MAPK Nomenclature

Mitogen-activated protein kinase cascades in plants: A new nomenclature

Mitogen-activated protein (MAP) kinase cascades are universal signal transduction modules in eukaryotes, including yeasts, animals and plants. These protein phosphorylation cascades link extracellular stimuli to a wide range of cellular responses. In plants, MAP kinase cascades are involved in responses to various biotic/abiotic stresses, hormonal responses, cell division, and developmental processes. Completion of the Arabidopsis genome-sequencing project has revealed the existence of 20 MAP kinases (MAPKs), 10 MAPK kinases (MAPKKs) and 60 MAPKK kinases (MAPKKKs). Here, we propose a simplified nomenclature for Arabidopsis MAPKs and MAPKKs that is based on genomics information for standard annotation of these gene families in Arabidopsis and may also serve as a basis for standard annotation of these gene families in all plants.

Trends in Plant Science, 2002

Kazuya Ichimura, Guillaume Tena, Yves Henry, Shuqun Zhang, Heribert Hirt, Brian E Ellis, Peter C. Morris, Cathal Wilson, Anthony Champion, Roger W. Innes, Jen Sheen, Joseph R. Ecker, Dierk Scheel, Daniel F. Klessig, Yasunori Machida, John Mundy, Yuko Ohashi, Martin Kreis, Erwin Heberle-Bors, John C. Walker and Kazuo Shinozaki

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Kazuya Ichimura, Kazuo Shinozaki
Laboratory of Plant Molecular Biology, RIKEN Tsukuba Institute, Koyadai 3-1-1, Tsukuba, Ibaraki 305-0074, Japan

Guillaume Tena, Jen Sheen
Department of Genetics, Harvard Medical School, Massachusetts General Hospital, Boston, MA 02114, USA

Yves Henry, Anthony Champion, Martin Kreis
Institut de Biotechnologie des Plantes (IBP), Laboratoire de Biologie du Developpement des Plantes, Batiment 630, UMR 8618, Universite de Paris-Sud, F-91405, Orsay Cedex, France

Shuqun Zhang
Department of Biochemistry, University of Missouri-Columbia, 117 Schweitzer Hall, Columbia, MO 65211, USA

Erwin Heberle-Bors, Heribert Hirt, Cathal Wilson
Institute of Microbiology and Genetics, Vienna Biocenter, University of Vienna, Dr. Bohrgasse 9, 1030 Vienna, Austria

Brian E. Ellis
Biotechnology Laboratory, University of British Columbia, 6174 University Blvd., Vancouver, British Columbia, Canada V6T 1Z3

Peter C. Morris
Department of Biological Sciences, Heriot-Watt University, Riccarton, Edinburgh EH14 4AS, UK

Roger W. Innes
Department of Biology, Indiana University, Bloomington, IN 47405-3700, USA

Joseph R. Ecker
Plant Biology Laboratory, The Salk Institute for Biological Studies, La Jolla, California 92037, USA

Dierk Scheel
Institute of Plant Biochemistry, Department of Stress and Developmental Biology, Weinberg 3, D-06120 Halle (Saale), Germany

Daniel F. Klessig
Boyce Thompson Institute for Plant Research, Tower Road, Ithaca, NY 14853, USA

Yasunori Machida
Laboratory of Plant Development, Division of Biological Science, Graduate School of Science, Nagoya University, Chikusa-ku, Nagoya 464-8602, Japan

John Mundy
Institute of Molecular Biology, Copenhagen University, ster Farimagsgade 2A, 1353 Copenhagen K, Denmark

John C. Walker
Division of Biological Sciences and Department of Biochemistry, University of Missouri-Columbia, Columbia, MO 65211, USA

Yuko Ohashi
National Institute of Agrobiological Sciences, Tsukuba, Ibaraki 305-8602, Japan

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Fig.1 Phylogenetic tree and domain structure of plant mitogen activated protein kinases (MPKs). To identify the species of origin for each mitogen activated kinase (MPK), a species acronym is included before the protein name.

Fig.2 Phylogenetic tree and domain structure of plant mitogen activated protein kinase kinases (MKKs)

Fig.3 Phylogenetic tree and domain structure of putative plant mitogen activated protein kinase kinase kinases (MAPKKKs).

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Table 1, Arabidopsis MAP kinases
Protein Name Gene code BAC clone Protein length (aa) Molecular Weight Genbank protein accession no. PlantsP ID T-loop Subgroup Reference
MPK1 At1g10210 F14N23 370 42645 BAA03535 21421 TEY C1 1
MPK2 At1g59580 T30E16 376 43125 BAA03536 21386 TEY C1 1
MPK3 At3g45640 F9K21 370 42717 BAA04866 21732 TEY A3 1
MPK4 At4g01370 F2N1 376 42852 BAA04867 21274 TEY B4 1
MPK5 At4g11330 F8L21 373 42882 BAA04868 21820 TEY B4 1
MPK6 At2g43790 F18O19 395 45058 BAA04869 21513 TEY A6 1
MPK7 At2g18170 F8D23 368 42299 BAA04870 21194 TEY C7 1
MPK8 At1g18150 T10O22 589 66232 AB038693 21494 TDY D8 2
MPK9 At3g18040 MRC8 613 70128 AB038694 21212 TDY D8 2
MPK10 At3g59790 F24G16 393 45175 CAB75798 21786 TEY A10 unpublished
MPK11 At1g01560 F22L4 369 42476 AAF81314 NR* TEY B4 unpublished
MPK12 At2g46070 T3F17 406 46137 AAC62906 21528 TEY B4 unpublished
MPK13 At1g07880 F24B9 363 42195 AAF75067 21381 TEY B13 unpublished
MPK14 At4g36450 ATAP22 361 41976 CAB16812 NR* TEY C7 unpublished
MPK15 At1g73670 F25P22 576 65251 AAG52072 21099 TDY D8 unpublished
MPK16 At5g19010 T16G12 567 64912 NP_197402 21303 TDY D8 unpublished
MPK17 At2g01450 F2I9 516 59038 AAC67338 21188 TDY D8 unpublished
MPK18 At1g53510 F22G10 603 67951 AAG51978 21 TDY D18 unpublished
MPK19 At3g14720 MIE1 586 65904 NP_188090 21700 TDY D18 unpublished
MPK20 At2g42880 F7D19 694 67275 AAD21721 21191 TDY D18 unpublished
* not registered                  
Table 2, Arabidopsis MAP kinase kinases
Name Gene code BAC clone Protein length (aa) Molecular Weight Genbank protein accession no. PlantsP ID Subgroup Reference
MKK1* At4g26070 F20B18 354 39210 AAB97145 21874 A1 23
MKK2 At4g29810 F27B13 363 39849 BAA28828 21892 A1 24
MKK3 At5g40440 MPO12 520 57530 BAA28829 21997 B3 25
MKK4 At1g51660 F19C24 366 40117 BAA28830 21493 C4 25, 26
MKK5 At3g21220 MXL8 348 38329 BAA28831 21701 C4 25-27
MKK6 At5g56580 MIK19 356 39837 NP_200469 22034 A6 unpublished
MKK7 At1g18350 F15H18 307 34272 AAF25995 21030 D7 unpublished
MKK8 At3g06230 F28L1 293 32521 AAF30316 21239 D7 unpublished
MKK9 At1g73500 T9L24 310 34347 AAG30984 21164 D7 unpublished
MKK10 At1g32320 F27G20 305 34023 AAF81327 21506 D7 unpublished
* Kindly renamed by Morris, PC from MEK1
Table 3, Arabidopsis MAP kinase kinase kinases
Protein name Gene code BAC clone ID Protein length Molecular weight Protein Genbank accession no. PlantsP ID Group Reference
  At1g04700 T1G11 1042 117987 AAB80620 21068 B4 unpublished
EDR1 =AtMAP3Kd3 At1g08720 F22O13 1015 113006 AAG31143 21410 B3 35,36
ANP1 At1g09000 F7G19 666 73453 BAA21854 21067 A3 37
  At1g14000 F7A19 438 49336 AAD39286 21080 C1 unpublished
  At1g16270 F3O9 1147 126730 AAD34679 21431 B4 unpublished
  At1g18160 T10O22 990 107871 AAF78373 21128 B3 unpublished
AtMAP3Ka At1g53570 F22G10 608 66334 CAA08994 21374 A2 35
ANP2 At1g54960 F14C21 585 64704 BAA21856 26217 A3 37
  At1g62400 F24O1 345 39197 AAF70839 21504 C5 unpublished
  At1g63700 F24D7 883 95961 AAG52426 21456 A2 unpublished
  At1g67890 T23K23 738 82224 AAG52018 21477 B2 unpublished
  At1g73660 F25P22 1030 112184 AAG52069 21462 B3 unpublished
  At1g79570 T8K14 1248 137192 AAD30219 21429 B4 unpublished
  At2g17700 T17A5 546 61508 AAB80785 NR* C2 unpublished
  At2g24360 T28I24 407 45774 AAD18109 21646 C6 unpublished
AtMAP3Kq1 At2g31010 F7F1 375 42589 AAC20735 21612 B1 35
  At2g31800 F20M17 489 55315 AAD32292 21523 C1 unpublished
  At2g35050 F19I3 1257 139666 AAC12844 21586 B4 unpublished
AtMAP3Kq2 At2g42630 F14N22 357 40198 AAD22991 21521 B1 35
  At2g43850 F18O19 477 54150 AAF18591 NR* C1 unpublished
  At3g01490 F4P13 411 46053 AAF01534 21661 C7 unpublished
ANP3 At3g06030 F24F17 651 71659 BAA21857 21692 A3 37
  At3g06620 F5E6 773 86058 AAG51332 21691 B2 unpublished
  At3g06630 F5E6 671 75178 AAG51330 21690 B2 unpublished
  At3g06640 F5E6 763 85597 AAG51328 21689 B2 unpublished
AtMAP3Ke2 At3g07980 F17A17 1367 151113 AAF21208 21681 A4 38
AtMAP3Ke1 At3g13530 MRP15 1368 1368 CAA12272 21714 A4 38
  At3g22750 MWI23 378 42717 BAB01250 21206 C7 unpublished
  At3g24720 K7P8 297 33300 NP_189116 21224 B4 unpublished
ATN1 At3g27560 MMJ24 356 40098 CAA63387 21214 C3 39
  At3g46920 T6H20 1171 128958 CAB51173 21747 B4 unpublished
  At3g46930 T6H20 475 53835 CAB51172 21748 C5 unpublished
  At3g50720 T3A5 377 43109 CAB62441 21758 C4 unpublished
ATN1-like At3g50730 T3A5 370 42331 CAB62442 21759 C4 unpublished
  At3g58640 F14P22 816 90836 CAB68202 21260 B1 unpublished
  At3g58760 T20N10 456 51595 CAB88293 21261 C1 unpublished
  At3g59830 F24G16 476 53904 CAB75802 21266 C1 unpublished
AtMRK1 At3g63260 F16M2 391 42578 BAA22079 21789 C7 40
AtMEKK3 At4g08470 T15F16 560 62298 AAC28187 21812 A1 41
AtMEKK2 At4g08480 T15F16 773 84953 AAC28188 21813 A1 41
AtMEKK1 At4g08500 T15F16 608 66014 BAA09057 11930 A1 41, 42
AtMEKK4 At4g12020 F16J13 1895 210307 CAB40943 21829 A1 40
  At4g14780 dl3430w 364 40727 CAB10257 21929 C7 unpublished
  At4g18950 F13C5 421 48174 CAA16752 21836 C1 unpublished
AtMAP3Kd4 At4g23050 F7H19 736 82276 CAB79260 21849 B2 35
AtMAP3Kd5 At4g24480 T22A6 963 107984 CAB45083 21870 B3 35
  At4g31170 F6E21 412 46077 CAB79835 21896 C6 unpublished
  At4g35780 F8D20 553 62633 CAA20048 21916 C2 unpublished
  At4g38470 F20M13 545 61965 CAB37503 21920 C2 unpublished
  At5g01850 T20L15 356 40154 CAB82755 21942 C3 unpublished
CTR1 At5g03730 F17C15 821 90290 AAA32779 21949 B3 43
AtMAP3Kd1 At5g11850 F14F18 886 98264 CAA74591 21971 B3 35
  At5g40540 MNF13 353 39661 NP_198870 21998 C3 unpublished
AtMAP3Kh3 At5g49470 K7J8 730 81159 BAB10760 22014 B2 35
  At5g50000 MPF21 385 42731 BAB10286 22019 C7 unpublished
  At5g50180 K6A12 346 38971 BAB09389 22020 C3 unpublished
  At5g57610 MUA2 1054 117410 BAB08796 22036 B4 unpublished
  At5g58950 K19M22 561 62979 BAB09638 21346 C5 unpublished
  At5g66710 MSN2 405 46018 BAA97277 22062 C4 unpublished
AtMAP3Kg At5g66850 MUD21 376 42148 AF360242 22064 A2 35
* not registered                

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1. Mizoguchi, T. et al.(1993) ATMPKs - a gene family of plant MAP kinases in Arabidopsis thaliana. FEBS Let. 336, 440-444

2. Mizoguchi, T. et al. (1997) Environmental stress response in plants - the role of mitogen-activated protein kinases. Trends Biotech. 15, 15-19

3. Huttly, A.K. and Phillips, A.L. (1995) Gibberellin-regulated expression in oat aleurone cells of two kinases that show homology to MAPkinase and a ribosomal protein kinase. Plant Mol.Biol. 27, 1043-1052

4. Shin, H.J. et al. (2001) Molecular cloning and cultivar specific expression of MAP kinases from Capsicum annuum. Mol. Cell 11, 48-54

5. To, K.Y. et al. (1999) Cloning of a sweet potato leaf cDNA encoding mitogen-activated protein kinase (PGR99-135). Plant Physiol. 121, 312

6. Jonak, C. et al. (1996) Stress signaling in plants: a mitogen-activated protein kinase pathway is activated by cold and drought. Proc. Natl. Acad. Sci. U. S. A. 93, 11274-11279

7. Munnik, T. et al. (1999) Distinct osmo-sensing protein kinase pathways are involved in signalling moderate and severe hyper-osmotic stress. Plant J.20, 381-388

8. Jonak, C. et al. (1995) MMK2, a novel alfalfa MAP kinase, specifically complements the yeast MPK1 function. Mol. Gen. Genet. 248, 686-694

9. Bgre, L. et al. (1999) A MAP kinase is activated late in plant mitosis and becomes localized to the plane of cell division. Plant Cell 11, 101-113

10. Schoenbeck, M.A. et al. (1999) The alfalfa (Medicago sativa) TDY1 gene encodes a mitogen-activated protein kinase homolog. Mol Plant Microbe Interact 12, 882-893

11. Seo, S. et al. (1995) Tobacco MAP kinase: a possible mediator in wound signal transduction pathways. Science 270, 1988-1992

12. Wilson, C. et al. (1995) Molecular cloning, functional expression in Escherichia coli, and characterization of multiple mitogen-activated-protein kinases from tobacco. Eur. J. Biochem.233, 249-257

13. Zhang, S. and Klessig, D.F. (1997) Salicylic acid activates a 48-kD MAP kinase in tobacco. Plant Cell 9,809-824

14. Calderini, O. et al. (1998) A cell cycle regulated MAP kinase with a possible role in cytokinesis in tobacco cells. J. Cell Sci. 111, 3091-3100

15. Wilson, C. et al. (1993) Isolation and characterization of a tobacco cDNA clone encoding a putative MAP kinase. Plant Mol. Biol. 23, 543-551

16. He, C. et al. (1999) BWMK1, a novel MAP kinase induced by fungal infection and mechanical wounding in rice. Mol. Plant Microbe Interact.12, 1064-1073

17. Ligterink, W. et al.(1997) Receptor-mediated activation of a MAP kinase in pathogen defense of plants. Science276, 2054-2057

18. Decroocq-Ferrant, V. et al. (1995) A homologue of the MAP/ERK family of protein kinase genes is expressed in vegetative and in female reproductive organs of Petunia hybrida. Plant Mol.Biol.27, 339-350

19. Marcote, M.J. and Carbonell, J. (2000) Transient expression of a pea MAP kinase gene induced by gibberellic acid and 6-benzyladenine in unpollinated pea ovaries. Plant Mol. Biol. 44, 177-186

20. Stafstrom, J.P. et al. (1993) Molecular cloning and expression of a MAP kinase homologue from pea. Plant Mol. Biol. 22, 83-90

21. Takezawa, D. (1999) Elicitor- and A23187-induced expression of WCK-1, a gene encoding mitogen-activated protein kinase in wheat. Plant Mol. Biol. 40, 921-933

22. Berberich, T. et al. (1999) Involvement of a MAP kinase, ZmMPK5, in senescence and recovery from low-temperature stress in maize. Mol. Gen. Genet. 262, 534-542

23. Morris, P.C. et al. (1997) Cloning and characterisation of MEK1, an Arabidopsis gene encoding a homologue of MAP kinase kinase. Plant Mol. Biol. 35, 1057-1064

24. Ichimura, K. et al. (1998) Isolation of ATMEKK1 (a MAP kinase kinase Kinase) - Interacting proteins and analysis of a MAP kinase cascade in Arabidopsis. Biochem. Biophys. Res. Comm. 253, 532-543

25. Ichimura, K. et al. (1998) Molecular cloning and characterization of three cDNAs encoding putative mitogen-activated protein kinase kinases (MAPKKs) in Arabidopsis thaliana. DNA Res. 5, 341-348

26. Ren, al. (2001) Cell death mediated by MAPK is associated with hydrogen peroxide production in Arabidopsis. J. Biol. Chem. 227, 559-565

27. Hamal, A. et al. (1999) Molecular characterisation and expression of an Arabidopsis thaliana L. MAP kinase kinase cDNA, AtMAP2Ka. Plant Sci. 140, 41-52

28. Hackett, R.M. et al. (1998) A tomato MAP kinase kinase gene (Accession No. AJ000728) differentially regulated during fruit development, leaf senescence and wounding (PGR98-151). Plant Physiol. 117, 1526

29. Kiegerl, S. et al.(2000) SIMKK, a mitogen-activated protein kinase (MAPK) kinase, is a specific activator of the salt stress-induced MAPK, SIMK. Plant Cell 12, 2247-2258

30. Calderini, O. et al. (2001) A novel tobacco mitogen-activated protein (MAP) kinase kinase, NtMEK1, activates the cell cycle-regulated p43Ntf6 MAP kinase. J. Biol. Chem. 276, 18139-18145

31. Yang, K.Y. et al.(2001) Activation of a mitogen-activated protein kinase pathway is involved in disease resistance in tobacco. Proc. Natl. Acad. Sci. U.S.A. 98, 741-746

32. Shibata, W. et al. (1995) A tobacco protein kinase, NPK2, has a domain homologous to a domain found in activators of mitogen-activated protein kinases (MAPKKs). Mol. Gen. Genet. 246, 401-410

33. Liu, Y. et al. (2000) Molecular cloning and characterization of a tobacco MAP kinase kinase that interacts with SIPK. Mol. Plant Microbe Interact. 13, 118-124

34. Hardin, S.C. and Wolniak, S.M. (1998) Molecular cloning and characterization of maize ZmMEK1, a protein kinase with a catalytic domain homologous to mitogen- and stress- activated protein kinase kinases. Planta 206, 577-584

35. Jouannic, S. et al. (1999) Plant MAP kinase kinase kinases structure, classification and evolution. Gene 233, 1-11

36. Frye, C.A. et al. (2001) Negative regulation of defense responses in plants by a conserved MAPKK kinase. Proc. Natl. Acad. Sci. U.S.A. 98, 373-378

37. Nishihama, R. et al. (1997) Possible involvement of differential splicing in regulation of the activity of Arabidopsis ANP1 that is related to mitogen-activated protein kinase kinase kinases (MAPKKKs). Plant J.12, 39-48

38. Jouannic, S. et al.(2001) The protein kinases AtMAP3Ke1 and BnMAP3Ke1 are functional homologues of S. pombe cdc7p and may be involved in cell division. Plant J. 26, 637-649

39. Tregear, J.W. et al.(1996) An unusual protein kinase displaying characteristics of both the serine/threonine and tyrosine families is encoded by the Arabidopsis thaliana gene Atn1. Plant Sci. 117, 107-119

40. Ichimura, K. et al. (1997) Atmrk1, an Arabidopsis protein kinase related to mammal mixed-lineage kinases and Raf protein kinases. Plant Sci. 130, 171-179

41. Mizoguchi, T. et al. (1996) A gene encoding a mitogen-activated protein kinase kinase kinase is induced simultaneously with genes for a mitogen-activated protein kinase and an S6 ribosomal protein kinase by touch, cold, and water stress in Arabidopsis thaliana. Proc.Natl.Acad.Sci.U.S.A.93, 765-769

42. Covic, L. and Lew, R.R. (1996) Arabidopsis thaliana cDNA isolated by functional complementation shows homology to serine/threonine protein kinases. Biochim. Biophys. Acta 1305, 125-129

43. Kieber, J.J. et al.(1993) CTR1, a negative regulator of the ethylene response pathway in Arabidopsis, encodes a member of the raf family of protein kinases. Cell 72, 427-441

44. Lorenzo, al.(2000) Characterization and expression of two protein kinase genes and EIN-3 like gene, which are regulated by ABA and GA3 in dormant Fagus sylvatica seeds, In Seed Biology-Advances and Applications (Vol. 32)(Black,M. et al., eds), pp. 329-340, CAB international, Oxfordshire, UK

45. Feng, X.H. et al. (1993) Cloning and characterization of a novel member of protein kinase family from soybean. Biochim. Biophys. Acta 1172, 200-204

46. Banno, H. et al.(1993) NPK1, a tobacco gene that encodes a protein with a domain homologous to yeast BCK1, STE11, and Byr2 protein kinases. Mol. Cell. Biol. 13, 4745-4752