PDBsum entry 1blx

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protein metals Protein-protein interface(s) links
Complex (inhibitor protein/kinase) PDB id
Protein chains
305 a.a. *
160 a.a. *
Waters ×294
* Residue conservation analysis
PDB id:
Name: Complex (inhibitor protein/kinase)
Title: P19ink4d/cdk6 complex
Structure: Cyclin-dependent kinase 6. Chain: a. Engineered: yes. P19ink4d. Chain: b. Engineered: yes
Source: Homo sapiens. Human. Organism_taxid: 9606. Cell_line: sf9. Expressed in: spodoptera frugiperda. Expression_system_taxid: 7108. Expression_system_cell_line: sf9. Mus musculus. House mouse.
Biol. unit: Dimer (from PQS)
1.90Å     R-factor:   0.200     R-free:   0.253
Authors: D.H.Brotherton,V.Dhanaraj,S.Wick,L.Brizuela,P.J.Domaille, E.Volyanik,X.Xu,E.Parisini,B.O.Smith,S.J.Archer,M.Serrano, S.L.Brenner,T.L.Blundell,E.D.Laue
Key ref:
D.H.Brotherton et al. (1998). Crystal structure of the complex of the cyclin D-dependent kinase Cdk6 bound to the cell-cycle inhibitor p19INK4d. Nature, 395, 244-250. PubMed id: 9751051 DOI: 10.1038/26164
21-Jul-98     Release date:   01-Jun-99    
Go to PROCHECK summary

Protein chain
Pfam   ArchSchema ?
Q00534  (CDK6_HUMAN) -  Cyclin-dependent kinase 6
326 a.a.
305 a.a.
Protein chain
Pfam   ArchSchema ?
Q60773  (CDN2D_MOUSE) -  Cyclin-dependent kinase 4 inhibitor D
166 a.a.
160 a.a.
Key:    PfamA domain  Secondary structure  CATH domain

 Enzyme reactions 
   Enzyme class: Chain A: E.C.  - Cyclin-dependent kinase.
[IntEnz]   [ExPASy]   [KEGG]   [BRENDA]
      Reaction: ATP + a protein = ADP + a phosphoprotein
+ protein
+ phosphoprotein
Molecule diagrams generated from .mol files obtained from the KEGG ftp site
 Gene Ontology (GO) functional annotation 
  GO annot!
  Cellular component     cell projection   10 terms 
  Biological process     type B pancreatic cell development   47 terms 
  Biochemical function     nucleotide binding     12 terms  


DOI no: 10.1038/26164 Nature 395:244-250 (1998)
PubMed id: 9751051  
Crystal structure of the complex of the cyclin D-dependent kinase Cdk6 bound to the cell-cycle inhibitor p19INK4d.
D.H.Brotherton, V.Dhanaraj, S.Wick, L.Brizuela, P.J.Domaille, E.Volyanik, X.Xu, E.Parisini, B.O.Smith, S.J.Archer, M.Serrano, S.L.Brenner, T.L.Blundell, E.D.Laue.
The crystal structure of the cyclin D-dependent kinase Cdk6 bound to the p19 INK4d protein has been determined at 1.9 A resolution. The results provide the first structural information for a cyclin D-dependent protein kinase and show how the INK4 family of CDK inhibitors bind. The structure indicates that the conformational changes induced by p19INK4d inhibit both productive binding of ATP and the cyclin-induced rearrangement of the kinase from an inactive to an active conformation. The structure also shows how binding of an INK4 inhibitor would prevent binding of p27Kip1, resulting in its redistribution to other CDKs. Identification of the critical residues involved in the interaction explains how mutations in Cdk4 and p16INK4a result in loss of kinase inhibition and cancer.
  Selected figure(s)  
Figure 2.
Figure 2 Three-dimensional structure of the p19^INK4d/Cdk6 complex. a, Stereo view of the C trace of the p19^INK4d/Cdk6 structure. Every tenth C atom is indicated by a solid ball. b, Schematic drawing of the same complex after rotation by 90 about the x axis. c, P19^INK4d and p27^Kip1 prevent each other binding to the CDK subunit in cyclin D/Cdk6. The structure in c is rotated 20 about the z axis compared to b. In a and b p19^INK4d is coloured yellow, apart from helix 3 (residues 46-50), which is red. The C-terminal domain of Cdk6 is coloured light blue, whereas the N-terminal domain, which undergoes extensive movement, is dark blue. In c, p19^INK4d is coloured yellow, Cdk6 is light/dark blue, p27^Kip1 is red and cyclin A is green. p27^Kip1 and cyclin A from the p27^Kip1/cyclin A/Cdk2 structure^40 were superimposed, as described in Fig. 5a, on p19^INK4d/Cdk6 (r.m.s. deviation was 0.46 ? over 70 residues). Figures 2 and 5a, b were produced using the programs MOLSCRIPT47 and RASTER3D^48.
Figure 5.
Figure 5 Comparison of ATP-binding sites in different kinases. a, Comparison of the structure of p19^INK4d/Cdk6 with those of Cdk2 (ref. 15) and Cdk2 from the cyclin A complex16,17. Cdk6 was superimposed on Cdk2 (r.m.s. deviation was 0.56 ? over 70 residues) and Cdk2 from the cyclin A complex (r.m.s. deviation was 0.49 ? over 70 residues) by aligning the C atoms in the -helices in the C-terminal domain of both proteins. Loop L5 and the linker (loop L7) (Fig. 3c), which anchor the N- and C-terminal domains together, are labelled. b, The same structures, showing only the N-terminal domain, illustrating the changes in position of the N-terminal -sheet and helix 1. In both a and b, the N-terminal domains and T-loop are coloured red (Cdk2), yellow (Cdk2 from the cyclin A complex) and blue (Cdk6); other regions of all three proteins are coloured grey. In b, helices 3 and 5 of cyclin A, which interact with PSTAIRE helix 1 in cyclin A/Cdk2 (refs 16, 17), are green. c, The ATP-binding site in Cdk6. Lys 29, His 100 and Asp 102, which together might inhibit ATP binding, are labelled, as are key active-site residues and the phosphate-binding loop L2. ATP, from the active Cdk2/cyclin A complex17, is superimposed on the structure with the N1 and N6 adenine nitrogens making conserved hydrogen bonds with the carbonyl of Glu 99 and amide of Val 101, respectively17,34. The structures of Cdk6 and ATP are yellow and green, respectively. Carbon, nitrogen, oxygen and phosphorus atoms are yellow, blue, red and yellow, respectively. The structure of Cdk2 from the cyclin A/Cdk2 complex17 is blue.
  The above figures are reprinted by permission from Macmillan Publishers Ltd: Nature (1998, 395, 244-250) copyright 1998.  
  Figures were selected by the author.  

Literature references that cite this PDB file's key reference

  PubMed id Reference
21077851 P.H.Cui, N.Petrovic, and M.Murray (2011).
The ω-3 epoxide of eicosapentaenoic acid inhibits endothelial cell proliferation by p38 MAP kinase activation and cyclin D1/CDK4 down-regulation.
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19470474 C.G.Mullighan, J.Zhang, R.C.Harvey, J.R.Collins-Underwood, B.A.Schulman, L.A.Phillips, S.K.Tasian, M.L.Loh, X.Su, W.Liu, M.Devidas, S.R.Atlas, I.M.Chen, R.J.Clifford, D.S.Gerhard, W.L.Carroll, G.H.Reaman, M.Smith, J.R.Downing, S.P.Hunger, and C.L.Willman (2009).
JAK mutations in high-risk childhood acute lymphoblastic leukemia.
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19260062 C.Kannengiesser, S.Brookes, A.G.del Arroyo, D.Pham, J.Bombled, M.Barrois, O.Mauffret, M.F.Avril, A.Chompret, G.M.Lenoir, A.Sarasin, G.Peters, B.Bressac-de Paillerets, F.Boitier, V.Bonadonna, J.M.Bonnetblanc, J.Chiesa, I.Coupier, S.Dalle, F.Grange, B.Guillot, P.Joly, C.Lasset, D.Leroux, J.M.Limacher, M.Longy, T.Martin-Denavit, L.Thomas, and P.Vabres (2009).
Functional, structural, and genetic evaluation of 20 CDKN2A germ line mutations identified in melanoma-prone families or patients.
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Biological regulation via ankyrin repeat folding.
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19237565 P.J.Day, A.Cleasby, I.J.Tickle, M.O'Reilly, J.E.Coyle, F.P.Holding, R.L.McMenamin, J.Yon, R.Chopra, C.Lengauer, and H.Jhoti (2009).
Crystal structure of human CDK4 in complex with a D-type cyclin.
  Proc Natl Acad Sci U S A, 106, 4166-4170.
PDB codes: 2w96 2w99 2w9f 2w9z
18305166 C.Löw, U.Weininger, P.Neumann, M.Klepsch, H.Lilie, M.T.Stubbs, and J.Balbach (2008).
Structural insights into an equilibrium folding intermediate of an archaeal ankyrin repeat protein.
  Proc Natl Acad Sci U S A, 105, 3779-3784.
PDB code: 2rfm
17502107 A.Schweizer, H.Roschitzki-Voser, P.Amstutz, C.Briand, M.Gulotti-Georgieva, E.Prenosil, H.K.Binz, G.Capitani, A.Baici, A.Plückthun, and M.G.Grütter (2007).
Inhibition of caspase-2 by a designed ankyrin repeat protein: specificity, structure, and inhibition mechanism.
  Structure, 15, 625-636.
PDB code: 2p2c
17373709 G.Lolli, and L.N.Johnson (2007).
Recognition of Cdk2 by Cdk7.
  Proteins, 67, 1048-1059.
PDB code: 2hic
17342741 G.M.Buchold, P.L.Magyar, R.Arumugam, M.M.Lee, and D.A.O'Brien (2007).
p19Ink4d and p18Ink4c cyclin-dependent kinase inhibitors in the male reproductive axis.
  Mol Reprod Dev, 74, 997.  
17625456 M.Széll, K.Balogh, A.Dobozy, L.Kemény, and J.Oláh (2007).
First detection of the melanoma-predisposing proline-48-threonine mutation of p16 in Hungarians: was there a common founder either in Italy or in Hungary?
  Melanoma Res, 17, 251-254.  
16953232 P.P.Joshi, M.V.Kulkarni, B.K.Yu, K.R.Smith, D.L.Norton, W.Veelen, J.W.Höppener, and D.S.Franklin (2007).
Simultaneous downregulation of CDK inhibitors p18(Ink4c) and p27(Kip1) is required for MEN2A-RET-mediated mitogenesis.
  Oncogene, 26, 554-570.  
16493461 C.Aubry, A.J.Wilson, P.R.Jenkins, S.Mahale, B.Chaudhuri, J.D.Maréchal, and M.J.Sutcliffe (2006).
Design, synthesis and biological activity of new CDK4-specific inhibitors, based on fascaplysin.
  Org Biomol Chem, 4, 787-801.  
16542156 J.Joy, N.Nalabothula, M.Ghosh, O.Popp, M.Jochum, W.Machleidt, S.Gil-Parrado, and T.A.Holak (2006).
Identification of calpain cleavage sites in the G1 cyclin-dependent kinase inhibitor p19(INK4d).
  Biol Chem, 387, 329-335.  
16584130 J.Sridhar, N.Akula, and N.Pattabiraman (2006).
Selectivity and potency of cyclin-dependent kinase inhibitors.
  AAPS J, 8, E204-E221.  
16084385 A.Kohl, P.Amstutz, P.Parizek, H.K.Binz, C.Briand, G.Capitani, P.Forrer, A.Plückthun, and M.G.Grütter (2005).
Allosteric inhibition of aminoglycoside phosphotransferase by a designed ankyrin repeat protein.
  Structure, 13, 1131-1141.
PDB code: 2bkk
15855166 A.Schulte, N.Czudnochowski, M.Barboric, A.Schönichen, D.Blazek, B.M.Peterlin, and M.Geyer (2005).
Identification of a cyclin T-binding domain in Hexim1 and biochemical analysis of its binding competition with HIV-1 Tat.
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15893667 M.Lei, M.A.Robinson, and S.C.Harrison (2005).
The active conformation of the PAK1 kinase domain.
  Structure, 13, 769-778.
PDB codes: 1yhv 1yhw
15851475 P.Amstutz, H.K.Binz, P.Parizek, M.T.Stumpp, A.Kohl, M.G.Grütter, P.Forrer, and A.Plückthun (2005).
Intracellular kinase inhibitors selected from combinatorial libraries of designed ankyrin repeat proteins.
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15664993 P.Kaldis (2005).
The N-terminal peptide of the Kaposi's sarcoma-associated herpesvirus (KSHV)-cyclin determines substrate specificity.
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15838515 X.H.Pei, and Y.Xiong (2005).
Biochemical and cellular mechanisms of mammalian CDK inhibitors: a few unresolved issues.
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14583612 B.Padmanabhan, N.Adachi, K.Kataoka, and M.Horikoshi (2004).
Crystal structure of the homolog of the oncoprotein gankyrin, an interactor of Rb and CDK4/6.
  J Biol Chem, 279, 1546-1552.
PDB code: 1ixv
15304098 G.Yang, K.B.Niendorf, and H.Tsao (2004).
A novel methionine-53-valine mutation of p16 in a hereditary melanoma kindred.
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15611164 J.W.Loar, R.M.Seiser, A.E.Sundberg, H.J.Sagerson, N.Ilias, P.Zobel-Thropp, E.A.Craig, and D.E.Lycan (2004).
Genetic and biochemical interactions among Yar1, Ltv1 and Rps3 define novel links between environmental stress and ribosome biogenesis in Saccharomyces cerevisiae.
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14570903 M.Y.Niv, H.Rubin, J.Cohen, L.Tsirulnikov, T.Licht, A.Peretzman-Shemer, E.Cna'an, A.Tartakovsky, I.Stein, S.Albeck, I.Weinstein, M.Goldenberg-Furmanov, D.Tobi, E.Cohen, M.Laster, S.A.Ben-Sasson, and H.Reuveni (2004).
Sequence-based design of kinase inhibitors applicable for therapeutics and target identification.
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14760739 P.Forrer, H.K.Binz, M.T.Stumpp, and A.Plückthun (2004).
Consensus design of repeat proteins.
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14701845 Q.Zhao, F.Boschelli, A.J.Caplan, and K.T.Arndt (2004).
Identification of a conserved sequence motif that promotes Cdc37 and cyclin D1 binding to Cdk4.
  J Biol Chem, 279, 12560-12564.  
15141161 Y.S.Heo, S.K.Kim, C.I.Seo, Y.K.Kim, B.J.Sung, H.S.Lee, J.I.Lee, S.Y.Park, J.H.Kim, K.Y.Hwang, Y.L.Hyun, Y.H.Jeon, S.Ro, J.M.Cho, T.G.Lee, and C.H.Yang (2004).
Structural basis for the selective inhibition of JNK1 by the scaffolding protein JIP1 and SP600125.
  EMBO J, 23, 2185-2195.
PDB codes: 1ukh 1uki
14621993 J.Li, S.H.Joo, and M.D.Tsai (2003).
An NF-kappaB-specific inhibitor, IkappaBalpha, binds to and inhibits cyclin-dependent kinase 4.
  Biochemistry, 42, 13476-13483.  
12517341 K.S.Tang, A.R.Fersht, and L.S.Itzhaki (2003).
Sequential unfolding of ankyrin repeats in tumor suppressor p16.
  Structure, 11, 67-73.  
12686541 S.Malek, D.B.Huang, T.Huxford, S.Ghosh, and G.Ghosh (2003).
X-ray crystal structure of an IkappaBbeta x NF-kappaB p65 homodimer complex.
  J Biol Chem, 278, 23094-23100.
PDB codes: 1k3z 1oy3
12789688 T.Honma (2003).
Recent advances in de novo design strategy for practical lead identification.
  Med Res Rev, 23, 606-632.  
  12537555 A.Krupa, and N.Srinivasan (2002).
The repertoire of protein kinases encoded in the draft version of the human genome: atypical variations and uncommon domain combinations.
  Genome Biol, 3, RESEARCH0066.  
12086621 C.I.Chang, B.E.Xu, R.Akella, M.H.Cobb, and E.J.Goldsmith (2002).
Crystal structures of MAP kinase p38 complexed to the docking sites on its nuclear substrate MEF2A and activator MKK3b.
  Mol Cell, 9, 1241-1249.
PDB codes: 1lew 1lez
11823456 F.L.Robinson, A.W.Whitehurst, M.Raman, and M.H.Cobb (2002).
Identification of novel point mutations in ERK2 that selectively disrupt binding to MEK1.
  J Biol Chem, 277, 14844-14852.  
11930009 L.J.Raffini, D.J.Slater, E.F.Rappaport, L.Lo Nigro, N.K.Cheung, J.A.Biegel, P.C.Nowell, B.J.Lange, and C.A.Felix (2002).
Panhandle and reverse-panhandle PCR enable cloning of der(11) and der(other) genomic breakpoint junctions of MLL translocations and identify complex translocation of MLL, AF-4, and CDK6.
  Proc Natl Acad Sci U S A, 99, 4568-4573.  
11959850 M.C.Morris, C.Gondeau, J.A.Tainer, and G.Divita (2002).
Kinetic mechanism of activation of the Cdk2/cyclin A complex. Key role of the C-lobe of the Cdk.
  J Biol Chem, 277, 23847-23853.  
12456646 P.Michaely, D.R.Tomchick, M.Machius, and R.G.Anderson (2002).
Crystal structure of a 12 ANK repeat stack from human ankyrinR.
  EMBO J, 21, 6387-6396.
PDB code: 1n11
12191603 R.A.Engh, and D.Bossemeyer (2002).
Structural aspects of protein kinase control-role of conformational flexibility.
  Pharmacol Ther, 93, 99.  
12370184 R.N.Venkataramani, T.K.MacLachlan, X.Chai, W.S.El-Deiry, and R.Marmorstein (2002).
Structure-based design of p18INK4c proteins with increased thermodynamic stability and cell cycle inhibitory activity.
  J Biol Chem, 277, 48827-48833.
PDB codes: 1mx2 1mx4 1mx6
11960696 S.Ortega, M.Malumbres, and M.Barbacid (2002).
Cyclin D-dependent kinases, INK4 inhibitors and cancer.
  Biochim Biophys Acta, 1602, 73-87.  
12417717 T.J.Huot, J.Rowe, M.Harland, S.Drayton, S.Brookes, C.Gooptu, P.Purkis, M.Fried, V.Bataille, E.Hara, J.Newton-Bishop, and G.Peters (2002).
Biallelic mutations in p16(INK4a) confer resistance to Ras- and Ets-induced senescence in human diploid fibroblasts.
  Mol Cell Biol, 22, 8135-8143.  
11828325 U.Schulze-Gahmen, and S.H.Kim (2002).
Structural basis for CDK6 activation by a virus-encoded cyclin.
  Nat Struct Biol, 9, 177-181.
PDB code: 1jow
11590016 C.Kleanthous, and D.Walker (2001).
Immunity proteins: enzyme inhibitors that avoid the active site.
  Trends Biochem Sci, 26, 624-631.  
11556834 G.Della Torre, B.Pasini, S.Frigerio, R.Donghi, D.Rovini, D.Delia, G.Peters, T.J.Huot, G.Bianchi-Scarra, F.Lantieri, M.Rodolfo, G.Parmiani, and M.A.Pierotti (2001).
CDKN2A and CDK4 mutation analysis in Italian melanoma-prone families: functional characterization of a novel CDKN2A germ line mutation.
  Br J Cancer, 85, 836-844.  
11479422 G.Walker, and N.Hayward (2001).
No evidence of a role for activating CDK2 mutations in melanoma.
  Melanoma Res, 11, 343-348.  
11398194 J.Chan, E.S.Robinson, J.Atencio, Z.Wang, S.Kazianis, L.D.Coletta, R.S.Nairn, and J.R.McCarrey (2001).
Characterization of the CDKN2A and ARF genes in UV-induced melanocytic hyperplasias and melanomas of an opossum (Monodelphis domestica).
  Mol Carcinog, 31, 16-26.  
11574463 K.Niefind, B.Guerra, I.Ermakowa, and O.G.Issinger (2001).
Crystal structure of human protein kinase CK2: insights into basic properties of the CK2 holoenzyme.
  EMBO J, 20, 5320-5331.
PDB code: 1jwh
11463838 N.Nekrep, M.Geyer, N.Jabrane-Ferrat, and B.M.Peterlin (2001).
Analysis of ankyrin repeats reveals how a single point mutation in RFXANK results in bare lymphocyte syndrome.
  Mol Cell Biol, 21, 5566-5576.  
  11739795 P.Kaldis, P.M.Ojala, L.Tong, T.P.Mäkelä, and M.J.Solomon (2001).
CAK-independent activation of CDK6 by a viral cyclin.
  Mol Biol Cell, 12, 3987-3999.  
11526321 U.Schulze-Gahmen, and S.H.Kim (2001).
Crystallization of a complex between human CDK6 and a virus-encoded cyclin is critically dependent on the addition of small charged organic molecules.
  Acta Crystallogr D Biol Crystallogr, 57, 1287-1289.  
11567160 Y.W.Chen (2001).
Solution solution: using NMR models for molecular replacement.
  Acta Crystallogr D Biol Crystallogr, 57, 1457-1461.  
  10892805 C.Yuan, T.L.Selby, J.Li, I.J.Byeon, and M.D.Tsai (2000).
Tumor suppressor INK4: refinement of p16INK4A structure and determination of p15INK4B structure by comparative modeling and NMR data.
  Protein Sci, 9, 1120-1128.
PDB code: 1dc2
10594039 F.Zindy, J.van Deursen, G.Grosveld, C.J.Sherr, and M.F.Roussel (2000).
INK4d-deficient mice are fertile despite testicular atrophy.
  Mol Cell Biol, 20, 372-378.  
10651629 J.Li, M.J.Poi, D.Qin, T.L.Selby, I.J.Byeon, and M.D.Tsai (2000).
Tumor suppressor INK4: quantitative structure-function analyses of p18INK4C as an inhibitor of cyclin-dependent kinase 4.
  Biochemistry, 39, 649-657.  
10734227 M.Thullberg, J.Bartkova, S.Khan, K.Hansen, L.Rönnstrand, J.Lukas, M.Strauss, and J.Bartek (2000).
Distinct versus redundant properties among members of the INK4 family of cyclin-dependent kinase inhibitors.
  FEBS Lett, 470, 161-166.  
11124804 P.D.Jeffrey, L.Tong, and N.P.Pavletich (2000).
Structural basis of inhibition of CDK-cyclin complexes by INK4 inhibitors.
  Genes Dev, 14, 3115-3125.
PDB code: 1g3n
11063931 S.V.Ekholm, and S.I.Reed (2000).
Regulation of G(1) cyclin-dependent kinases in the mammalian cell cycle.
  Curr Opin Cell Biol, 12, 676-684.  
11076027 T.L.Blundell, D.F.Burke, D.Chirgadze, V.Dhanaraj, M.Hyvönen, C.A.Innis, E.Parisini, L.Pellegrini, M.Sayed, and B.L.Sibanda (2000).
Protein-protein interactions in receptor activation and intracellular signalling.
  Biol Chem, 381, 955-959.  
11080645 Y.W.Chen, E.J.Dodson, and G.J.Kleywegt (2000).
Does NMR mean "not for molecular replacement"? Using NMR-based search models to solve protein crystal structures.
  Structure, 8, R213-R220.  
  10454538 B.J.Warner, S.W.Blain, J.Seoane, and J.Massagué (1999).
Myc downregulation by transforming growth factor beta required for activation of the p15(Ink4b) G(1) arrest pathway.
  Mol Cell Biol, 19, 5913-5922.  
10203759 C.Swanton, G.L.Card, D.Mann, N.McDonald, and N.Jones (1999).
Overcoming inhibitions: subversion of CKI function by viral cyclins.
  Trends Biochem Sci, 24, 116-120.  
10064597 H.Yasukawa, H.Misawa, H.Sakamoto, M.Masuhara, A.Sasaki, T.Wakioka, S.Ohtsuka, T.Imaizumi, T.Matsuda, J.N.Ihle, and A.Yoshimura (1999).
The JAK-binding protein JAB inhibits Janus tyrosine kinase activity through binding in the activation loop.
  EMBO J, 18, 1309-1320.  
10607671 J.A.Endicott, M.E.Noble, and J.A.Tucker (1999).
Cyclin-dependent kinases: inhibition and substrate recognition.
  Curr Opin Struct Biol, 9, 738-744.  
10074345 J.Li, I.J.Byeon, K.Ericson, M.J.Poi, P.O'Maille, T.Selby, and M.D.Tsai (1999).
Tumor suppressor INK4: determination of the solution structure of p18INK4C and demonstration of the functional significance of loops in p18INK4C and p16INK4A.
  Biochemistry, 38, 2930-2940.
PDB code: 1bu9
10361086 M.R.Groves, and D.Barford (1999).
Topological characteristics of helical repeat proteins.
  Curr Opin Struct Biol, 9, 383-389.  
9989501 M.R.Groves, N.Hanlon, P.Turowski, B.A.Hemmings, and D.Barford (1999).
The structure of the protein phosphatase 2A PR65/A subunit reveals the conformation of its 15 tandemly repeated HEAT motifs.
  Cell, 96, 99.
PDB code: 1b3u
10580009 M.Sugimoto, T.Nakamura, N.Ohtani, L.Hampson, I.N.Hampson, A.Shimamoto, Y.Furuichi, K.Okumura, S.Niwa, Y.Taya, and E.Hara (1999).
Regulation of CDK4 activity by a novel CDK4-binding protein, p34(SEI-1).
  Genes Dev, 13, 3027-3033.  
10431175 S.G.Sedgwick, and S.J.Smerdon (1999).
The ankyrin repeat: a diversity of interactions on a common structural framework.
  Trends Biochem Sci, 24, 311-316.  
11232330 T.Huxford, S.Malek, and G.Ghosh (1999).
Structure and mechanism in NF-kappa B/I kappa B signaling.
  Cold Spring Harb Symp Quant Biol, 64, 533-540.  
10368294 U.Schulze-Gahmen, J.U.Jung, and S.H.Kim (1999).
Crystal structure of a viral cyclin, a positive regulator of cyclin-dependent kinase 6.
  Structure, 7, 245-254.
PDB code: 1bu2
The most recent references are shown first. Citation data come partly from CiteXplore and partly from an automated harvesting procedure. Note that this is likely to be only a partial list as not all journals are covered by either method. However, we are continually building up the citation data so more and more references will be included with time. Where a reference describes a PDB structure, the PDB codes are shown on the right.