDK2 complex reaches a threshold level, it terminates the assembly of the pre-replication complex made by
566:
of cyclin A is tightly regulated and synchronized with cell cycle progression. Initiation of transcription of cyclin A is coordinated with passage of the
364:, the somatic form. Cyclin A1 is prevalently expressed during meiosis and early on in embryogenesis. Cyclin A2 is expressed in dividing somatic cells.
193:
44:
624:
feedback loop. Phosphorylation of E2F turns the transcription factor off, providing another level of controlling the transcription of cyclin A.
1455:
1265:
Coverley D, Laman H, Laskey RA (July 2002). "Distinct roles for cyclins E and A during DNA replication complex assembly and activation".
430:. Association of cyclin A with CDK2 is required for passage into S phase while association with CDK1 is required for entry into M phase.
356:
form. A single cyclin A gene has been identified in
Drosophila while Xenopus, mice and humans contain two distinct types of cyclin A:
1747:
1641:
648:
597:. E2F is responsible for initiating the transcription of many critical S phase genes. Cyclin A transcription is off during most of G
372:
49:
1953:
1165:"Deregulated expression of E2F-1 induces cyclin A- and E-associated kinase activities independently from cell cycle position"
777:
198:
1958:
1687:
659:. Cell cycle arrest allows the cell to repair DNA damage before the cell divides and passes damaged DNA to daughter cells.
217:
68:
1448:
540:
in the centrosome with mitotic events in the nucleus was lost in the cyclin A knockout/CDK2 inhibited mutant cells.
493:
In late S phase, cyclin A can also associate with CDK1. Cyclin A remains associated with CDK1 from late S into late
205:
56:
1973:
1115:
De Boer L, Oakes V, Beamish H, Giles N, Stevens F, Somodevilla-Torres M, Desouza C, Gabrielli B (July 2008).
867:"Cyclin: a protein specified by maternal mRNA in sea urchin eggs that is destroyed at each cleavage division"
1441:
633:
536:. These mutants entered mitosis late due to a delayed activation of the cyclin B/CDK1 complex. Coupling of
348:, mice, and in humans but has not been found in lower eukaryotes like yeast. The protein exists in both an
698:"Cyclin A2 mutagenesis analysis: a new insight into CDK activation and cellular localization requirements"
1757:
210:
61:
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1631:
114:
118:
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819:
709:
469:. As the amount of Cyclin A/CDK2 complex increases, the complex initiates DNA replication.
263:
574:
into S phase. Transcription peaks and plateaus mid-S phase and abruptly declines in late G
332:
Cyclin A was first identified in 1983 in sea urchin embryos. Since its initial discovery,
287:
138:
8:
1963:
267:
1222:
1073:
1048:
823:
713:
1893:
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1968:
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1384:
1338:
1282:
1234:
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1078:
1000:
940:
911:"Cyclin A expression is under negative transcriptional control during the cell cycle"
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773:
737:
353:
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1396:
415:. Depending on which CDK partner cyclin A binds, the cell will continue through the
1374:
1328:
1294:
1274:
1246:
1226:
1176:
1163:
Soucek T, Pusch O, Hengstschläger-Ottnad E, Adams PD, Hengstschläger M (May 1997).
1128:
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that binds to several cyclin/CDK complexes, including cyclin A-CDK2/1 and cyclin D/
258:
109:
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722:
474:
443:
234:
85:
297:
148:
532:. Cells were modified so their CDK2 was inhibited and their cyclin A2 gene was
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1947:
1827:
808:"Structure and cell cycle-regulated transcription of the human cyclin A gene"
533:
528:
A study in 2008 provided further evidence of cyclin A/CDK2 complex's role in
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832:
1849:
1342:
1286:
1181:
1164:
1142:
1082:
741:
616:
510:
391:(CDKs), which thereby activates the enzymatic activity of its CDK partner.
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1238:
1190:
1004:
944:
926:
892:
851:
1870:
1839:
518:
403:, a region conserved across cyclins, and a region of the CDK, called the
246:
97:
1133:
1116:
1791:
1464:
1333:
1316:
640:
383:
Cyclin A, along with the other members of the cyclin family, regulates
377:
338:
321:
241:
92:
1433:
1914:
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1523:
1519:
1515:
1505:
1501:
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1162:
620:
570:, a critical transition point that is required for progression from G
506:
462:
361:
357:
163:
1919:
1859:
1844:
1813:
1799:
1529:
1511:
1497:
1278:
969:
Pagano M, Pepperkok R, Verde F, Ansorge W, Draetta G (March 1992).
609:
502:
454:
447:
420:
317:
1117:"Cyclin A/cdk2 coordinates centrosomal and nuclear mitotic events"
865:
Evans T, Rosenthal ET, Youngblom J, Distel D, Hunt T (June 1983).
336:
of cyclin A have been identified in numerous eukaryotes including
1928:
1835:
1808:
1467:
1317:"Cyclin-dependent kinases and S phase control in mammalian cells"
806:
Henglein B, Chenivesse X, Wang J, Eick D, Bréchot C (June 1994).
567:
529:
427:
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344:
222:
73:
1767:
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349:
313:
229:
80:
1742:
1737:
1626:
1621:
1616:
864:
457:. Cyclin E is responsible for initiating the assembly of the
178:
37:
22:
971:"Cyclin A is required at two points in the human cell cycle"
695:
586:
Transcription of cyclin A is predominantly regulated by the
477:
of particular DNA replication machinery components, such as
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909:
Huet X, Rech J, Plet A, Vié A, Blanchard JM (July 1996).
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632:
Transcription of cyclin A is indirectly regulated by the
587:
442:
where it is involved in the initiation and completion of
1114:
1208:
543:
Cyclin A has been shown to play a crucial role in the G
453:
into S phase, cyclin A associates with CDK2, replacing
1363:"p53, the cellular gatekeeper for growth and division"
696:
Bendris N, Lemmers B, Blanchard JM, Arsic N (2011).
320:
that function in regulating progression through the
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367:
1945:
908:
509:pathway. Degradation of cyclin A/CDK1 induces
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1202:
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691:
1049:"Cyclin A in cell cycle control and cancer"
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615:E2F initiates transcription of cyclin A by
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601:and the begins shortly after the R point.
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1748:Cellular apoptosis susceptibility protein
1426:at the U.S. National Library of Medicine
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619:the promoter. The promoter is bound by a
1047:Yam CH, Fung TK, Poon RY (August 2002).
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376:Expression of human cyclins through the
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1018:
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438:Cyclin A resides in the nucleus during
1946:
1301:
1437:
899:
1011:
387:through physically interacting with
360:, the embryonic-specific form, and
13:
987:10.1002/j.1460-2075.1992.tb05135.x
371:
14:
1985:
1406:
484:
627:
581:
368:Role in cell cycle progression
1:
1380:10.1016/S0092-8674(00)81871-1
772:. New York: Garland Science.
662:
558:
1954:Genes on human chromosome 13
884:10.1016/0092-8674(83)90420-8
812:Proc. Natl. Acad. Sci. U.S.A
723:10.1371/journal.pone.0022879
399:The interaction between the
7:
1959:Genes on human chromosome 4
1758:Maturation promoting factor
1361:Levine AJ (February 1997).
419:or it will transition from
10:
1990:
1899:Postreplication checkpoint
446:. As the cell passes from
433:
1907:
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1826:
1790:
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1725:
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1065:10.1007/s00018-002-8510-y
634:tumor suppressor protein
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1428:Medical Subject Headings
1315:Woo RA, Poon RY (2003).
389:cyclin-dependent kinases
327:
833:10.1073/pnas.91.12.5490
501:when it is replaced by
459:pre-replication complex
395:CDK partner association
1881:Cell cycle checkpoints
1182:10.1038/sj.onc.1201061
639:. P53 is activated by
606:retinoblastoma protein
595:negative feedback loop
538:microtubule nucleation
385:cell cycle progression
380:
1974:Cell cycle regulators
1908:Other cellular phases
1632:CDK-activating kinase
1419:- The Interactive Fly
927:10.1128/mcb.16.7.3789
770:The biology of cancer
588:transcription factor
461:. This complex makes
375:
768:Weinberg RE (2007).
1223:1995Natur.376..313J
1134:10.1038/onc.2008.74
1053:Cell. Mol. Life Sci
824:1994PNAS...91.5490H
714:2011PLoSO...622879B
316:family, a group of
312:is a member of the
1894:Spindle checkpoint
1699:P53 p63 p73 family
1334:10.4161/cc.2.4.468
381:
1941:
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1889:Restriction point
779:978-0-8153-4076-8
547:/M transition in
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1279:10.1038/ncb813
1267:Nat. Cell Biol
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988:
984:
981:(3): 961–71.
980:
976:
972:
965:
963:
961:
959:
957:
955:
946:
942:
937:
932:
928:
924:
920:
916:
912:
905:
903:
894:
890:
885:
880:
877:(2): 389–96.
876:
872:
868:
861:
853:
849:
844:
839:
834:
829:
825:
821:
817:
813:
809:
802:
800:
798:
796:
794:
792:
790:
781:
775:
771:
764:
762:
760:
758:
756:
754:
752:
743:
739:
734:
729:
724:
719:
715:
711:
708:(7): e22879.
707:
703:
699:
692:
690:
688:
686:
684:
682:
680:
678:
676:
674:
672:
667:
660:
654:
650:
649:CDK inhibitor
646:
642:
638:
637:
625:
622:
618:
617:de-repressing
613:
611:
607:
602:
596:
592:
591:
579:
569:
565:
564:Transcription
556:
554:
550:
541:
539:
535:
531:
526:
520:
514:
512:
508:
504:
500:
482:
480:
476:
470:
468:
467:cyclin E/CDK2
464:
460:
456:
452:
445:
441:
431:
429:
425:
418:
414:
410:
406:
402:
392:
390:
386:
379:
374:
365:
363:
359:
355:
351:
347:
346:
341:
340:
335:
325:
323:
319:
315:
311:
299:
296:
292:
289:
286:
282:
277:
273:
270:
269:
265:
262:
260:
256:
251:
248:
245:
243:
239:
236:
233:
231:
227:
224:
221:
219:
215:
212:
209:
207:
203:
200:
197:
195:
191:
187:
183:
180:
177:
173:
168:
165:
161:
150:
147:
143:
140:
137:
133:
128:
124:
121:
120:
116:
113:
111:
107:
102:
99:
96:
94:
90:
87:
84:
82:
78:
75:
72:
70:
66:
63:
60:
58:
54:
51:
48:
46:
42:
39:
36:
32:
27:
24:
20:
1850:Prometaphase
1483:
1416:
1413:
1370:
1366:
1324:
1320:
1273:(7): 523–8.
1270:
1266:
1214:
1210:
1172:
1168:
1124:
1120:
1056:
1052:
978:
974:
918:
914:
874:
870:
860:
815:
811:
769:
705:
701:
635:
631:
614:
603:
589:
585:
562:
552:
548:
542:
527:
515:
511:mitotic exit
494:
492:
471:
439:
437:
398:
384:
382:
343:
337:
331:
309:
308:
266:
185:Alt. symbols
117:
1871:Cytokinesis
1840:Preprophase
1784:checkpoints
647:. P21 is a
628:p53 and p21
582:E2F and pRb
534:knocked out
519:centrosomes
288:Swiss-model
170:Identifiers
139:Swiss-model
29:Identifiers
1964:Cell cycle
1948:Categories
1792:Interphase
1782:Phases and
1465:Cell cycle
1414:Drosophila
1321:Cell Cycle
663:References
641:DNA damage
559:Regulation
549:Drosophila
401:cyclin box
378:cell cycle
339:Drosophila
334:homologues
322:cell cycle
284:Structures
279:Search for
253:Other data
188:CCNA, CCN1
135:Structures
130:Search for
104:Other data
1915:Apoptosis
1865:Telophase
1855:Metaphase
1650:INK4a/ARF
621:repressor
555:embryos.
521:in late G
507:ubiquitin
489:/ M phase
463:chromatin
352:form and
350:embryonic
235:NM_001237
194:NCBI gene
164:cyclin A2
119:q12.3-q13
86:NM_003914
45:NCBI gene
23:cyclin A1
1969:Proteins
1860:Anaphase
1845:Prophase
1468:proteins
1424:Cyclin+A
1417:Cyclin A
1397:17192968
1343:12851482
1287:12080347
1169:Oncogene
1143:18372919
1121:Oncogene
1083:12363035
1074:11337442
742:21829545
702:PLOS ONE
610:cyclin D
503:cyclin B
455:cyclin E
318:proteins
310:Cyclin A
298:InterPro
149:InterPro
1929:Meiosis
1836:Mitosis
1828:M phase
1809:S phase
1676:cip/kip
1389:9039259
1295:4667087
1247:4361179
1239:7630397
1219:Bibcode
1191:9178900
1005:1312467
945:8668196
893:6134587
852:8202514
820:Bibcode
733:3145769
710:Bibcode
568:R point
553:Xenopus
530:mitosis
440:S phase
434:S phase
428:M phase
426:to the
417:S phase
405:PSTAIRE
354:somatic
345:Xenopus
294:Domains
242:UniProt
145:Domains
115:Chr. 13
93:UniProt
1768:Cullin
1654:p14arf
1476:Cyclin
1430:(MeSH)
1395:
1387:
1341:
1293:
1285:
1245:
1237:
1211:Nature
1189:
1141:
1089:
1081:
1071:
1003:
996:556537
993:
975:EMBO J
943:
936:231375
933:
891:
850:
840:
776:
740:
730:
314:cyclin
264:Chr. 4
247:P20248
230:RefSeq
223:123835
175:Symbol
98:P78396
81:RefSeq
74:604036
34:Symbol
1924:phase
1818:phase
1804:phase
1743:Cdc42
1738:Cdc25
1726:Other
1508:, B3)
1393:S2CID
1291:S2CID
1243:S2CID
1091:97398
1087:S2CID
843:44021
593:in a
499:phase
328:Types
259:Locus
179:CCNA2
110:Locus
38:CCNA1
1772:CUL7
1733:Cdc2
1385:PMID
1367:Cell
1339:PMID
1283:PMID
1235:PMID
1187:PMID
1139:PMID
1079:PMID
1001:PMID
941:PMID
889:PMID
871:Cell
848:PMID
774:ISBN
738:PMID
653:CDK4
604:The
551:and
479:CDC6
413:CDK1
411:and
409:CDK2
218:OMIM
211:1578
206:HGNC
69:OMIM
62:1577
57:HGNC
50:8900
1763:Wee
1753:E2F
1717:p73
1712:p63
1707:p53
1688:p57
1684:p27
1680:p21
1670:p19
1666:p18
1662:p15
1658:p16
1612:11B
1607:11A
1549:CDK
1375:doi
1329:doi
1275:doi
1227:doi
1215:376
1177:doi
1129:doi
1069:PMC
1061:doi
991:PMC
983:doi
931:PMC
923:doi
879:doi
838:PMC
828:doi
728:PMC
718:doi
645:p21
636:p53
590:E2F
268:q27
199:890
1950::
1686:,
1682:,
1668:,
1664:,
1660:,
1627:14
1622:13
1617:12
1602:10
1538:E2
1536:,
1534:E1
1524:D3
1522:,
1520:D2
1518:,
1516:D1
1506:B2
1504:,
1502:B1
1492:A2
1490:,
1488:A1
1391:.
1383:.
1371:88
1369:.
1365:.
1351:^
1337:.
1323:.
1319:.
1303:^
1289:.
1281:.
1269:.
1255:^
1241:.
1233:.
1225:.
1213:.
1199:^
1185:.
1173:14
1171:.
1167:.
1151:^
1137:.
1125:27
1123:.
1119:.
1099:^
1085:.
1077:.
1067:.
1057:59
1055:.
1051:.
1013:^
999:.
989:.
979:11
977:.
973:.
953:^
939:.
929:.
919:16
917:.
913:.
901:^
887:.
875:33
873:.
869:.
846:.
836:.
826:.
816:91
814:.
810:.
788:^
750:^
736:.
726:.
716:.
704:.
700:.
670:^
578:.
513:.
362:A2
358:A1
342:,
1922:0
1920:G
1867:)
1838:(
1816:2
1814:G
1802:1
1800:G
1774:)
1770:(
1690:)
1678:(
1672:)
1656:/
1652:(
1597:9
1592:8
1587:7
1582:6
1577:5
1572:4
1567:3
1562:2
1557:1
1540:)
1532:(
1530:E
1526:)
1514:(
1512:D
1500:(
1498:B
1494:)
1486:(
1484:A
1457:e
1450:t
1443:v
1399:.
1377::
1345:.
1331::
1325:2
1297:.
1277::
1271:4
1249:.
1229::
1221::
1193:.
1179::
1145:.
1131::
1093:.
1063::
1007:.
985::
947:.
925::
895:.
881::
854:.
830::
822::
782:.
744:.
720::
712::
706:6
657:1
599:1
576:2
572:1
545:2
523:2
497:2
495:G
487:2
485:G
450:1
448:G
423:2
421:G
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