能源植物小桐子赤霉素合成代谢及信号转导相关基因的鉴定及序列分析

doi:10.7677/ynzwyj201514076

摘要/Abstract

摘要：

赤霉素 (gibberellin, GA) 是一类非常重要的植物激素，在植物种子萌发、茎干伸长、叶片生长、腺毛发育、花粉成熟、开花诱导和果实成熟等生长发育过程中都发挥着重要的作用。GA在一年生草本植物中可以促进开花，而在大多数多年生木本植物中则抑制成花诱导。为了更好地研究赤霉素在木本油料能源植物小桐子 (Jatropha curcas) 开花调控方面的作用机理，我们对小桐子整个基因组中参与GA合成代谢和信号转导的全部基因进行了鉴定和序列分析。这些基因包括6个多基因家族编码的蛋白, 即GA2氧化酶（GA2oxidase, GA2ox）、GA3氧化酶（GA3oxidase, GA3ox）、GA20氧化酶（GA20oxidase, GA20ox）、GID1 （GIBBERELLIN INSENSITIVE DWARF1）、 DELLAs和Fbox蛋白，以及2个单基因编码的蛋白，EL1 （EARLY FLOWERING1）和SPY （SPINDLY）。采用拟南芥和水稻中已经鉴定的上述基因编码的蛋白序列在小桐子基因组序列数据库和本实验的小桐子转录组数据库中进行BLASTP分析，找到17个同源蛋白的全长序列，并将其与28个拟南芥的、16个水稻的、24个葡萄的和22个蓖麻的同源蛋白构建系统发育树进行比对分析。结果表明，小桐子中参与赤霉素合成代谢及信号转导的大多数基因与蓖麻和葡萄同源基因的相似度更高。

关键词: 赤霉素, 合成代谢, 信号转导, 小桐子, 系统发育

Abstract:

Gibberellins (GAs) are essential phytohormones that control many aspects of plant development, including seed germination, stem elongation, leaf growth, flowering induction, development of glandular hairs, and pollen maturation. However, there are different mechanisms underlying GAregulated flowering in perennial woody plants and annual herb plants. To facilitate study about the role of GAs in the biofuel plant Jatropha curcas, we identified all genes involved in GA metabolism and signaling pathways. These genes include members of six gene families, ie., GA2oxidase (GA3ox), GA3oxidase (GA3ox), GA20oxidase (GA20ox), GA receptor GIBBERELLIN INSENSITIVE DWARF1 (GID1), DELLA growth inhibitors (DELLAs), and Fbox proteins, and two single genes SPINDLY (SPY) and EARLY FLOWERING1 (EL1). Jatropha homologs of genes from Arabidopsis and rice (Oryza sativa) were identified by blasting the genome and transcriptome database of Jatropha. Total 17 genes involved in GA metabolism and signaling pathway were identified from Jatropha, and were phylogenetically analyzed with homologs from Arabidopsis, rice, grape (Vitis vinifera), and castor bean (Ricinus communis). Our results showed that compared to Arabidopsis and rice, protein sequences of genes involved in GA metabolism and signaling pathways in Jatropha showed a higher similarity to those from castor bean and grape.

Key words: Gibberellin, Metabolism, Phytohormone signaling, Jatropha curcas, Phylogenetic analysis

中图分类号:

Q 75

高聪聪1、2, 倪军1、3, 陈茂盛1、2, 徐增富1. 能源植物小桐子赤霉素合成代谢及信号转导相关基因的鉴定及序列分析[J]. Plant Diversity, 2015, 37(2): 157-167.

GAO Cong-Cong-, NI Jun-, CHEN Mao-Sheng-, XU Zeng-Fu. Characterization of Genes Involved in Gibberellin Metabolism and Signaling Pathway in the Biofuel Plant Jatropha curcas[J]. Plant Diversity, 2015, 37(2): 157-167.

参考文献

Achard P, Genschik P, 2009. Releasing the brakes of plant growth: how GAs shutdown DELLA proteins［J］. Journal of Experimental Botany, 60: 1085—1092
Bennett MD, Leitch IJ, Price HJ et al., 2003. Comparisons with Caenorhabditis (~100Mb) and Drosophila (~175Mb) using flow cytometry show genome size in Arabidopsis to be ~157Mb and thus ~25% larger than the Arabidopsis Genome Initiative estimate of ~125Mb［J］. Annals of Botany, 91: 547—557
Bolle C, 2004. The role of GRAS proteins in plant signal transduction and development［J］. Planta, 218: 683—692
Boss PK, Thomas MR, 2002. Association of dwarfism and floral induction with a grape ‘green revolution’ mutation［J］. Nature, 416: 847—850
CanoAsseleih LM, 1986. Chemical investigation of Jatropha curcas L. seeds (PhD. Thesis ) ［D］. London: University of London, UK
Carvalho CR, Clarindo WR, Praa MM et al., 2008. Genome size, base composition and karyotype of Jatropha curcas L., an important biofuel plant［J］. Plant Science, 174: 613—617
Chan AP, Crabtree J, Zhao Q et al., 2010. Draft genome sequence of the oilseed species Ricinus communis［J］. Nature Biotechnology, 28: 951—956
Cheng H, Qin L, Lee S et al., 2004. Gibberellin regulates Arabidopsis floral development via suppression of DELLA protein function［J］. Development, 131: 1055—1064
Costa GGL, Cardoso KC, Del Bem LEV et al., 2010. Transcriptome analysis of the oilrich seed of the bioenergy crop Jatropha curcas L.［J］. BMC Genomics, 11: 462
Dai C, Xue HW, 2010. Rice early flowering1, a CKI, phosphorylates DELLA protein SLR1 to negatively regulate gibberellin signalling［J］. EMBO Journal, 29: 1916—1927
Davière JM, Achard P, 2013. Gibberellin signaling in plants［J］. Development, 140: 1147—1151
Dill A, Sun TP, 2001. Synergistic derepression of gibberellin signaling by removing RGA and GAI function in Arabidopsis thaliana［J］. Genetics, 159: 777—785
Fairless D, 2007. Biofuel: the little shrub that couldmaybe［J］. Nature, 449: 652—655
Ghosh A, Chikara J, Chaudhary DR, 2011. Diminution of economic yield as affected by pruning and chemical manipulation of Jatropha curcas L.［J］. Biomass and Bioenergy, 35: 1021—1029
Ghosh A, Chikara J, Chaudhary DR et al., 2010. Paclobutrazol arrests vegetative growth and unveils unexpressed yield potential of Jatropha curcas［J］. Journal of Plant Growth Regulation, 29: 307—315
Giacomelli L, RotaStabelli O, Masuero D et al., 2013. Gibberellin metabolism in Vitis vinifera L. during bloom and fruitset: functional characterization and evolution of grapevine gibberellin oxidases［J］. Journal of Experimental Botany, 64: 4403—4419
GoldbergMoeller R, Shalom L, Shlizerman L et al., 2013. Effects of gibberellin treatment during flowering induction period on global gene expression and the transcription of floweringcontrol genes in Citrus buds［J］. Plant Science, 198: 46—57
Harberd NP, 2003. Relieving DELLA restraint［J］. Science, 299: 1853—1854
Hauvermale AL, Ariizumi T, Steber CM, 2012. Gibberellin signaling: a theme and variations on DELLA repression［J］. Plant Physiology, 160: 83—92
Hirakawa H, Tsuchimoto S, Sakai H et al., 2012. Upgraded genomic information of Jatropha curcas L.［J］. Plant Biotechnology, 29: 123—130
Immanen J, Nieminen K, Duchens Silva H et al., 2013. Characterization of cytokinin signaling and homeostasis gene families in two hardwood tree species: Populus trichocarpa and Prunus persica［J］. BMC Genomics, 14: 885
Jaillon O, Aury JM, Noel B et al., 2007. The grapevine genome sequence suggests ancestral hexaploidization in major angiosperm phyla［J］. Nature, 449: 463—467
Kaul S, Koo HL, Jenkins J et al., 2000. Analysis of the genome sequence of the flowering plant Arabidopsis thaliana［J］. Nature, 408: 796—815
Makkar HP, Becker K, 2009. Jatropha curcas, a promising crop for the generation of biodiesel and valueadded coproducts［J］. European Journal of Lipid Science and Technology, 111: 773—787
Makwana V, Shukla P, Robin P, 2010. GA application induces alteration in sex ratio and cell death in Jatropha curcas［J］. Plant Growth Regulation, 61: 121—125
Matsumoto T, Wu JZ, Kanamori H et al., 2005. The mapbased sequence of the rice genome［J］. Nature, 436: 793—800
Mitchum MG, Yamaguchi S, Hanada A et al., 2006. Distinct and overlapping roles of two gibberellin 3oxidases in Arabidopsis development［J］. Plant Journal, 45: 804—818
MuozFambuena N, Mesejo C, GonzálezMas MC et al., 2012. Gibberellic acid reduces flowering intensity in sweet orange［Citrus sinensis (L.) Osbeck］ by repressing CiFT gene expression［J］. Journal of Plant Growth Regulation, 31: 529—536
Nakajima M, Shimada A, Takashi Y et al., 2006. Identification and characterization of Arabidopsis gibberellin receptors［J］. The Plant Journal, 46 (5): 880—889
Olszewski N, Sun TP, Gubler F, 2002. Gibberellin signaling: Biosynthesis, catabolism, and response pathways［J］. Plant Cell, 14: S61—S80
Pi XJ (皮雪静), Pan BZ (潘帮珍), Xu ZF (徐增富), 2013. Induction of bisexual flowers by gibberellin in monoecious biofuel plant Jatropha curcas (Euphorbiaceae) ［J］. Plant Diversity and Resources (植物分类与资源学报), 35 (1): 26—32
sterlund T, 2001. Structurefunction relationships of hormonesensitive lipase［J］. European Journal of Biochemistry, 268: 1899—1907
Peter H, Stephen GT, 2012. Gibberellin biosynthesis and its regulation［J］. Biochemical Journal, 444: 11—25
Randoux M, Jeauffre J, Thouroude T et al., 2012. Gibberellins regulate the transcription of the continuous flowering regulator, RoKSN, a rose TFL1 homologue［J］. Journal of Experimental Botany, 63: 6543—6554
Robertson M, Swain SM, Chandler PM et al., 1998. Identification of a negative regulator of gibberellin action, HvSPY, in barley［J］. Plant Cell, 10: 995—1007
Sakamoto T, Miura K, Itoh H et al., 2004. An overview of gibberellin metabolism enzyme genes and their related mutants in rice［J］. Plant Physiology, 134: 1642—1653
Sato S, Hirakawa H, Isobe S et al., 2011. Sequence analysis of the genome of an oilbearing tree, Jatropha curcas L.［J］. DNA Research, 18: 65—76
Shimada A, UeguchiTanaka M, Nakatsu T et al., 2008. Structural basis for gibberellin recognition by its receptor GID1［J］. Nature, 456: 520—523
Shimada A, UeguchiTanaka M, Sakamoto T et al., 2006. The rice SPINDLY gene functions as a negative regulator of gibberellin signaling by controlling the suppressive function of the DELLA protein, SLR1, and modulating brassinosteroid synthesis［J］. Plant Journal, 48: 390—402
Song J (宋娟), Chen MS (陈茂盛), Li JL (李家龙) et al., 2013. Effects of soilapplied paclobutrazol on the vegetative and reproductive growth of biofuel plant Jatropha curcas［J］. Plant Diversity and Resources (植物分类与资源学报), 35 (2): 173—179
Sun TP, 2010. GibberellinGID1DELLA: a pivotal regulatory module for plant growth and development［J］. Plant Physiology, 154: 567—570
Swain SM, Tseng TS, Olszewski NE, 2001. Altered expression of SPINDLY affects gibberellin response and plant development［J］. Plant Physiology, 126: 1174—1185
Thomas SG, Phillips AL, Hedden P, 1999. Molecular cloning and functional expression of gibberellin 2oxidases, multifunctional enzymes involved in gibberellin deactivation［J］. Proceedings of the National Academy of Sciences, 96: 4698—4703
Tongumpai P, Jutamanee K, Sethapakdi R et al., 1991. Variation in level of gibberellinlike substances, during vegetative growth and flowering of mango cv. Khiew Sawoey. ISHS Acta Horticulturae 291 (III International Mango Symposium), 291
Tyler L, Thomas SG, Hu J et al., 2004. DELLA proteins and gibberellinregulated seed germination and floral development in Arabidopsis［J］. Plant Physiology, 135: 1008—1019
UeguchiTanaka M, Ashikari M, Nakajima M et al., 2005. GIBBERELLIN INSENSITIVE DWARF1 encodes a soluble receptor for gibberellin［J］. Nature, 437: 693—698
Wang H, Zou Z, Wang S et al., 2013. Global analysis of transcriptome responses and gene expression profiles to cold stress of Jatropha curcas L［J］. PLoS ONE, 8: e82817
Wild M, Daviere JM, Cheminant S et al., 2012. The Arabidopsis DELLA RGALIKE3 is a direct target of MYC2 and modulates jasmonate signaling responses［J］. Plant Cell, 24: 3307—3319
Wilkie JD, Sedgley M, Olesen T, 2008. Regulation of floral initiation in horticultural trees［J］. Journal of Experimental Botany, 59: 3215—3228
Williams J, Phillips AL, Gaskin P et al., 1998. Function and substrate specificity of the gibberellin 3 betahydroxylase encoded by the Arabidopsis GA4 gene［J］. Plant Physiology, 117: 559—563
Willige BC, Ghosh S, Nill C et al., 2007. The DELLA domain of GA INSENSITIVE mediates the interaction with the GA INSENSITIVE DWARF1A gibberellin receptor of Arabidopsis［J］. Plant Cell, 19: 1209—1220
Wilson RN, Heckman JW, Somerville CR, 1992. Gibberellin is required for flowering in Arabidopsis thaliana under short days［J］. Plant Physiology, 100: 403—408
Winston E, 1992. Evaluation of paclobutrazol on growth, flowering and yield of mango cv. Kensington Pride［J］. Australian Journal of Experimental Agriculture, 32: 97—104
Xu YL, Li L, Wu KQ et al., 1995. The Ga5 locus of Arabidopsis thaliana encodes a multifunctional gibberellin 20oxidasemolecular cloning and functional expression［J］. Proceedings of the National Academy of Sciences of the United States of America, 92: 6640—6644
Yamaguchi N, Winter CM, Wu MF et al., 2014. Gibberellin acts positively then negatively to control onset of flower formation in Arabidopsis［J］. Science, 344: 638—641
Zentella R, Zhang ZL, Park M et al., 2007. Global analysis of DELLA direct targets in early gibberellin signaling in Arabidopsis［J］. Plant Cell, 19: 3037—3057

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