Contents 1 Etymology 2 Function 3 Domain structure 4 Types 4.1 Main groups 4.2 Subtypes 5 Other Proteins Containing This Domain 6 History 7 References 8 External links 9 Further reading

Etymology[edit] Cyclins were originally discovered by R. Timothy Hunt in 1982 while studying the cell cycle of sea urchins.[2][3] In an interview for "The Life Scientific" (aired on 13/12/2011) hosted by Jim Al-Khalili, R. Timothy Hunt explained that the name "cyclin" was originally named after his hobby cycling. It was only after the naming did its importance in the cell cycle become apparent. As it was appropriate the name stuck.[4] R. Timothy Hunt: "By the way, the name cyclin, which I coined, was really a joke, it's because I liked cycling so much at the time, but they did come and go in the cell..."[4]

Function[edit] Expression of human cyclins through the cell cycle. Cyclins were originally named because their concentration varies in a cyclical fashion during the cell cycle. (Note that the cyclins are now classified according to their conserved cyclin box structure, and not all these cyclins alter in level through the cell cycle.[5]) The oscillations of the cyclins, namely fluctuations in cyclin gene expression and destruction by the ubiquitin mediated proteasome pathway, induce oscillations in Cdk activity to drive the cell cycle. A cyclin forms a complex with Cdk, which begins to activate the Cdk, but the complete activation requires phosphorylation, as well. Complex formation results in activation of the Cdk active site. Cyclins themselves have no enzymatic activity but have binding sites for some substrates and target the Cdks to specific subcellular locations.[5] Cyclins, when bound with the dependent kinases, such as the p34/cdc2/cdk1 protein, form the maturation-promoting factor. MPFs activate other proteins through phosphorylation. These phosphorylated proteins, in turn, are responsible for specific events during cycle division such as microtubule formation and chromatin remodeling. Cyclins can be divided into four classes based on their behavior in the cell cycle of vertebrate somatic cells and yeast cells: G1 cyclins, G1/S cyclins, S cyclins, and M cyclins. This division is useful when talking about most cell cycles, but it is not universal as some cyclins have different functions or timing in different cell types. G1/S Cyclins rise in late G1 and fall in early S phase. The Cdk- G1/S cyclin complex begins to induce the initial processes of DNA replication, primarily by arresting systems that prevent S phase Cdk activity in G1. The cyclins also promote other activities to progress the cell cycle, such as centrosome duplication in vertebrates or spindle pole body in yeast. The rise in presence of G1/S cyclins is paralleled by a rise in S cyclins. G1 cyclins do not behave like the other cyclins, in that the concentrations increase gradually (with no oscillation), throughout the cell cycle based on cell growth and the external growth-regulatory signals. The presence of G cyclins coordinate cell growth with the entry to a new cell cycle. S cyclins bind to Cdk and the complex directly induces DNA replication. The levels of S cyclins remain high, not only throughout S phase, but through G2 and early mitosis as well to promote early events in mitosis. M cyclin concentrations rise as the cell begins to enter mitosis and the concentrations peak at metaphase. Cell changes in the cell cycle like the assembly of mitotic spindles and alignment of sister-chromatids along the spindles are induced by M cyclin- Cdk complexes. The destruction of M cyclins during metaphase and anaphase, after the Spindle Assembly Checkpoint is satisfied, causes the exit of mitosis and cytokinesis.[6] Expression of cyclins in individual cells in relation to cellular DNA content (cell cycle phase) is often measured by flow cytometry.[7] Their expression is also being measured by flow cytometry concurrently with the initiation and termination of DNA replication during S-phase [8]

Domain structure[edit] Cyclins are generally very different from each other in primary structure, or amino acid sequence. However, all members of the cyclin family are similar in 100 amino acids that make up the cyclin box. Cyclins contain two domains of a similar all-α fold, the first located at the N-terminus and the second at the C-terminus. All cyclins are believed to contain a similar tertiary structure of two compact domains of 5 α helices. The first of which is the conserved cyclin box, outside of which cyclins are divergent. For example, the amino-terminal regions of S and M cyclins contain short destruction-box motifs that target these proteins for proteolysis in mitosis. Cyclin, N-terminal domain Structure of bovine cyclin A.[9] Identifiers Symbol Cyclin_N Pfam PF00134 Pfam clan CL0065 InterPro IPR006671 PROSITE PDOC00264 SCOP 1vin SUPERFAMILY 1vin Available protein structures: Pfam structures PDB RCSB PDB; PDBe; PDBj PDBsum structure summary Cyclin, C-terminal domain Structure of CDK2-cyclin A/indirubin-5-sulphonate.[10] Identifiers Symbol Cyclin_C Pfam PF02984 Pfam clan CL0065 InterPro IPR004367 PROSITE PDOC00264 SCOP 1vin SUPERFAMILY 1vin Available protein structures: Pfam structures PDB RCSB PDB; PDBe; PDBj PDBsum structure summary K cyclin, C terminal structure of a p18(ink4c)-cdk6-k-cyclin ternary complex Identifiers Symbol K-cyclin_vir_C Pfam PF09080 InterPro IPR015164 SCOP 1g3n SUPERFAMILY 1g3n Available protein structures: Pfam structures PDB RCSB PDB; PDBe; PDBj PDBsum structure summary

Types[edit] There are several different cyclins that are active in different parts of the cell cycle and that cause the Cdk to phosphorylate different substrates. There are also several "orphan" cyclins for which no Cdk partner has been identified. For example, cyclin F is an orphan cyclin that is essential for G2/M transition.[11][12] A study in C. elegans revealed the specific roles of mitotic cyclins.[13][14] Notably, recent studies have shown that cyclin A creates a cellular environment that promotes microtubule detachment from kinetochores in prometaphase to ensure efficient error correction and faithful chromosome segregation. Cells must separate their chromosomes precisely, an event that relies on the bi-oriented attachment of chromosomes to spindle microtubules through specialized structures called kinetochores. In the early phases of division, there are numerous errors in how kinetochores bind to spindle microtubules. The unstable attachments promote the correction of errors by causing a constant detachment, realignment and reattachment of microtubules from kinetochores in the cells as they try to find the correct attachment. Protein cyclin A governs this process by keeping the process going until the errors are eliminated. In normal cells, persistent cyclin A expression prevents the stabilization of microtubules bound to kinetochores even in cells with aligned chromosomes. As levels of cyclin A decline, microtubule attachments become stable, allowing the chromosomes to be divided correctly as cell division proceeds. In contrast, in cyclin A-deficient cells, microtubule attachments are prematurely stabilized. Consequently, these cells may fail to correct errors, leading to higher rates of chromosome mis-segregation.[15] Main groups[edit] There are two main groups of cyclins: G1/S cyclins – essential for the control of the cell cycle at the G1/S transition, Cyclin A / CDK2 – active in S phase. Cyclin D / CDK4, Cyclin D / CDK6, and Cyclin E / CDK2 – regulates transition from G1 to S phase. G2/M cyclins – essential for the control of the cell cycle at the G2/M transition (mitosis). G2/M cyclins accumulate steadily during G2 and are abruptly destroyed as cells exit from mitosis (at the end of the M-phase). Cyclin B / CDK1 – regulates progression from G2 to M phase. Subtypes[edit] Specific cyclin subtypes include: Species G1 G1/S S M S. cerevisiae Cln3 (Cdk1) Cln 1,2 (Cdk1) Clb 5,6 (Cdk1) Clb 1,2,3,4 (Cdk 1) S. pombe Puc1? (Cdk1) Puc1, Cig1? (Cdk1) Cig2, Cig1? (Cdk1) Cdc13 (Cdk1) D. melanogaster cyclin D (Cdk4) cyclin E (Cdk2) cyclin E, A (Cdk2,1) cyclin A, B, B3 (Cdk1) X. laevis either not known or not present cyclin E (Cdk2) cyclin E, A (Cdk2,1) cyclin A, B, B3 (Cdk1) H. sapiens cyclin D 1,2,3 (Cdk4, Cdk6) cyclin E (Cdk2) cyclin A (Cdk2, Cdk1) cyclin B (Cdk1) family members A CCNA1, CCNA2 B CCNB1, CCNB2, CCNB3 C CCNC D CCND1, CCND2, CCND3 E CCNE1, CCNE2 F CCNF G CCNG1, CCNG2 H CCNH I CCNI, CCNI2 J CCNJ, CCNJL K CCNK L CCNL1, CCNL2 O CCNO T CCNT1, CCNT2 Y CCNY, CCNYL1, CCNYL2, CCNYL3

Other Proteins Containing This Domain[edit] In addition, the following human protein contains a cyclin domain: CNTD1

History[edit] Leland H. Hartwell, R. Timothy Hunt, and Paul M. Nurse won the 2001 Nobel Prize in Physiology or Medicine for their discovery of cyclin and cyclin-dependent kinase.[16]

References[edit] ^ Galderisi U, Jori FP, Giordano A (August 2003). "Cell cycle regulation and neural differentiation". Oncogene. 22 (33): 5208–19. doi:10.1038/sj.onc.1206558. PMID 12910258.  ^ Evans et al., 1983, Cell 33, p389-396 ^ ^ a b "The Life Scientific". BBC Radio 4. BBC. Retrieved 13 December 2011.  ^ a b Morgan, DO (2007) 'The Cell Cycle: Principles of Control, Oxford University Press ^ Clute and Pines, (1999) Nature Cell Biology, 1, p82-87 ^ Darzynkiewicz Z, Gong J, Juan G, Ardelt B, Traganos F. Cytometry of cyclin proteins. (1996) Cytometry. 25(1):1-13. Review. PMID 8875049; doi:10.1002/(SICI)1097-0320(19960901)25:1<1::AID-CYTO1>3.0.CO;2-N ^ Darzynkiewicz Z, Zhao H, Zhang S, Lee MY, Lee EY, Zhang Z. (2015)Initiation and termination of DNA replication during S phase in relation to cyclins D1, E and A, p21WAF1, Cdt1 and the p12 subunit of DNA polymerase δ revealed in individual cells by cytometry. Oncotarget. 6(14):11735-50. Review. PMID 26059433; doi:10.18632/oncotarget.4149 ;PMCID: PMC4494901 ^ Brown NR, Noble ME, Endicott JA, et al. (November 1995). "The crystal structure of cyclin A". Structure. 3 (11): 1235–47. doi:10.1016/S0969-2126(01)00259-3. PMID 8591034.  ^ Davies TG, Tunnah P, Meijer L, et al. (May 2001). "Inhibitor binding to active and inactive CDK2: the crystal structure of CDK2-cyclin A/indirubin-5-sulphonate". Structure. 9 (5): 389–97. doi:10.1016/S0969-2126(01)00598-6. PMID 11377199.  ^ Fung TK, Poon RY (2005). "A roller coaster ride with the mitotic cyclins". Semin. Cell Dev. Biol. 16 (3): 335–42. doi:10.1016/j.semcdb.2005.02.014. PMID 15840442.  ^ Gerald Karp (2007). Cell and Molecular Biology: Concepts and Experiments. New York: Wiley. pp. 148, 165–170, and 624–664. ISBN 0-470-04217-6.  ^ van der Voet, Monique; Lorson, Monique; Srinivasan, Dayalan G.; Bennett, Karen L.; van den Heuvel, Sander (2009). "C. elegans mitotic cyclins have distinct as well as overlapping functions in chromosome segregation". Cell Cycle. 8 (24): 4091–4102. doi:10.4161/cc.8.24.10171. ISSN 1538-4101. PMC 3614003 . PMID 19829076.  ^ Rahman, Mohammad M.; Kipreos, Edward (2010). "The specific roles of mitotic cyclins revealed". Cell Cycle. 9 (1): 22–27. doi:10.4161/cc.9.1.10577. ISSN 1538-4101.  ^ "Cell cycle: Cyclin A corrections". Nature Reviews Molecular Cell Biology. 14: 692. doi:10.1038/nrm3680.  ^ "The Nobel Prize in Physiology or Medicine 2001". The Nobel Foundation. Retrieved 2009-03-15. 

External links[edit] Eukaryotic Linear Motif resource motif class LIG_CYCLIN_1

Further reading[edit] Monty Krieger; Matthew P Scott; Matsudaira, Paul T.; Lodish, Harvey F.; Darnell, James E.; Lawrence Zipursky; Kaiser, Chris; Arnold Berk (2004). Molecular cell biology (Fifth ed.). New York: W.H. Freeman and CO. ISBN 0-7167-4366-3.  v t e Intracellular signaling peptides and proteins MAP see MAP kinase pathway Calcium Intracellular calcium-sensing proteins Calcineurin Ca2+/calmodulin-dependent protein kinase G protein Heterotrimeric cAMP: Heterotrimeric G protein Gs/Gi Adenylate cyclase cAMP 3',5'-cyclic-AMP phosphodiesterase Protein kinase A cGMP: Transducin Gustducin Guanylate cyclase cGMP 3',5'-cyclic-GMP phosphodiesterase Protein kinase G G alpha subunit Gα GNAO1 GNAI1 GNAI2 GNAI3 GNAT1 GNAT2 GNAT3 GNAZ GNAS GNAL GNAQ GNA11 GNA12 GNA13 GNA14 GNA15/GNA16 G beta-gamma complex Gβ GNB1 GNB2 GNB3 GNB4 GNB5 Gγ GNGT1 GNGT2 GNG2 GNG3 GNG4 GNG5 GNG7 GNG8 GNG10 GNG11 GNG12 GNG13 BSCL2 G protein-coupled receptor kinase AMP-activated protein kinase Monomeric ARFs Rabs Ras HRAS KRAS NRAS Rhos Arfs Ran Rhebs Raps RGKs Cyclin Cyclin-dependent kinase inhibitor protein Cyclin-dependent kinase Cyclin Lipid Phosphoinositide phospholipase C Phospholipase C Other protein kinase Serine/threonine: Casein kinase 1 2 eIF-2 kinase EIF2AK3 Glycogen synthase kinase GSK1 GSK2 GSK-3 GSK3A GSK3B IκB kinase CHUK IKK2 IKBKG Interleukin-1 receptor associated kinase IRAK1 IRAK2 IRAK3 IRAK4 Lim kinase LIMK1 LIMK2 p21-activated kinases PAK1 PAK2 PAK3 PAK4 Rho-associated protein kinase ROCK1 ROCK2 Ribosomal s6 kinase RPS6KA1 Tyrosine: ZAP70 Focal adhesion protein-tyrosine kinase PTK2 PTK2B BTK both Dual-specificity kinase Other protein phosphatase Serine/threonine: Protein phosphatase 2 Tyrosine: protein tyrosine phosphatase: Receptor-like protein tyrosine phosphatase Sh2 domain-containing protein tyrosine phosphatase both: Dual-specificity phosphatase Apoptosis see apoptosis signaling pathway GTP-binding protein regulators see GTP-binding protein regulators Other Activating transcription factor 6 Signal transducing adaptor protein I-kappa B protein Mucin-4 Olfactory marker protein Phosphatidylethanolamine binding protein EDARADD PRKCSH see also deficiencies of intracellular signaling peptides and proteins v t e Cell cycle proteins Cyclin A (A1, A2) B (B1, B2, B3) D (D1, D2, D3) E (E1, E2) CDK 1 2 3 4 5 6 7 8 9 10 CDK-activating kinase CDK inhibitor INK4a/ARF (p14arf/p16, p15, p18, p19) cip/kip (p21, p27, p57) P53 p63 p73 family p53 p63 p73 Other Cdc2 Cdc25 Cdc42 Cellular apoptosis susceptibility protein E2F Maturation promoting factor Wee Cullin (CUL7) Phases and checkpoints Interphase G1 phase S phase G2 phase M phase Mitosis (Preprophase Prophase Prometaphase Metaphase Anaphase Telophase) Cytokinesis Cell cycle checkpoints Restriction point Spindle checkpoint Postreplication checkpoint Other cellular phases Apoptosis G0 phase Meiosis This article incorporates text from the public domain Pfam and InterPro IPR006671 Retrieved from "" Categories: Cell cycleProteins

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Cyclin - Photos and All Basic Informations

Cyclin More Links

Protein FamilyCell CycleCyclin-dependent KinaseEnzymeR. Timothy HuntJim Al-KhaliliR. Timothy HuntR. Timothy HuntEnlargeCell CycleActive SiteKinaseCdk1Cdk1Cyclin-dependent Kinase 1Maturation-promoting FactorPhosphorylationMicrotubuleChromatin Structure Remodeling (RSC) ComplexCentrosomeSpindle Pole BodyFlow CytometryFlow CytometryDNA ReplicationProtein DomainAll-α ProteinsN-terminusC-terminusPfamPfamInterProPROSITEStructural Classification Of ProteinsSUPERFAMILYPfamProtein Data BankPDBsumPfamPfamInterProPROSITEStructural Classification Of ProteinsSUPERFAMILYPfamProtein Data BankPDBsumPfamInterProStructural Classification Of ProteinsSUPERFAMILYPfamProtein Data BankPDBsumCell Cycle CheckpointCyclin ACyclin-dependent Kinase 2Cyclin DCyclin-dependent Kinase 4Cyclin-dependent Kinase 6Cyclin ECell Cycle CheckpointMitosisMetaphaseCyclin BCdk1Saccharomyces CerevisiaeSchizosaccharomyces PombeDrosophila MelanogasterXenopus LaevisHomo SapiensCyclin DCdk4Cdk6Cyclin ECdk2Cyclin ACdk2Cdk1Cyclin BCdk1Cyclin ACyclin A1Cyclin A2Cyclin BCyclin B1Cyclin B2Cyclin CCyclin DCyclin D1Cyclin D2Cyclin D3Cyclin ECyclin E1Cyclin E2CCNFCCNG1CCNG2Cyclin HCCNI (gene)Cyclin KCCNL1CCNL2Cyclin OCyclin T1Cyclin T2Leland H. HartwellR. Timothy HuntPaul M. NurseNobel Prize In Physiology Or MedicineDigital Object IdentifierPubMed IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierInternational Standard Book NumberSpecial:BookSources/0-470-04217-6Digital Object IdentifierInternational Standard Serial NumberPubMed CentralPubMed IdentifierDigital Object IdentifierInternational Standard Serial NumberDigital Object IdentifierEukaryotic Linear Motif ResourceInternational Standard Book NumberSpecial:BookSources/0-7167-4366-3Template:Intracellular Signaling Peptides And ProteinsTemplate Talk:Intracellular Signaling Peptides And ProteinsIntracellular Signaling Peptides And ProteinsTemplate:MAP Kinase PathwayIntracellular Calcium-sensing ProteinsCalcineurinCa2+/calmodulin-dependent Protein KinaseG ProteinHeterotrimeric G ProteinCAMP-dependent PathwayHeterotrimeric G ProteinGs Alpha SubunitGi Alpha SubunitAdenylate CyclaseCyclic Adenosine Monophosphate3',5'-cyclic-AMP PhosphodiesteraseProtein Kinase ACyclic Guanosine MonophosphateTransducinGustducinGuanylate CyclaseCyclic Guanosine Monophosphate3',5'-cyclic-GMP PhosphodiesteraseCGMP-dependent Protein KinaseG Alpha SubunitGNAO1GNAI1GNAI2GNAI3GNAT1GNAT2GNAT3GNAZGNAS Complex LocusGNALGNAQGNA11GNA12GNA13G Beta-gamma ComplexGNB1GNB2GNB3GNB4GNB5GNGT1GNGT2GNG2GNG3GNG4GNG5GNG7GNG11GNG12GNG13BSCL2G Protein-coupled Receptor KinaseAMP-activated Protein KinaseSmall GTPaseADP Ribosylation FactorRab (G-protein)Ras SubfamilyHRASKRASNeuroblastoma RAS Viral Oncogene HomologRho Family Of GTPasesADP Ribosylation FactorRan (gene)RHEBRap GTP-binding ProteinRRADCyclin-dependent Kinase Inhibitor ProteinCyclin-dependent KinasePhosphoinositide Phospholipase CPhospholipase CProtein KinaseSerine/threonine-specific Protein KinaseCasein KinaseCasein Kinase 1Casein Kinase 2EIF-2 KinaseEIF2AK3Glycogen Synthase KinaseGSK-3GSK3AGSK3BIκB KinaseCHUKIKK2IKBKGInterleukin-1 Receptor Associated KinaseIRAK1IRAK2IRAK3IRAK4Lim KinaseLIMK1LIMK2P21-activated KinasesPAK1PAK2PAK3PAK4Rho-associated Protein KinaseROCK1ROCK2Ribosomal S6 KinaseRPS6KA1Non-receptor Tyrosine KinaseZAP70PTK2PTK2BBruton's Tyrosine KinaseDual-specificity KinaseProtein PhosphataseProtein Serine/threonine PhosphataseProtein Phosphatase 2Protein Tyrosine PhosphataseProtein Tyrosine PhosphataseDual-specificity PhosphataseApoptosisTemplate:Apoptosis Signaling PathwayGTP-binding Protein RegulatorsTemplate:GTP-binding Protein RegulatorsATF6Signal Transducing Adaptor ProteinIκBαMUC4Olfactory Marker ProteinPhosphatidylethanolamine Binding ProteinEDARADDPRKCSHTemplate:Deficiencies Of Intracellular Signaling Peptides And ProteinsTemplate:Cell Cycle ProteinsTemplate Talk:Cell Cycle ProteinsCell CycleProteinCyclin ACyclin A1Cyclin A2Cyclin BCyclin B1Cyclin B2Cyclin DCyclin D1Cyclin D2Cyclin D3Cyclin ECyclin E1Cyclin E2Cyclin-dependent KinaseCyclin-dependent Kinase 1Cyclin-dependent Kinase 2Cyclin-dependent Kinase 3Cyclin-dependent Kinase 4Cyclin-dependent Kinase 5Cyclin-dependent Kinase 6Cyclin-dependent Kinase 7Cyclin-dependent Kinase 8Cyclin-dependent Kinase 9Cyclin-dependent Kinase 10CDK-activating KinaseCyclin-dependent Kinase Inhibitor ProteinCell CycleP14arfP16CDKN2BCDKN2CCDKN2DCell CycleP21CDKN1BCyclin-dependent Kinase Inhibitor 1CP53 P63 P73 FamilyP53TP63P73Cdk1Cdc25CDC42Cellular Apoptosis Susceptibility ProteinE2FMaturation Promoting FactorWee1CullinCUL7InterphaseG1 PhaseS PhaseG2 PhaseCell DivisionMitosisPreprophaseProphasePrometaphaseMetaphaseAnaphaseTelophaseCytokinesisCell Cycle CheckpointRestriction PointSpindle CheckpointPostreplication CheckpointApoptosisG0 PhaseMeiosisPublic DomainPfamInterProHelp:CategoryCategory:Cell CycleCategory:ProteinsDiscussion About Edits From This IP Address [n]A List Of Edits Made From This IP Address [y]View The Content Page [c]Discussion About The Content Page [t]Edit This Page [e]Visit The Main Page [z]Guides To Browsing WikipediaFeatured Content – The Best Of WikipediaFind Background Information On Current EventsLoad A Random Article [x]Guidance On How To Use And Edit WikipediaFind Out About WikipediaAbout The Project, What You Can Do, Where To Find ThingsA List Of Recent Changes In The Wiki [r]List Of All English Wikipedia Pages Containing Links To This Page [j]Recent Changes In Pages Linked From This Page [k]Upload Files [u]A List Of All Special Pages [q]Wikipedia:AboutWikipedia:General Disclaimer

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