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大学・研究所にある論文を検索できる 「Comparative analysis of plant isochorismate synthases reveals structural mechanisms underlying their distinct biochemical properties」の論文概要。リケラボ論文検索は、全国の大学リポジトリにある学位論文・教授論文を一括検索できる論文検索サービスです。

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Comparative analysis of plant isochorismate synthases reveals structural mechanisms underlying their distinct biochemical properties

Yokoo, Shohei Inoue, Seiya Suzuki, Nana Amakawa, Naho Matsui, Hidenori Nakagami, Hirofumi Takahashi, Akira Arai, Ryoichi Katou, Shinpei 信州大学 DOI:10.1042/BSR20171457

2023.02.03

概要

Isochorismate synthase (ICS) converts chorismate into isochorismate, a precursor of primary and secondary metabolites including salicylic acid (SA). SA plays important roles in responses to stress conditions in plants. Many studies have suggested that the function of plant ICSs is regulated at the transcriptional level. In Arabidopsis thaliana, the expression of AtICS1 is induced by stress conditions in parallel with SA synthesis, and AtICS1 is required for SA synthesis. In contrast, the expression of NtICS is not induced when SA synthesis is activated in tobacco, and it is unlikely to be involved in SA synthesis. Studies on the biochemical properties of plant ICSs are limited, compared with those on transcriptional regulation. We analyzed the biochemical properties of four plant ICSs: AtICS1, NtICS, NbICS from Nicotiana benthamiana, and OsICS from rice. Multiple sequence alignment analysis revealed that their primary structures were well conserved, and predicted key residues for ICS activity were almost completely conserved. However, AtICS1 showed much higher activity than the other ICSs when expressed in Escherichia coli and N. benthamiana leaves. Moreover, the levels of AtICS1 protein expression in N. benthamiana leaves were higher than the other ICSs. Construction and analysis of chimeras between AtICS1 and OsICS revealed that the putative chloroplast transit peptides (TPs) significantly affected the levels of protein accumulation in N. benthamiana leaves. Chimeric and point-mutation analyses revealed that Thr(531), Ser(537), and Ile(550) of AtICS1 are essential for its high activity. These distinct biochemical properties of plant ICSs may suggest different roles in their respective plant species.

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参考文献

1 Dosselaere, F. and Vanderleyden, J. (2001) A metabolic node in action: chorismate-utilizing enzymes in microorganisms. Crit. Rev. Microbiol. 27,

75–131, https://doi.org/10.1080/20014091096710

2 Kerbarh, O., Chirgadze, D.Y., Blundell, T.L. and Abell, C. (2006) Crystal structures of Yersinia enterocolitica salicylate synthase and its complex with the

reaction products salicylate and pyruvate. J. Mol. Biol. 357, 524–534, https://doi.org/10.1016/j.jmb.2005.12.078

3 Dahm, C., Muller,

R., Schulte, G., Schmidt, K. and Leistner, E. (1998) The role of isochorismate hydroxymutase genes entC and menF in enterobactin

and menaquinone biosynthesis in Escherichia coli. Biochim. Biophys. Acta 1425, 377–386, https://doi.org/10.1016/S0304-4165(98)00089-0

4 Serino, L., Reimmann, C., Baur, H., Beyeler, M., Visca, P. and Haas, D. (1995) Structural genes for salicylate biosynthesis from chorismate in

Pseudomonas aeruginosa. Mol. Gen. Genet. 249, 217–228, https://doi.org/10.1007/BF00290369

5 Raymond, K.N., Dertz, E.A. and Kim, S.S. (2003) Enterobactin: an archetype for microbial iron transport. Proc. Natl. Acad. Sci. U.S.A. 100, 3584–3588,

https://doi.org/10.1073/pnas.0630018100

6 Pelludat, C., Brem, D. and Heesemann, J. (2003) Irp9, encoded by the highpathogenicity island of Yersinia enterocolitica, is able to convert chorismate

into salicylate, the precursor of the siderophore yersiniabactin. J. Bacteriol. 185, 5648–5653, https://doi.org/10.1128/JB.185.18.5648-5653.2003

7 Reumann, S. (2013) Biosynthesis of vitamin K1 (phylloquinone) by plant peroxisomes and its integration into signaling molecule synthesis pathways.

Subcell. Biochem. 69, 213–229, https://doi.org/10.1007/978-94-007-6889-5˙12

8 Mustafa, N.R., Kim, H.K., Choi, Y.H., Erkelens, C., Lefeber, A.W., Spijksma, G. et al. (2009) Biosynthesis of salicylic acid in fungus elicited Catharanthus

roseus cells. Phytochemistry 70, 532–539, https://doi.org/10.1016/j.phytochem.2009.01.009

9 Bartsch, M., Bednarek, P., Vivancos, P.D., Schneider, B., von Roepenack-Lahaye, E., Foyer, C.H. et al. (2010) Accumulation of isochorismate-derived

2,3-dihydroxybenzoic 3-O-β-D-xyloside in Arabidopsis resistance to pathogens and ageing of leaves. J. Biol. Chem. 285, 25654–25665,

https://doi.org/10.1074/jbc.M109.092569

10 Boatwright, J.L. and Pajerowska-Mukhtar, K. (2013) Salicylic acid: an old hormone up to new tricks. Mol. Plant Pathol. 14, 623–634,

https://doi.org/10.1111/mpp.12035

11 Wildermuth, M., Dewdney, J., Wu, G. and Ausubel, F.M. (2001) Isochorismate synthase is required to synthesize salicylic acid for plant defence. Nature

414, 562–565, https://doi.org/10.1038/35107108

12 Ogawa, D., Nakajima, N., Tamaoki, M., Aono, M., Kubo, A., Kamada, H. et al. (2007) The isochorismate pathway is negatively regulated by salicylic acid

signaling in O3 -exposed Arabidopsis. Planta 226, 1277–1285, https://doi.org/10.1007/s00425-007-0556-5

13 Hunter, L.J.R., Westwood, J.H., Heath, G., Macaulay, K., Smith, A.G., MacFarlane, S.A. et al. (2013) Regulation of RNA-dependent RNA polymerase 1

and isochorismate synthase gene expression in Arabidopsis. PLoS ONE 8, e66530, https://doi.org/10.1371/journal.pone.0066530

14 Zimmermann, P., Hennig, L. and Gruissem, W. (2005) Gene-expression analysis and network discovery using Genevestigator. Trends Plant Sci. 10,

407–409, https://doi.org/10.1016/j.tplants.2005.07.003

12

c 2018 The Author(s). This is an open access article published by Portland Press Limited on behalf of the Biochemical Society and distributed under the Creative Commons

Attribution License 4.0 (CC BY).

Bioscience Reports (2018) 38 BSR20171457

https://doi.org/10.1042/BSR20171457

` E., Catinot, J., Buchala, A., Doermann, P. et al. (2008) Characterization and biological function of the

15 Garcion, C., Lohmann, A., Lamodiere,

ISOCHORISMATE SYNTHASE2 gene of Arabidopsis. Plant Physiol. 147, 1279–1287, https://doi.org/10.1104/pp.108.119420

16 Shibata, Y., Kawakita, K. and Takemoto, D. (2010) Age-related resistance of Nicotiana benthamiana against hemibiotrophic pathogen Phytophthora

infestans requires both ethylene- and salicylic acid-mediated signaling pathways. Mol. Plant Microbe Interact. 23, 1130–1142,

https://doi.org/10.1094/MPMI-23-9-1130

17 Catinot, J., Buchala, A., Abou-Mansour, E. and Metraux,

J.-P. (2008) Salicylic acid production in response to biotic and abiotic stress depends on

isochorismate in Nicotiana benthamiana. FEBS Lett. 582, 473–478, https://doi.org/10.1016/j.febslet.2007.12.039

18 Ogawa, D., Nakajima, N., Sano, T., Tamaoki, M., Aono, M., Kubo, A. et al. (2005) Salicylic acid accumulation under O3 exposure is regulated by ethylene

in tobacco plants. Plant Cell Physiol. 46, 1062–1072, https://doi.org/10.1093/pcp/pci118

19 Ogawa, D., Nakajima, N., Seo, S., Mitsuhara, I., Kamada, H. and Ohashi, Y. (2006) The phenylalanine pathway is the main route of salicylic acid

biosynthesis in tobacco mosaic virus -infected tobacco leaves. Plant Biotechnol. 23, 395–398, https://doi.org/10.5511/plantbiotechnology.23.395

´ J. and Raskin, I. (1995) Biosynthesis and metabolism of salicylic acid. Proc. Natl. Acad. Sci. U.S.A. 92, 4076–4079,

20 Lee, H.I., Leon,

https://doi.org/10.1073/pnas.92.10.4076

21 Silverman, P., Seskar, M., Kanter, D., Schweizer, P., Metraux,

J.P. and Raskin, I. (1995) Salicylic acid in rice: biosynthesis, conjugation, and possible role.

Plant Physiol. 108, 633–639, https://doi.org/10.1104/pp.108.2.633

22 Kolappan, S., Zwahlen, J., Zhou, R., Truglio, J.J., Tonge, P.J. and Kisker, C. (2007) Lysine 190 is the catalytic base in MenF, the menaquinone-specific

isochorismate synthase from Escherichia coli: implications for an enzyme family. Biochemistry 46, 946–953, https://doi.org/10.1021/bi0608515

23 Sridharan, S., Howard, N., Kerbarh, O., Blaszczyk, M., Abell, C. and Blundell, T.L. (2010) Crystal structure of Escherichia coli enterobactin-specific

isochorismate synthase (EntC) bound to its reaction product isochorismate: implications for the enzyme mechanism and differential activity of

chorismate-utilizing enzymes. J. Mol. Biol. 397, 290–300, https://doi.org/10.1016/j.jmb.2010.01.019

24 van Tegelen, L.J.P., Moreno, P.R.H., Croes, A.F., Verpoorte, R. and Wullems, G.J. (1999) Purification and cDNA cloning of isochorismate synthase from

elicited cell suspension cultures of Catharanthus roseus. Plant Physiol. 119, 705–712, https://doi.org/10.1104/pp.119.2.705

25 Strawn, M.A., Marr, S.K., Inoue, K., Inada, N., Zubieta, C. and Wildermuth, M.C. (2007) Arabidopsis isochorismate synthase functional in

pathogen-induced salicylate biosynthesis exhibits properties consistent with a role in diverse stress responses. J. Biol. Chem. 282, 5919–5933,

https://doi.org/10.1074/jbc.M605193200

26 Macaulay, K.M., Heath, G.A., Ciulli, A., Murphy, A.M., Abell, C., Carr, J.P. et al. (2017) The biochemical properties of the two Arabidopsis thaliana

isochorismate synthases. Biochem. J. 474, 1579–1590, https://doi.org/10.1042/BCJ20161069

27 Verberne, M.C., Verpoorte, R., Bol, J.F., Mercado-Blanco, J. and Linthorst, H.J.M. (2000) Overproduction of salicylic acid in plants by bacterial

transgenes results in enhanced resistance to pathogens. Nat. Biotechnol. 18, 779–783, https://doi.org/10.1038/77347

28 Ohtsubo, N., Mitsuhara, I., Koga, M., Seo, S. and Ohashi, Y. (1999) Ethylene promotes the necrotic lesion formation and basic PR gene expression in

TMV-infected tobacco. Plant Cell Physiol. 40, 808–817, https://doi.org/10.1093/oxfordjournals.pcp.a029609

29 Seo, S., Okamoto, M., Seto, H., Ishizuka, K., Sano, H. and Ohashi, Y. (1995) Tobacco MAP kinase: a possible mediator in wound signal transduction

pathways. Science 270, 1988–1992, https://doi.org/10.1126/science.270.5244.1988

30 Young, I.G. and Gibson, F. (1969) Regulation of the enzymes involved in the biosynthesis of 2,3-dihydroxybenzoic acid in Aerobacter aerogenes and

Escherichia coli. Biochim. Biophys. Acta. 177, 401–411, https://doi.org/10.1016/0304-4165(69)90302-X

31 Biasini, M., Bienert, S., Waterhouse, A., Arnold, K., Studer, G., Schmidt, T. et al. (2014) SWISS-MODEL: modelling protein tertiary and quaternary

structure using evolutionary information. Nucleic Acids Res. 42, W252–W258, https://doi.org/10.1093/nar/gku340

32 Guex, N., Peitsch, M.C. and Schwede, T. (2009) Automated comparative protein structure modeling with SWISS-MODEL and Swiss-PdbViewer: a

historical perspective. Electrophoresis 30 (Suppl. 1), S162–S173, https://doi.org/10.1002/elps.200900140

33 Kiefer, F., Arnold, K., Kunzli, M., Bordoli, L. and Schwede, T. (2009) The SWISS-MODEL repository and associated resources. Nucleic Acids Res. 37,

D387–D392, https://doi.org/10.1093/nar/gkn750

34 Caligiuri, M.G. and Bauerle, R. (1991) Identification of amino acid residues involved in feedback regulation of the anthranilate synthase complex from

Salmonella typhimurium. J. Biol. Chem. 266, 8328–8335

35 Mauch, F., Mauch-Mani, B., Gaille, C., Kull, B., Haas, D. and Reimmann, C. (2001) Manipulation of salicylate content in Arabidopsis thaliana by the

expression of an engineered bacterial salicylate synthase. Plant J. 25, 67–77, https://doi.org/10.1046/j.1365-313x.2001.00940.x

36 Jang, I.C., Lee, K.H., Nahm, B.H. and Kim, J.K. (2002) Chloroplast targeting signal of a rice rbcS gene enhances transgene expression. Mol. Breed. 9,

81–91, https://doi.org/10.1023/A:1026787308828

37 Xu, L., Zhao, H., Ruan, W., Deng, M., Wang, F., Peng, J. et al. (2017) ABNORMAL INFLORESCENCE MERISTEM1 functions in salicylic acid biosynthesis to

maintain proper reactive oxygen species levels for root meristem activity in rice. Plant Cell 29, 560–574, https://doi.org/10.1105/tpc.16.00665

38 Shine, M.B., Yang, J.W., El-Habbak, M., Nagyabhyru, P., Fu, D.Q., Navarre, D. et al. (2016) Cooperative functioning between phenylalanine ammonia

lyase and isochorismate synthase activities contributes to salicylic acid biosynthesis in soybean. New Phytol. 212, 627–636,

https://doi.org/10.1111/nph.14078

39 Gross, J., Cho, W.K., Lezhneva, L., Falk, J., Krupinska, K., Shinozaki, K. et al. (2006) J. Biol. Chem. 281, 17189–17196,

https://doi.org/10.1074/jbc.M601754200

40 Kim, E.H., Suh, S.C., Park, B.S., Shin, K.S., Kweon, S.J., Han, E.J. et al. (2009) Chloroplast-targeted expression of synthetic cry1Ac in transgenic rice as

an alternative strategy for increased pest protection. Planta 230, 397–405, https://doi.org/10.1007/s00425-009-0955-x

41 Yuan, Y., Chung, J.-D., Fu, X., Johnson, V.E., Ranjan, P., Booth, S.L. et al. (2009) Alternative splicing and gene duplication differentially shaped the

regulation of isochorismate synthase in Populus and Arabidopsis. Proc. Natl. Acad. Sci. U.S.A. 106, 22020–22025,

https://doi.org/10.1073/pnas.0906869106

c 2018 The Author(s). This is an open access article published by Portland Press Limited on behalf of the Biochemical Society and distributed under the Creative Commons Attribution

License 4.0 (CC BY).

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523

LVEKGLGALIYAGTGIVAGSDPSSEWNELDLKISQFTKSIEYEATTSLQAIN------------------------.................S..N........E.........L.H.SALQPIN--------------------------.................E...S.L..E..E..T.....LMKL..PLVTRGQRRIINQNQKVQRA------------.................E...S.L..E..E..T.....LMKL..PLLTRGQSRIINQNQKGLSCLPHRI-------.................E...S.L..E..E..T.....LMKL..PLLTRREIRTINQNQKGLSCLQHHI-------.................E...S.L..E..E..T.....LMKL..PLLTRGEIRTINQNQKGLSCLQHHILVCLHHSF

.IG.DA.......L.V.E....AL..Q..E..A...M.LMKL..PALK----------------------------.A...............K..N..L..D..E..T.....LLKL.GPSRQKIE.SGIIN-------------------.LG..HST.V...A...E.TN..F..D.....A...A.LLQ.QEQHICLQEAENMGTVI-----------------* *. .:*:*** *:* *::.: **:**:** *** * :: :

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