10 Vol. 48, No. 1, a GA bar = 1 mm b - f GA GA 6 b 9 c 12 d 15 e 18 f bar = 10 µm Nakayama et al GA GA 6 6 DAA : days after anthesis GA 1

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1 学会賞植物の生長調節 Regulation of Plant Growth & Development Vol.48, No.1, 9-16, 2013 Studies on molecular mechanism for gibberellin - triggered actions Masatoshi Nakajima Dept. Appl. Biol. Chem., Univ. of Tokyo : Gibberellin GA regulates various developmental processes in plants. To study on some actions triggered by GA biochemically, we prepared antisera against GA. At first, we used the antisera for the detection of GA in plants. In developing seeds of morning glory and also of Arabidopsis, GA was immunologically detected in their integuments where starch granules were observed and then disappeared. Taking together with other result for related genes expression, we proposed that GA plays a role for seed maturation of dicotyledonous plants at least through the degradation of starch. Secondly, we established a biochemical method for a detection of water - soluble GA receptor candidates in crude solution, which led to the identification of a rice GA receptor, Gibberellin Insensitive Dwarf1 GID1. Its GA - binding activity and ligand selectivity were confirmed. The suppressor of GA signaling called DELLA boosted the GA - binding activity of GID1 in vitro, which means the formation of a GID1 - DELLA complex tightens the binding of GA to GID1. In Arabidopsis, we found three GA receptors. By using loss - of - function mutants, we elucidated the functional redundancy and partial specificity of their receptors. After some approaches, we proposed that the phenotype s appeared in the mutants depends on the abundance of the remaining receptor and on its preference to DELLAs. 1. GA GA GA GA GA GA LC/MS GC/MS GC/MS GA GA Nakajima et al. 1991a GA 10 GA GA Nakajima et al. 1991b GA GA GA GA Hasegawa et al ; Hasegawa et al GA Ipomoea nil The Japanese Society for Chemical Regulation of Plants

10 Vol. 48, No. 1, 2013 1 a GA bar = 1 mm b - f GA GA 6 b 9 c 12 d 15 e 18 f bar = 10 µm Nakayama et al. 2002 GA GA 6 6 DAA : days after anthesis GA 12 DAA 1 9 DAA integument GA Nakayama et al.

2 10 Vol. 48, No. 1, a GA bar = 1 mm b - f GA GA 6 b 9 c 12 d 15 e 18 f bar = 10 µm Nakayama et al GA GA 6 6 DAA : days after anthesis GA 12 DAA 1 9 DAA integument GA Nakayama et al GA GA GA α - 2 InAmy1 2 InAmy1 InAmy1 6 DAA 9 12 DAA 15 DAA 9 DAA InAmy1 GA 0.1 µm GA 3 6 GA Nakayama et al InAmy1 45 kda InAmy DAA InAmy1 12 DAA GA InAmy1 Nakayama et al ; Nakajima et al i In- Amy1 GA ii InAmy1 GA GA α - GA InAmy1 GA α - GA GA GA GA InAmy1 GA in situ GA cdna 4 InGA20ox1 2 InGA3ox1 2 GC/MS in situ In- GA20ox1 InGA3ox2 InAmy1 seed coat GA InAmy1 InAmy1 Nakajima et

situ 9 12 15 c 12 InGA20ox1 in situ 2 bar = 1 mm Nakayama et al.

3 Vol. 48, No. 1, GA a1 - a3 GA InGA20ox1 in situ b1 - b3 α - InAmy1 in situ c 12 InGA20ox1 in situ 2 bar = 1 mm Nakayama et al GA a1 - a3 GA AtGA20ox3 in situ o1 o2 e b1 - b3 c1 - c3 d1 - d3 AtGA3ox4 - KO GA bar = 50 µm Kim et al. 2005

4 12 Vol. 48, No. 1, 2013 al ; Nakayama et al GA InAmy1 GA InAmy1 GA α - 2 Nakayama et al GA 1 GA20 GA20 3. GA α - GA GA GA GA in situ AtGA20ox3 AtGA- 20ox2 AtGA3ox4 2 Kim et al GA GA 3 α - α - 3 AtAmy3 AtAmy3 in situ AtGA20ox2 AtGA3ox4 AtAmy3 GA GA α - Kim et al At- GA3ox4 T - DNA AtGA3ox4 - KO AtGA3ox4 - KO 3 α - AtAmy3 Kim et al GA α - AtGA3ox4 - KO AtGA- 3ox4 - KO 4 GA ga1 ga1 GA GA GA GA GA GA ga1 Kim et al GA mucilage GA GA

5 Vol. 48, No. 1, GA GA GA 90 GA α - GA Hooley et al ; Gilroy and Jones 1994 GA GA GA 100 GA GA GA GA Nakajima et al GA Nakajima et al N GA Park et al GA Ueguchi - Tanaka et al GA GA GA GA GA gid1 gid1 5 cm GA GA GID1 GID1 GA 5 gid1 GA two - hybrid GA DELLA GID1 GA GID1 DELLA GA Ueguchi - Tanaka et al GID1 3 Nakajima et al GA Iuchi et al GUS pgid1::gid1 - GUS 1 2 Suzuki et al.

6 14 Vol. 48, No. 1, GA AtGA3ox4 - KO 2 bar = 50 µm 10 µm GA AtGA3ox4 - KO bar = 200 µm Kim et al GA GA DELLA - DELLA DELLA DELLA GA DELLA DELLA Suzuki et al GA GA 3 7 GA GA DELLA DELLA Otani et al ; Yoon et al GA GA GA GA4 Ueguchi - Tanaka et al. 2005

7 Vol. 48, No. 1, GA 3 GA 3KO a GA b 3 2 AtGID1a AtGID1c ac c ab - 2KO, AtGID1a AtGID1b d 7 GA GA GA X1 DELLA X2 DELLA X3 GA X4 GA

8 16 Vol. 48, No. 1, 2013 Gilroy, S and Jones, RL 1994 Perception of gibberellin and abscisic acid at the external face of the plasma membrane of barley Hordeum vulgare L. aleurone protoplasts. Plant Physiol 104 : Hasegawa, M, Hashimoto, N, Zhang, J, Nakajima, M, Takeda, K, Yamaguchi, I and Murofushi, N 1995 Immunohistochemistry of gibberellin in rice anthers. Biosci Biotech Biochem 59 : Hasegawa, M, Nakajima, M, Takeda, K, Yamaguchi, I and Murofushi, N 1994 A novel gibberellin, 16α,17 - dihydroxy - 16,17 - dihydrogibberellin A O - β - D - glucopyranoside, from rice anthers. Phytochemistry 37 : Hooley, R, Beale, MH and Smith, SJ 1991 Gibberellin perception at the plasma membrane of Avena fatua aleurone protoplasts. Planta 183 : Iuchi, S, Suzuki, H, Kim, Y - C, Iuchi, A, Kuromori, T, Ueguchi - Tanaka, M, Asami, T, Yamaguchi, I, Matsuoka, M, Kobayashi, M and Nakajima, M 2007 Multiple loss - of - function of Arabidopsis gibberellin receptor AtGID1s completely shuts down a gibberellin signal. Plant J 50 : Kim, Y - C, Nakajima, M, Nakayama, A and Yamaguchi, I 2005 Contribution of gibberellins to the formation of Arabidopsis seed coat through starch degradation. Plant Cell Physiol 46 : Nakajima, M, Nakayama, A, Xu, Z - J and Yamaguchi, I 2004 Gibberellin induces alpha - amylase gene in seed coat of Ipomoea nil immature seeds. Biosci Biotech Biochem 68 : Nakajima, M, Shimada, A, Takashi, Y, Kim, Y - C, Park, S - H, Ueguchi - Tanaka, M, Suzuki, H, Katoh, E, Iuchi, S, Kobayashi, M, Maeda, T, Matsuoka, M and Yamaguchi, I 2006 Identification and characterization of Arabidopsis gibberellin receptors. Plant J 46 : Nakajima, M, Takita, K, Wada, H, Mihara, K, Hasegawa, M, Yamaguchi, I and Murofushi, N 1997 Partial purification and characterization of a gibberellin - binding protein from seedling of Azukia angularis. Biochem Biophys Res Comm 241 : Nakajima, M, Yamaguchi, I, Kizawa, S, Murofushi, N and Takahashi, N 1991a Preparation and validation of an antiserum specific for non - derivatized gibberellins. Plant Cell Physiol 32 : Nakajima, M, Yamaguchi, I, Nagatani, A, Kizawa, S, Murofushi, N, Furuya, M and Takahashi, N 1991b Monoclonal antibodies specific for non - derivatized gibberellins. I. Preparation of monoclonal antibodies against GA4 and their use in immunoaffinity column chromatography. Plant Cell Physiol 32 : Nakayama, A, Nakajima, M and Yamaguchi, I 2005 Distribution of gibberellins and expressional analysis of GA 20 - oxidase genes of morning glory during fruit maturation. Biosci Biotech Biochem 69 : Nakayama, A, Park, S - J, Xu, Z - J, Nakajima, M and Yamaguchi, I 2002 Immunohistochemistry of active gibberellins and gibberellin - inducible α - amylase in developing seeds of Morning glory. Plant Physiol 129 : Otani, M, Yoon, J - M, Park, S - H, Asami, T and Nakajima, M 2010 Screening and characterization of an inhibitory chemical specific to Arabidopsis gibberellin 2 - oxidases. Bioorg Med Chem Lett 20 : Park, S - H, Nakajima, M, Hasegawa, M and Yamaguchi, I 2005 Similarity and difference between the characteristics of gibberellin - binding protein and gibberellin 2 - oxidases in Adzuki bean Vigna angularis seedlings. Biosci Biotechnol Biochem 69 : Suzuki, H, Park, S - H, Okubo, K, Kitamura, J, Ueguchi - Tanaka, M, Iuchi, S, Katoh, E, Kobayashi, M, Yamaguchi, I, Matsuoka, M, Asami, T and Nakajima, M 2009 Differential expression and affinities of Arabidopsis gibberellin receptors can explain variation in phenotypes of multiple knock - out mutants. Plant J 60 : Ueguchi - Tanaka, M, Ashikari, M, Nakajima, M, Itoh, H, Katoh, E, Kobayashi, M, Chow, T - Y, Hsing, Y - C, Kitano, H, Yamaguchi, I and Matsuoka, M 2005 GIBBERELLIN INSENSITIVE DWARF1 encodes a soluble receptor for gibberellin. Nature 437 : Yoon, J - M, Nakajima, M, Mashiguchi, K, Park, S - H, Otani, M and Asami, T 2013 Chemical screening of an inhibitor for gibberellin receptors based on a yeast two - hybrid system. Bioorg Med Chem Lett 23 : : TEL : FAX : E - mail : nkjm@pgr1.ch.a.u - tokyo.ac.jp

GID1, Hiroaki KATO, Tomomi SATO and Miyako UEGUCHI-TANAKA: Gibberellin Receptor GID1: Gibberellin Recognition and Molecular Evolution

52 31-36 2010 2 GID1, Hiroaki KATO, Tomomi SATO and Miyako UEGUCHI-TANAKA: Gibberellin Receptor GID1: Gibberellin Recognition and Molecular Evolution Gibberellins GAs are phytohormones essential for many