Vol. 49, No. 1, Chrysanthemum seticuspe f. boreale 2. FLOWERING LOCUS T FT / Heading date 3a Hd3a Corbesier et al ; Tamaki et al F

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1 特集 植物の生長調節 Regulation of Plant Growth & Development Vol.49, No.1, 34-40, 2014 Florigen and antiflorigen determine the flowering in chrysanthemum Atsushi Oda NARO Institute of Floricultural Science NIFS, National Agriculture and Food Research Organization NARO : Chrysanthemum is a typical short - day plant and responds to shortening of daylength in the transition from vegetative to reproductive phase. The manipulation of daylength is widely used in the commercial production of chrysanthemum. In chrysanthemum, daylength and growth temperature affect the development of floral organs after the floral transition. Recent advances in molecular genetics have identified a florigen in model plants. Recently, an antiflorigen which inhibits flowering antagonizing with florigen has been found in chrysanthemum. This review summarizes the flowering regulated by daylength and temperature through the balance of florigen and antiflorigen production in chrysanthemum. Chrysanthemum morifolium RAMAT. Garner Allard Chailakhyan and Krikorian The Japanese Society for Chemical Regulation of Plants

2 Vol. 49, No. 1, Chrysanthemum seticuspe f. boreale 2. FLOWERING LOCUS T FT / Heading date 3a Hd3a Corbesier et al ; Tamaki et al FT/Hd3a b - ZIP FD/OsFD1 APETALA1 AP1 / OsMADS15 FRUITFULL FUL Abe et al ; Taoka et al ; Wigge et al Hd3a RICE FLOWERING LOCUS T 1 RFT1 Hd3a RFT1 Komiya et al FT/Hd3a 3 CsFTL1 CsFTL2 Cs- FTL3CsFTL1 CsFTL2 CsFTL3 CsFTL3 1 Cs- FTL3 Oda et al CsFTL3 CsFTL3 CsFTL1 Higuchi et al Lang et al CsFTL3 Anti - florigenic FT/TFL1 family protein CsAFT Higuchi et al. 2013CsAFT BROTHER OF FT AND TFL1 BFT TERMINAL FLOWER 1 TFL1 FT Ratcliffe et al ; Hanano and Goto 2011BFT TFL1 FT TFL1 Yoo et al TFL1 CsTFL1 Higuchi et al CsAFT CsAFT CsFTL3 1 CsFDL1 CsFDL2 2 FD CsFTL3 CsFDL1 AP1/FUL CsAFL1 CsAFL2 Higuchi et al. 2013CsAFT CsFTL3 CsFDL1 CsAFL1 CsAFL2 CsFTL3 CsAFT CsAFT 発現量 16 栄養生長 日長 ( 時間 ) 12 開花 花器官の発達遅延 1 CsFTL3 CsAFT CsFTL3 CsAFT CsFTL3 CsAFT

3 36 Vol. 49, No. 1, 2014 Hd3a 10 Itoh et al CsFTL3 CsAFT 1 Hd3a CsFTL3 CsAFT 2 CsFTL3CsAFT CsFTL3 CsAFT CsFTL3 CsAFT CsFTL3 CsAFT Oda et al ; Higuchi et al. 2013CsFTL3 CsFTL3 CsFTL3 CsFTL3 CsAFT Cs- FTL3 CsFTL CsFTL3 Oda et al CsFTL3 CsAFT 1 PnFT1 1 Hayama et al CsFTL3 1 2 Oda et al. 2012CsFTL3 3 CsFTL3 CsFDL1 CsFTL3 CsFTL3 Higuchi et al CsFTL3 CsFTL3 Cs- FTL C 20 C Nakano et al CsFTL3 Nakano et al CmFTL3 FTL3 FTL3 CmFTL3 CsFTL3 CsAFT CsFTL3 5.

4 Vol. 49, No. 1, Sumitomo et al CsFTL3 CsAFT Higuchi et al B B CsPHYB RNAi CsFTL3 CsAFT Higuchi et al B CsFTL3 CsAFT B Hd3a Ishikawa et al CCT Grain number, plant height and heading date 7 Ghd7B - type Early heading date 1 Ehd1Hd3a Itoh et al. 2010Hd3a Ehd1 Ghd7 Ghd7 Itoh et al Ghd7 1 1 Ghd A CsAFT Higuchi et al CsAFT 8 4BGhd7 CsAFT Ghd7 CsAFT Ghd7 CsAFT Ghd7 CsAFT Ishikawa et al ; 2013 ; Higuchi et al CsAFT Ghd7 CsAFT Izawa et al CsFTL3 CsAFT Higuchi et al CsFTL3 CsFTL3 Oda et al CsAFT 8 CsAFT CsFTL3 CsAFT PnFT1 14 Hayama et al GIGANTEA GI FLAVIN - BINDING KELCH REPEAT, F - BOX FKF1 CYCLING DOF FACTOR 1 CDF1 CONSTANS CO FT Sawa et al GI CO OsGI Heading date 1 Hd1 Hd3a Ghd7 Ehd1 Hd3a Hd3a Hayama et al ; Itoh et al FT/Hd3a 1 CsFTL3 CsAFT 2 CsFTL3 CsAFT CsFTL3 CsAFT

5 38 Vol. 49, No. 1, 2014 SD 16 SD 12 SD 8 SD 4 SD 2 : SD : 4 4SD SD - 12SD - 16SD - 発現量 長日条件 短日条件 栄養生長 花序分裂組織の分化 CsFTL3 開花 CsAFT 短日処理の回数 3 CsFTL3 CsAFT CsFTL3 CsAFT Cs- FTL3 CsAFT 1 CsFTL3 GI/ OsGI CO/Hd1 Ghd7 Ehd1 CsFTL3 CsAFT GI CO FT BvFT1 BvFT2 BvFT2 BvFT1 Pin et al. 2010BvFT1 BvFT2 TFL1 FvTFL1 Koskela et al ; Mouhu et al CsFTL3 CsAFT FT/TFL1 CsAFT TFL1 8. CsFTL3 CsAFT

6 Vol. 49, No. 1, A B 長日条件 長日条件 Ghd7 CsAFT Ehd1 花成 短日条件 短日条件 Ehd1 Hd3a 花成 CsFTL3 花成 4 A : Ghd7 Ghd Ghd7 Hd3a Ehd1 Ghd7 Ehd1 Hd3a B : CsAFT CsAFT 8 CsAFT CsAFT CsFTL3 CsFTL3 CsAFT CsFTL3 CsAFT : : Abe, M, Kobayashi, Y, Yamamoto, S, Daimon, Y, Yamaguchi, A, Ikeda, Y, Ichinoki, H, Notaguchi, M, Goto, K and Araki, T 2005FD, a bzip protein mediating signals from the floral pathway integrator FT at the shoot apex. Science 309 : Chailakhyan, MK and Krikorian, AD 1975Forty years of research on the hormonal basis of plant development some personal reflections. The Botanical Review 41 : Corbesier, L, Vincent, C, Jang, S, Fornara, F, Fan, Q, Searle, I, Giakountis, A, Farrona, S, Gissot, L, Turnbull, C and Coupland, G 2007FT protein movement contributes to long - distance signaling in floral induction of Arabidopsis. Science 316 : Garner, WW and Allard, HA 1920Effect of the relative length of day and night and other factors of the environment on growth and reproduc-

7 40 Vol. 49, No. 1, 2014 tion in plants. J Agric Res 18 : Hanano, S and Goto, K 2011Arabidopsis TERMINAL FLOWER1 is involved in the regulation of flowering time and inflorescence development through transcriptional repression. Plant Cell 23 : Hayama, R, Agashe, B, Luley, E, King, R and Coupland, G 2007A circadian rhythm set by dusk determines the expression of FT homologs and the short - day photoperiodic flowering response in Pharbitis. Plant Cell 19 : Hayama, R, Yokoi, S, Tamaki, S, Yano, M and Shimamoto, K 2003 Adaptation of photoperiodic control pathways produces short - day flowering in rice. Nature 422 : Higuchi, Y, Narumi, T, Oda, A, Nakano, Y, Sumitomo, K, Fukai, S and Hisamatsu, T 2013The gated induction system of a systemic floral inhibitor, antiflorigen, determines obligate short - day flowering in chrysanthemums. Proc Natl Acad Sci U S A 110 : Ishikawa, R, Tamaki, S, Yokoi, S, Inagaki, N, Shinomura, T, Takano, M and Shimamoto, K 2005Suppression of the floral activator Hd3a is the principal cause of the night break effect in rice. Plant Cell 17 : Itoh, H, Nonoue, Y, Yano, M and Izawa, T 2010A pair of floral regulators sets critical day length for Hd3a florigen expression in rice. Nature Genetics 43 : Izawa, T, Oikawa, T, Sugiyama, N, Tanisaka, T, Yano, M and Shimamoto, K 2002Phytochrome mediates the external light signal to repress FT orthologs in photoperiodic flowering of rice. Genes Dev 16 : Komiya, R, Yokoi, S and Shimamoto, K 2009A gene network for long - day flowering activates RFT1 encoding a mobile flowering signal in rice. Development 136 : Koskela, EA, Mouhu, K, Albani, MC, Kurokura, T, Rantanen, M, Sargent, DJ, Battey, NH, Coupland, G, Elomaa, P and Hytönen, T 2012Mutation in TERMINAL FLOWER1 reverses the photoperiodic requirement for flowering in the wild strawberry Fragaria vesca. Plant Physiol 159 : Lang, A, Chailakhyan, MK and Frolova, IA 1977Promotion and inhibition of flower formation in a day neutral plant in grafts with a short - day plant and a long - day plant. Proc Natl Acad Sci USA 74 : Mouhu, K, Kurokura, T, Koskela, EA, Albert, VA, Elomaa, P, Hytönen, T 2013The Fragaria vesca homolog of SUPPRESSOR OF OVEREXPRESSION OF CONSTANS represses flowering and promotes vegetative growth. Plant Cell 25 : Nakano, Y, Higuchi, Y, Sumitomo, K and Hisamatsu, T 2013Flowering retardation by high temperature in chrysanthemums : involvement of FLOWERING LOCUS T - like 3 gene repression. J Exp Bot 64 : Oda, A, Narumi, T, Li, T, Kando, T, Higuchi, Y, Sumitomo, K, Fukai, S and Hisamatsu, T 2012CsFTL3, a chrysanthemum FLOWERING LOCUS T - like gene, is a key regulator of photoperiodic flowering in chrysanthemums. J Exp Bot 63 : Pin, PA, Benlloch, R, Bonnet, D, Wremerth-Weich, E, Kraft, T, Gielen, JJL and Nilsson, O 2010An antagonistic pair of FT homologs mediates the control of flowering time in sugar beet. Science 330 : Ratcliffe, OJ, Amaya, I, Vincent, CA, Rothstein, S, Carpenter, R, Coen, ES and Bradley, DJ 1998A common mechanism controls the life cycle and architecture of plants. Development 125 : Sawa, M, Nusinow, DA, Kay, SA and Imaizumi, T 2007FKF1 and GIGANTEA complex formation is required for day - length measurement in Arabidopsis. Science 318 : Sumitomo, K, Higuchi, Y, Aoki, K, Miyamae, H, Oda, A, Ishiwata, M, Yamada, M, Nakayama, M and Hisamatsu, T 2012Spectral sensitivity of flowering and FT - like gene expression in response to night - break light treatments in the chrysanthemum cultivar, Reagan. J Hort Sci Biotechnol 87 : Tamaki, S, Matsuo, S, Wong, HL, Yokoi, S and Shimamoto, K 2007 Hd3a protein is a mobile flowering signal in rice. Science 316 : Taoka, K, Ohki, I, Tsuji, H, Furuita, K, Hayashi, K, Yanase, T, Yamaguchi, M, Nakashima, C, Purwestri, YA, Tamaki, S, Ogaki, Y, Shimada, C, Nakagawa, A, Kojima, C and Shimamoto, K proteins act as intracellular receptors for rice Hd3a florigen. Nature 476 : Yoo, SJ, Chung, KS, Jung, SH, Yoo, SY, Lee, JS and Ahn, JH 2010 BROTHER OF FT AND TFL1 BFT has TFL1-like activity and functions redundantly with TFL1 in inflorescence meristem development in Arabidopsis. Plant J 63 : Wigge, PA, Kim, MC, Jaeger, KE, Busch, W, Schmid, M, Lohmann, JU and Weigel, D 2005Integration of spatial and temporal information during floral induction in Arabidopsis. Science 309 : : TEL : FAX : E - mail : atsushio@affrc.go.jp

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