陆地棉开花相关基因GhFLP5的表达及功能分析
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摘要
【目的】克隆陆地棉开花促进因子家族的一个基因Flowering Promoting Factor1-like Protein 5(Gh FLP5),并对其时空表达模式进行研究,解析其在开花调控过程中的作用,为创制早熟陆地棉材料奠定基础。【方法】根据NCBI数据库中的序列,用Oligo7软件设计引物,以中棉所50(CCRI50)的c DNA为模板,克隆获得Gh FLP5。在Ex PASy网站上预测其蛋白的理化性质,同时在NCBI中检索其他物种中的FPF蛋白,使用Clustal X2进行多重序列比对,并用MEGA6构建系统进化树;取陆地棉早熟品种CCRI50和中晚熟品种鲁棉研28(Lu28)不同生长时期、不同组织的样品,对Gh FLP5的时间和空间表达模式进行分析;从基因组数据库中调取Gh FLP5起始密码子上游1 500 bp的片段,用Plant CARE在线工具预测其顺式作用元件,选取相关激素对二叶期棉花幼苗进行叶面喷施处理,研究Gh FLP5的应答反应;构建植物表达载体p BI121-Gh FLP5,转化拟南芥,观察转基因株系的表型,并用q RT-PCR对其内源基因表达的变化进行测定。【结果】Gh FLP5开放阅读框长300 bp,编码的蛋白质分子量为11.4 k D,共包含3个比较保守的区域,与大豆、苜蓿和毛果杨的开花促进因子亲缘关系较近。空间表达模式分析表明,Gh FLP5在叶片中优势表达,且在早熟品种CCRI50中的表达量显著高于晚熟品种Lu28;时间表达模式分析表明,在CCRI50中其表达量高峰出现在三叶期,而在Lu28中四叶期表达量最高。Gh FLP5的启动子上主要存在两大类顺式作用元件,一类是光响应元件和生物钟元件,一类是胁迫响应元件。根据顺式作用元件的分布和功能选取水杨酸(SA)、脱落酸(ABA)和茉莉酸(JA)喷施处理棉花幼苗,结果表明,Gh FLP5能响应外源SA和ABA而上调,也会被外施JA抑制。组成型表达Gh FLP5的拟南芥株系抽薹时间提前约9 d,开花时间提前约7 d,莲座叶数目减少,差异达到极显著水平。荧光定量的结果表明转基因拟南芥中促进开花的基因LEAFY(At LFY)、SUPPRESSOR OF OVEREXPRESSION OF CONSTANS(At SOC1)、FLOWERING LOCUS T(At FT)、APETALA1(At AP1)和FRUITFULL(At FUL)表达量显著升高,抑制开花的基因Flowering Locus C(At FLC)表达量显著降低。在转基因拟南芥中生长素应答基因SMALL AUXIN UPREGULATED 20(At SAUR20)和SMALL AUXIN UPREGULATED22(At SAUR22)显著上调,具有生物活性的赤霉素(GA)合成的相关基因GIBBERELLIN 20-OXIDASE 1(GA20OX1)的表达量提高2倍。【结论】转基因拟南芥早花表型明显,Gh FLP5可能在调控中发挥了双重作用,通过IAA和GA途径促进拟南芥的开花转型。
【Objective】To provide more information for breeding, the gene Flowering Promoting Factor1-like Protein 5(Gh FLP5), a member of the flowering promoting factor family in Gossypium hirsutum L., was cloned, and then its expression patterns were studied to characterize its functions in flowering process. 【Method】According to the sequences on NCBI, specific primers were designed by Oligo7 and Gh FLP5 was cloned from c DNA of cultivar CCRI50. The protein properties were predicted via Ex PASy. FPFs of other species were retrieved from NCBI. Clustal X2 was used for multiple sequence alignment and phylogenetic tree was constructed by MEGA6. The samples of different stages and different tissues of early maturity variety CCRI50 and later maturity variety Lu28 were used to study the spatial and temporal expression profiles of Gh FLP5. A 1 500 bp fragment upstream the start codon was taken from the genomic database, then the cis-acting elements were analysed with Plant CARE. Based on the predictions, several phytohormones were selected to explore the responses of Gh FLP5. p BI121-Gh FLP5, a recombinant expression plasmid was constructed and transformed into Arabidopsis. The homozygous overexpression lines were observed and expression profiles were performed with quantitative real-time PCR(q RT-PCR). 【Result】With a 300 bp coding frame, Gh FLP5 encoded a 11.4 k D protein. There were three pretty conserved domains, which reveals that Gh FLP5 has a close relationship with those of Glycine max, Medicago truncatula and Polulus trichocarpa. Spatial expression patterns showed that it expressed predominantly in leaves. And Gh FLP5 was transcribed at a higher level in early maturity variety CCRI50 than in later maturity variety Lu28. Temporal expression patterns showed that it hit a peak at three-true-leaf stage in CCRI50 but at four-true-leaf stage in Lu28. There were mainly two kinds of cis-acting elements in promoter region: one was light-response elements and circadian elements, and the other was stress-response elements. According to the elements' distributions and functions, salicylic acid(SA), abscisic acid(ABA) and jasmonic acid(JA) were selected to treat cotton seedlings. As a result, Gh FLP5 was activated by SA and ABA, while it was suppressed by JA. The overexpression plants bolted about 9 days and flowered about 7 days earlier than the wild type. Meanwhile, the rosette leaves decreased, and all the differences were extremely significant. Quantitative analysis showed that the genes promoting flowering such as LEAFY(At LFY), SUPPRESSOR OF OVEREXPRESSION OF CONSTANS(At SOC1), FLOWERING LOCUS T(At FT), APETALA1(At AP1) and FRUITFULL(At FUL) were up-regulated in 35S::Gh FLP5 lines, while the gene At FLC, delaying flowering, was down-regulated. Moreover, the auxin-responsive genes SMALL AUXIN UPREGULATED 20(At SAUR20) and SMALL AUXIN UPREGULATED 22(At SAUR22) were induced in transgenic Arabidopsis lines. GIBBERELLIN 20-OXIDASE 1(GA20OX1), a gene involved in gibberellin(GA) biosynthesis, was also up-regulated more than two folds. 【Conclusion】The Arabidopsis thaliana lines over-expressing Gh FLP5 performed an obvious early-flowering phenotype. Moreover, the gene expression profiles indicated that Gh FLP5 may play dual roles in the transition to flowering via both GA and IAA signaling pathways.
【Objective】To provide more information for breeding, the gene Flowering Promoting Factor1-like Protein 5(Gh FLP5), a member of the flowering promoting factor family in Gossypium hirsutum L., was cloned, and then its expression patterns were studied to characterize its functions in flowering process. 【Method】According to the sequences on NCBI, specific primers were designed by Oligo7 and Gh FLP5 was cloned from c DNA of cultivar CCRI50. The protein properties were predicted via Ex PASy. FPFs of other species were retrieved from NCBI. Clustal X2 was used for multiple sequence alignment and phylogenetic tree was constructed by MEGA6. The samples of different stages and different tissues of early maturity variety CCRI50 and later maturity variety Lu28 were used to study the spatial and temporal expression profiles of Gh FLP5. A 1 500 bp fragment upstream the start codon was taken from the genomic database, then the cis-acting elements were analysed with Plant CARE. Based on the predictions, several phytohormones were selected to explore the responses of Gh FLP5. p BI121-Gh FLP5, a recombinant expression plasmid was constructed and transformed into Arabidopsis. The homozygous overexpression lines were observed and expression profiles were performed with quantitative real-time PCR(q RT-PCR). 【Result】With a 300 bp coding frame, Gh FLP5 encoded a 11.4 k D protein. There were three pretty conserved domains, which reveals that Gh FLP5 has a close relationship with those of Glycine max, Medicago truncatula and Polulus trichocarpa. Spatial expression patterns showed that it expressed predominantly in leaves. And Gh FLP5 was transcribed at a higher level in early maturity variety CCRI50 than in later maturity variety Lu28. Temporal expression patterns showed that it hit a peak at three-true-leaf stage in CCRI50 but at four-true-leaf stage in Lu28. There were mainly two kinds of cis-acting elements in promoter region: one was light-response elements and circadian elements, and the other was stress-response elements. According to the elements' distributions and functions, salicylic acid(SA), abscisic acid(ABA) and jasmonic acid(JA) were selected to treat cotton seedlings. As a result, Gh FLP5 was activated by SA and ABA, while it was suppressed by JA. The overexpression plants bolted about 9 days and flowered about 7 days earlier than the wild type. Meanwhile, the rosette leaves decreased, and all the differences were extremely significant. Quantitative analysis showed that the genes promoting flowering such as LEAFY(At LFY), SUPPRESSOR OF OVEREXPRESSION OF CONSTANS(At SOC1), FLOWERING LOCUS T(At FT), APETALA1(At AP1) and FRUITFULL(At FUL) were up-regulated in 35S::Gh FLP5 lines, while the gene At FLC, delaying flowering, was down-regulated. Moreover, the auxin-responsive genes SMALL AUXIN UPREGULATED 20(At SAUR20) and SMALL AUXIN UPREGULATED 22(At SAUR22) were induced in transgenic Arabidopsis lines. GIBBERELLIN 20-OXIDASE 1(GA20OX1), a gene involved in gibberellin(GA) biosynthesis, was also up-regulated more than two folds. 【Conclusion】The Arabidopsis thaliana lines over-expressing Gh FLP5 performed an obvious early-flowering phenotype. Moreover, the gene expression profiles indicated that Gh FLP5 may play dual roles in the transition to flowering via both GA and IAA signaling pathways.
引文
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[2]李威.陆地棉花芽分化诱导阶段茎尖的表达谱分析[D].武汉:华中农业大学,2011.LI W.Transcriptome profiling of upland cotton shoot apex during floral induction[D].Wuhan:Huazhong Agricultural University,2011.(in Chinese)
[3]吴嫚.中棉所36花发育相关功能基因的发掘及其验证[D].杨陵:西北农林科技大学,2010.WU M.Research on the flowering-related gene in CCRI36[D].Yangling:Northwest A&F University,2010.(in Chinese)
[4]YU S,CAO L,ZHOU C M,ZHANG T Q,LIAN H,SUN Y,WU J Q,HUANG J R,WANG G D,WANG J W.Sugar is an endogenous cue for juvenile-to-adult phase transition in plants.Elife Sciences,2013,2(2):e00269.
[5]孙昌辉,邓晓建,方军,储成才.高等植物开花诱导研究进展.遗传,2007,29(10):1182-1190.SUN C H,DENG X J,FANG J,CHU C C.An overview of flowering transition in higher plants.Hereditas,2007,29(10):1182-1190.(in Chinese)
[6]MELZER S,MAJEWSKI D M,APEL K.Early changes in gene expression during the transition from vegetative to generative growth in the long-day plant Sinapis alba.The Plant Cell,1990,2(10):953-961.
[7]KANIA T,RUSSENBERGER D,PENG S,APEL K,MELZER S.FPF1 promotes flowering in Arabidopsis.The Plant Cell,1997,9(8):1327-1338.
[8]MELZER S,KAMPMANN G,CHANDLER J,APEL K.FPF1modulates the competence to flowering in Arabidopsis.The Plant Journal,1999,18(4):395-405.
[9]CHANDLER J W,MELZER S.An alpha-crystallin gene,ACD31.2from Arabidopsis is negatively regulated by FPF1 overexpression,floral induction,gibberellins,and long days.Journal of Experimental Botany,2004,55(401):1433-1435.
[10]GREENUP A G,SASANI S,OLIVER S N,TALBOT M J,DENNIS E S,HEMMING M N,TREVASKIS B.ODDSOC2 is a MADS box floral repressor that is down-regulated by vernalization in temperate cereals.Plant Physiology,2010,153(3):1062-1073.
[11]HOENICKA H,LAUTNER S,KLINGBERG A,KOCH G,EL-SHERIF F,LEHNHARDT D,ZHANG B,BURGERT I,ODERMATT J,MELZER S,FROMM J,FLADUNG M.Influence of over-expression of the Flowering Promoting Factor 1 gene(FPF1)from Arabidopsis on wood formation in hybrid poplar(Populus tremula L.×P.tremuloides Michx.).Planta,2012,235(2):359-373.
[12]XU M L,JIANG J F,GE L,XU Y Y,CHEN H,ZHAO Y,BI Y R,WEN J Q,CHONG K.FPF1 transgene leads to altered flowering time and root development in rice.Plant Cell Reports,2005,24(2):79-85.
[13]GE L,CHEN H,JIANG J F,ZHAO Y,XU M L,XU Y Y,TAN K H,XU Z H,CHONG K.Overexpression of Os RAA1 causes pleiotropic phenotypes in transgenic rice plants,including altered leaf,flower,and root development and root response to gravity.Plant Physiology,2004,135(3):1502-1513.
[14]Xu Y Y,CAO H,CHONG K.APC-targeted RAA1 degradation mediates the cell cycle and root development in plants.Plant Signaling&Behavior,2010,5(3):218-223.
[15]Wang H,Ge L,Ye H C,CHONG K,LIU B Y,LI G F.Studies on the effects of fpf1 gene on Artemisia annua flowering time and on the linkage between flowering and artemisinin biosynthesis.Planta Medica,2004,70(70):347-352.
[16]黄琼华,杨光伟,李德谋,罗小英,裴炎.农杆菌介导法将FPF1基因导入油菜的研究初报.西南大学学报,2002,24(2):124-127.HUANG Q H,YANG G W,LI D M,LUO X Y,PEI Y.Preliminary studies on transgenic rapeseed(Brassica Napus L.)with FPF1 gene mediated by Agrobacterium Tumefacien.Journal of Southwest Agricultural University,2002,24(2):124-127.(in Chinese)
[17]皮伟,李名扬.根癌农杆菌介导FPF1基因转化菊花的研究.西南大学学报,2007,29(4):70-73.PI W,LI M Y.Transferring FPF1 into Chrysanthemum mediated by Agrobacterium tumefaciens.Journal of Southwest Agricultural University,2007,29(4):70-73.(in Chinese)
[18]许园园,王燕,柳李旺,徐良,王克磊,张凤娇,聂姗姗,龚义勤.萝卜抽薹开花促进因子Rs FPF1基因克隆与功能分析.南京农业大学学报,2014,37(4):31-38.XU Y Y,WANG Y,LIU L W,XU L,WANG K L,ZHANG F J,NIE S S,GONG Y Q.Molecular cloning and functional analysis of flowering promoting factor 1 gene(Rs FPF1)involved in bolting and flowering control in radish(Raphanus sativus L.).Journal of Nanjing Agricultural University,2014,37(4):31-38.(in Chinese)
[19]WANG X Y,PANG C Y,WEI H L,YU S X.Involvement of cotton gene Gh FPF1 in the regulation of shade avoidance responses in Arabidopsis thaliana.Plant Signaling&Behavior,2015,10(9):631-634.
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