48 Takashi HIRAYAMA and Tsutomu UGAJIN This idea is consistent with the weak ethylene insensitive phenotype of the ein3 mutants. Future perspective As described above, the over all structure of the ethylene-signaling pathway, from the perception to the gene expression, has been elucidated through the studies using Arabidopsis (Figure 3). Orthologues for Arabidopsis ethylene-signaling components have been found in other plant systems, including tomato, maize and rice. Therefore the ethylene-signaling pathway described here presumably is common in large parts in plant kingdom. Figure 3 Schematic representation of a model for the ethylene-signaling pathway. Ethylene receptor binds a copper ion presumably during protein modification in the Golgi apparatus. Copper ion is delivered by RAN1 copper transporter. Ethylene receptor seems to localize to the ER membrane although this is still controversial. The subcellular localization of EIN2 is not elucidated yet. EIN2 might sense some signals although there is no evidence. EIN3 functions in the nuclear although it is not clear if EIN3 localizes to the nuclear in the absence of ethylene or not. In the absence of ethylene, ethylene receptor and CTR1 are active and presumably inhibit EIN2. When ethylene receptor recognizes ethylene molecule, it turns off and consequently allows EIN2 to activate EIN3 somehow. Activated EIN3 induces the expressions of the ERF1 gene and some ethylene responsive genes (not illustrated). Produced ERF1, a transcriptional factor, in turn activates ethylene responsive genes.
ETHYLENE-SIGNALING PATHWAY 49 In spite of amount of efforts made by many researchers world wide, there are still many unsolved issues. For example, how do ethylene receptors activate down stream signaling molecules?, what is the CTR1 substrate?, what is the EIN2 function?, how is EIN3 regulated?, how is EIN5 or EIN6 involved in the ethylene-signaling pathway?, etc. Additional genetic studies, such as the isolation of suppressor mutants for existing mutants, may be required to identify new components. Furthermore, describing the biochemical properties, cellular functions and threedimensional structures of known components is necessary for understanding the ethylene-signaling pathway. Reference 1. ABRAHAM, D., PODAR, K., PACHER, M., KUBICEK, M., WELZEL, N., HEMMINGS, B.A., DILWORTH, S.M., MISCHAK, H., KOLCH, W. and BACCARINI, M. (2000). Raf-1-associated protein phosphatase 2A as a positive regulator of kinase activation. J Biol Chem. 275: 22300-22304. 2. ALONSO, J.M., HIRAYAMA, T., ROMAN, G., NOURIZADEH, S. and ECKER, J.R. (1999). EIN2, a bifunctional transducer of ethylene and stress responses in Arabidopsis. Science. 284: 2148-2152. 3. ALONSO, J.M., STEPANOVA, A.N., SOLANO, R., WISMAN, E., FERRARI, S., AUSUBEL, F.M. and ECKER, J.R. (2003). Five components of the ethylene-response pathway identified in a screen for weak ethyleneinsensitive mutants in Arabidopsis. Proc Natl Acad Sci U S A. 100: 2992-2297. 4. BLEECKER, A.B., ESTELLE, M.A., SOMERVILLE, C. and KENDE, H. (1988). Insensitivity to ethylene conferred by a dominant mutation in Arabidopsis thaliana. Science. 241: 1086-1089. 5. CHAO, Q., ROTHENBERG, M., SOLAN, R., ROMAN, G., TERZAGHI, W. and ECKER, J.R. (1997). Activation of the ethylene gas response pathway in Arabidopsis by the nuclear protein ETHYLENE-INSENSITIVE3 and related proteins. Cell. 89: 1133-1144. 6. CHEN, Y.F., RANDLETT, M.D., FINDELL, J.L. and SCHALLER, G.E. (2002). Localization of the ethylene receptor ETR1 to the endoplasmic reticulum of Arabidopsis. Journal of Biological Chemistry. 277: 19861- 19866. 7. CLARK, K.L., LARSEN, P.B., WANG, X. and CHANG, C. (1998). Association of the Arabidopsis CTR1 Raflike kinase with the ETR1 and ERS ethylene receptors. Proc Natl Acad Sci U S A. 95: 5401-5406. 8. GAO, Z., CHEN, Y.F., RANDLETT, M.D., ZHAO, X.C., FINDELL, J.L., KIEBER, J.J. and SCHALLER, G.E. (2003). Localization of the Raf-like Kinase CTR1 to the Endoplasmic Reticulum of Arabidopsis through Participation in Ethylene Receptor Signaling Complexes. J Biol Chem. 278: 34725-34732. 9. GUZMAN, P. and ECKER, J.R. (1990). Exploiting the triple response of Arabidopsis to identify ethylenerelated mutants. Plant Cell. 2: 513-523. 10. HIRAYAMA, T., KIEBER, J.J., HIRAYAMA, N., KOGAN, M., GUZMAN, P., NOURIZADEH, S., ALONSO, J.M., DAILEY, W.P., DANCIS, A. and ECKER, J.R. (1999). RESPONSIVE-TO-ANTAGONIST1, a Menkes/Wilson disease-related copper transporter, is required for ethylene signaling in Arabidopsis. Cell. 97: 383-393. 11. HUA, J. and MEYEROWITZ, E.M. (1998). Ethylene responses are negatively regulated by a receptor gene family in Arabidopsis thaliana. Cell. 94: 261-271. 12. HUANG, Y., LI, H., HUTCHISON, C.E., LASKEY, J. and KIEBER, J.J. (2003). Biochemical and functional analysis of CTR1, a protein kinase that negatively regulates ethylene signaling in Arabidopsis. Plant Journal. 33: 221-233. 13. LARSEN, P.B. and CANCEL, J.D. (2003). Enhanced ethylene responsiveness in the Arabidopsis eer1 mutant results from a loss-of-function mutation in the protein phosphatase 2A A regulatory subunit, RCN1. Plant Journal. 34: 709-718.
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ISSN 0435-1096 Gamma Field Symposia
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The lecturers and the members of th
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KANEKO, T. KARITA, E. KASHIMA, M. K
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