外源葡萄糖对‘长富2号’苹果花芽生理分化期可溶性糖和相关基因表达的影响
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摘要
以‘长富2号’苹果为材料,于花后25和30 d喷施浓度1.5%和3%的葡萄糖,研究处理后短枝顶芽的发育状况、可溶性糖、淀粉含量及相关代谢酶活性的动态变化,并利用实时荧光定量PCR检测糖代谢相关基因与成花相关基因的表达模式。结果表明:葡萄糖处理后,短枝顶芽长度、宽度及质量有一定程度增加;短枝顶芽中,蔗糖、葡萄糖、山梨醇在生理分化中前期显著高于对照,淀粉含量整体升高,而果糖含量在整个生理分化期显著降低;山梨醇脱氢酶、山梨醇氧化酶、蔗糖合酶合成方向以及蔗糖磷酸合成酶活性在不同时期有所增加,而酸性与中性蔗糖转化酶,及蔗糖合酶分解方向酶活性普遍降低;叶片中碳水化合物积累明显,糖代谢相关酶活性也发生相似改变;顶芽糖代谢相关基因及2个MdTPS基因都响应葡萄糖处理;成花关键基因MdFT 1、MdFD、MdLFY、MdSOC 1和MdAP 1均显著上调,而成花抑制基因MdTFL 1从花后30 d开始,表达显著下调。
‘Nagafu 2'apple trees were sprayed with 1.5% and 3% glucose solutions at 25 days and 30 days after full bloom,respectively. Bud growth,endogenous sugar levels,starch content and activities of related enzymes in terminal spur buds were examined. Genes associated with sugar metabolism and flowering were studied by quantitative real-time PCR. The results revealed that glucose treatments increased bud size and fresh weight. The contents of sucrose,glucose and sorbitol were significantly higher than in the control during early stage of floweral bud differentiation,while starch increased generally,but the content of fructose significantly decreased during the whole period. The activities of sorbitol dehydrogenase,sorbitol oxidase,synthesis direction of sucrose synthase and sucrose phosphate synthase increased at different stages,while acid and neutral invertases,decomposition direction of sucrose synthase decreased. Carbohydrate accumulation in leaves was evident,and the activities of enzymes involved in sugar metabolism also changed similarly. Genes associated with sugar metabolism and two MdTPS genes in terminal buds all responded to glucose treatment. Furthermore,flowering genes like MdFT 1,MdFD,MdLFY,MdSOC 1,and MdAP 1 were significantly up-regulated,while the suppressor MdTFL 1 was down-regulated from beginning of floral induction.
‘Nagafu 2'apple trees were sprayed with 1.5% and 3% glucose solutions at 25 days and 30 days after full bloom,respectively. Bud growth,endogenous sugar levels,starch content and activities of related enzymes in terminal spur buds were examined. Genes associated with sugar metabolism and flowering were studied by quantitative real-time PCR. The results revealed that glucose treatments increased bud size and fresh weight. The contents of sucrose,glucose and sorbitol were significantly higher than in the control during early stage of floweral bud differentiation,while starch increased generally,but the content of fructose significantly decreased during the whole period. The activities of sorbitol dehydrogenase,sorbitol oxidase,synthesis direction of sucrose synthase and sucrose phosphate synthase increased at different stages,while acid and neutral invertases,decomposition direction of sucrose synthase decreased. Carbohydrate accumulation in leaves was evident,and the activities of enzymes involved in sugar metabolism also changed similarly. Genes associated with sugar metabolism and two MdTPS genes in terminal buds all responded to glucose treatment. Furthermore,flowering genes like MdFT 1,MdFD,MdLFY,MdSOC 1,and MdAP 1 were significantly up-regulated,while the suppressor MdTFL 1 was down-regulated from beginning of floral induction.
引文
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Cao Shang-yin,Zhang Jun-chang,Jiang Ai-hua,Wei Li-hua.2001.Study on the best period of regulation of apple flower buds.China Fruits,(1):14-17.(in Chinese)曹尚银,张俊昌,江爱华,魏立华.2001.苹果花芽孕育调控的最佳时期研究.中国果树,(1):14-17.
Cao Shang-yin,Zhang Qiu-ming,Wu Shun.2003.Advances in reaserch on the mechanism of flower-bud differentiation of fruit trees.Journal of Fruit Science,20(5):345-350.(in Chinese)曹尚银,张秋明,吴顺.2003.果树花芽分化机理研究进展.果树学报,20(5):345-350.
Chang Qiang.2010.Study on relationship between off-season floral induction in longan and nutrition of the carbon and nitrogen[Ph.D.Dissertation].Fuzhou:Fujian Agriculture and Forestry University.(in Chinese)常强.2010.龙眼反季节成花诱导与碳氮营养关系的研究[博士论文].福州:福建农林大学.
Chen Su-li,Peng Yu,Zhou Hua,Yu Bo,Dong Yan-jun,Teng Sheng.2014.Research advances in trehalose metabolism and trehalose-6-phosphate signaling in plants.Plant Physiology Journal,50(3):233-242.(in Chinese)陈素丽,彭瑜,周华,于波,董彦君,滕胜.2014.植物海藻糖代谢及海藻糖-6-磷酸信号研究进展.植物生理学报,50(3):233-242.
Cousson A,Van K T.1983.Light-and sugar-mediated control of direct de novo flower differentiation from tobacco thin cell layers.Plant Physiology,72(1):33-36.
Du L S,Qi S Y,Ma J J,Xing L B,Fan S,Zhang S W,Li Y M,Shen Y W,Zhang D,Han M Y.2017.Identification of TPS family members in apple(Malus×domestica Borkh.)and the effect of sucrose sprays on TPS expression and floral induction.Plant Physiology and Biochemistry,120:10-23.
Hanke M V,Henryk B,Bullet F,Peil A,H?ttasch C.2007.No flower no fruit-genetic potentials to trigger flowering in fruit trees.Genes Genomes Genomics,1(1):1-20.
Han Ming-yu,Li Yong-wu,Fan Chong-hui,Zhao Cai-ping.2008.Effects of branch bending angle on physiological characteristics and fruit quality of fuji apple.Acta Horticulturae Sinica,35(9):1345-1350.(in Chinese)韩明玉,李永武,范崇辉,赵彩萍.2008.拉枝角度对富士苹果树生理特性和果实品质的影响.园艺学报,35(9):1345-1350.
Huang Hai,Qiao Xian-sheng,Cao Shang-yin.1986.Studies on the period of physiological differentiation of flower bud in apple trees.Acta Horticulturae Sinica,3(3):181-186.(in Chinese)黄海,乔宪生,曹尚银.1986.关于苹果花芽生理分化时期的研究.园艺学报,3(3):181-186.
Ito A,Hayama H,Kashimura Y.2002.Sugar metabolism in buds during flower bud formation:a comparison of two Japanese pear[Pyrus pyrifolia (Burm.)Nak.]cultivars possessing different flowering habits.Scientia Horticulturae,96(1-4):163-175.
Ito A,Hayama H,Kashimura Y.2004.Possible roles of sugar concentration and its metabolism in the regulation of flower bud information in japanese pear(Pyrus pyrifolia).Acta Horticulturae,636(44):365-373.
Kanamaru N,Ito Y,Komori S,Saito M,Kato H,Takahashi S,Omura M,Soejima J,Shiratake K,Yamada K.2004.Transgenic apple transformed by sorbitol-6-phosphate dehydrogenase cDNA:switch between sorbitol and sucrose supply due to its gene expression.Plant Science,167(1):55-61.
Karve R,Lauria M,Virnig A,Xia X,Rauh B L,Moore B D.2010.Evolutionary lineages and functional diversification of plant hexokinases.Molecular Plant,3(2):334-346.
Kotoda N,Wada M,Komori S,Kidou S I,Abe K,Masuda T,Soejima J.2000.Expression pattern of homologues of floral meristem identity genes LFY and AP1 during flower development in apple.Journal of the American Society for Horticultural Science,125(4):398-403.
Li Jin-yan,Huang Ji,Yao Run-yu,Xiao Wen-jun,Guo Xin-hong.2017.Progresses on the cross-talk between sugar and hormone signaling.Life Science Research,21(1):79-84.(in Chinese)李金燕,黄吉,姚润屿,肖文君,郭新红.2017.植物糖与激素信号途径的交叉会话.生命科学研究,21(1):79-84.
Livak K J,Schmittgen T D.2001.Analysis of relative gene expression data using real-time quantitative PCR and the 2-ΔΔCT method.Methods,25(4):402-408.
Ma Xin-li,Qin Yuan,Wei Xiao-yu,Ma Feng-wang,Li Ming-jun.2014.Sequence and expression analysis of apple tonoplast monosaccharide transporter TMT genes and their relationship with sugar accumulation in fruit.Acta Horticulturae Sinica,41(7):1317-1325.(in Chinese)马新立,秦源,魏晓钰,马锋旺,李明军.2014.苹果糖转运蛋白TMT基因的表达及其与糖积累的关系.园艺学报,41(7):1317-1325.
Ohto M,Onai K,Furukawa Y,Aoki E,Araki T,Nakamura K.2001.Effects of sugar on vegetative development and floral transition in Arabidopsis.Plant Physiology,127(1):252-261.
Qian Hai-feng,Zhao Xiao-juan,Zhao Xin-ai.2003.Regulation ofα-amylase gene expression.Acta Agriculturae Boreali-Occidentalis Sinica,12(4):87-90.(in Chinese)钱海丰,赵晓娟,赵心爱.2003.α-淀粉酶基因表达的调控.西北农业学报,12(4):87-90.
Rolland F,Baena-Gonzalez E,Sheen J.2006.Sugar sensing and signaling in plants:conserved and novel mechanisms.Usaannual Reviews,57(1):675-709.
Rolland F,Sheen J.2005.Sugar sensing and signalling networks in plants.Biochemical Society Transactions,33(1):269-271.
Rosa M,Hilal M,Gonzalez J A,Prado F E.2009.Low-temperature effect on enzyme activities involved in sucrose-starch partitioning in salt-sressed and salt-acclimated cotyledons of quinoa(Chenopodium quinoa Willd.)seedlings.Plant Physiology and Biochemistry,47(4):300-307.
Tang Zhang-cheng.1999.Modern plant physiology experiment guide.Beijing:Science Press.(in Chinese)汤章城.1999.现代植物生理学实验指南.北京:科学出版社.
Tao R,Uratsu S L,Dandekar A M.1995.Sorbitol synthesis in transgenic tobacco with apple cDNA encoding NADP-dependent sorbitol-6-phosphate dehydrogenase.Plant&Cell Physiology,36(3):525-532.
Wahl V,Ponnu J,Schlereth A,Arrivault S,Langenecker T,Franke A,Feil R,Lunn J E,Stitt M,Schmid M.2013.Regulation of flowering by trehalose-6-phosphate signaling in Arabidopsis thaliana.Science,339(6120):704-707.
Xing L B,Zhang D,Li Y M,Shen Y W,Zhao C P,Ma J J,An Na,Han M Y.2015.Transcription profiles reveal sugar and hormone signaling pathways mediating flower induction in apple(Malus domestica Borkh.).Plant and Cell Physiology,56(10):2052-2068.
Yamaki S.1986.Roles of four sorbitol related enzymes and invertase in the seasonal alteration of sugar metabolism in apple tissue.J Amer Soc Hort Sci,111:134-137.
Zeng Xian-bin,Cao Xian-zu,Liang Jian-sheng.1993.Dynamic characteristics of invertase of sugarbeet roots(Beta vulgaris L.).Journal of Jiangsu Agricultural College,14(1):21-25.(in Chinese)曾宪彬,曹显祖,梁建生.1993.甜菜块根中蔗糖酶的若干动力学性质.江苏农学院学报,14(1):21-25.
Cao Shang-yin,Zhang Jun-chang,Jiang Ai-hua,Wei Li-hua.2001.Study on the best period of regulation of apple flower buds.China Fruits,(1):14-17.(in Chinese)曹尚银,张俊昌,江爱华,魏立华.2001.苹果花芽孕育调控的最佳时期研究.中国果树,(1):14-17.
Cao Shang-yin,Zhang Qiu-ming,Wu Shun.2003.Advances in reaserch on the mechanism of flower-bud differentiation of fruit trees.Journal of Fruit Science,20(5):345-350.(in Chinese)曹尚银,张秋明,吴顺.2003.果树花芽分化机理研究进展.果树学报,20(5):345-350.
Chang Qiang.2010.Study on relationship between off-season floral induction in longan and nutrition of the carbon and nitrogen[Ph.D.Dissertation].Fuzhou:Fujian Agriculture and Forestry University.(in Chinese)常强.2010.龙眼反季节成花诱导与碳氮营养关系的研究[博士论文].福州:福建农林大学.
Chen Su-li,Peng Yu,Zhou Hua,Yu Bo,Dong Yan-jun,Teng Sheng.2014.Research advances in trehalose metabolism and trehalose-6-phosphate signaling in plants.Plant Physiology Journal,50(3):233-242.(in Chinese)陈素丽,彭瑜,周华,于波,董彦君,滕胜.2014.植物海藻糖代谢及海藻糖-6-磷酸信号研究进展.植物生理学报,50(3):233-242.
Cousson A,Van K T.1983.Light-and sugar-mediated control of direct de novo flower differentiation from tobacco thin cell layers.Plant Physiology,72(1):33-36.
Du L S,Qi S Y,Ma J J,Xing L B,Fan S,Zhang S W,Li Y M,Shen Y W,Zhang D,Han M Y.2017.Identification of TPS family members in apple(Malus×domestica Borkh.)and the effect of sucrose sprays on TPS expression and floral induction.Plant Physiology and Biochemistry,120:10-23.
Hanke M V,Henryk B,Bullet F,Peil A,H?ttasch C.2007.No flower no fruit-genetic potentials to trigger flowering in fruit trees.Genes Genomes Genomics,1(1):1-20.
Han Ming-yu,Li Yong-wu,Fan Chong-hui,Zhao Cai-ping.2008.Effects of branch bending angle on physiological characteristics and fruit quality of fuji apple.Acta Horticulturae Sinica,35(9):1345-1350.(in Chinese)韩明玉,李永武,范崇辉,赵彩萍.2008.拉枝角度对富士苹果树生理特性和果实品质的影响.园艺学报,35(9):1345-1350.
Huang Hai,Qiao Xian-sheng,Cao Shang-yin.1986.Studies on the period of physiological differentiation of flower bud in apple trees.Acta Horticulturae Sinica,3(3):181-186.(in Chinese)黄海,乔宪生,曹尚银.1986.关于苹果花芽生理分化时期的研究.园艺学报,3(3):181-186.
Ito A,Hayama H,Kashimura Y.2002.Sugar metabolism in buds during flower bud formation:a comparison of two Japanese pear[Pyrus pyrifolia (Burm.)Nak.]cultivars possessing different flowering habits.Scientia Horticulturae,96(1-4):163-175.
Ito A,Hayama H,Kashimura Y.2004.Possible roles of sugar concentration and its metabolism in the regulation of flower bud information in japanese pear(Pyrus pyrifolia).Acta Horticulturae,636(44):365-373.
Kanamaru N,Ito Y,Komori S,Saito M,Kato H,Takahashi S,Omura M,Soejima J,Shiratake K,Yamada K.2004.Transgenic apple transformed by sorbitol-6-phosphate dehydrogenase cDNA:switch between sorbitol and sucrose supply due to its gene expression.Plant Science,167(1):55-61.
Karve R,Lauria M,Virnig A,Xia X,Rauh B L,Moore B D.2010.Evolutionary lineages and functional diversification of plant hexokinases.Molecular Plant,3(2):334-346.
Kotoda N,Wada M,Komori S,Kidou S I,Abe K,Masuda T,Soejima J.2000.Expression pattern of homologues of floral meristem identity genes LFY and AP1 during flower development in apple.Journal of the American Society for Horticultural Science,125(4):398-403.
Li Jin-yan,Huang Ji,Yao Run-yu,Xiao Wen-jun,Guo Xin-hong.2017.Progresses on the cross-talk between sugar and hormone signaling.Life Science Research,21(1):79-84.(in Chinese)李金燕,黄吉,姚润屿,肖文君,郭新红.2017.植物糖与激素信号途径的交叉会话.生命科学研究,21(1):79-84.
Livak K J,Schmittgen T D.2001.Analysis of relative gene expression data using real-time quantitative PCR and the 2-ΔΔCT method.Methods,25(4):402-408.
Ma Xin-li,Qin Yuan,Wei Xiao-yu,Ma Feng-wang,Li Ming-jun.2014.Sequence and expression analysis of apple tonoplast monosaccharide transporter TMT genes and their relationship with sugar accumulation in fruit.Acta Horticulturae Sinica,41(7):1317-1325.(in Chinese)马新立,秦源,魏晓钰,马锋旺,李明军.2014.苹果糖转运蛋白TMT基因的表达及其与糖积累的关系.园艺学报,41(7):1317-1325.
Ohto M,Onai K,Furukawa Y,Aoki E,Araki T,Nakamura K.2001.Effects of sugar on vegetative development and floral transition in Arabidopsis.Plant Physiology,127(1):252-261.
Qian Hai-feng,Zhao Xiao-juan,Zhao Xin-ai.2003.Regulation ofα-amylase gene expression.Acta Agriculturae Boreali-Occidentalis Sinica,12(4):87-90.(in Chinese)钱海丰,赵晓娟,赵心爱.2003.α-淀粉酶基因表达的调控.西北农业学报,12(4):87-90.
Rolland F,Baena-Gonzalez E,Sheen J.2006.Sugar sensing and signaling in plants:conserved and novel mechanisms.Usaannual Reviews,57(1):675-709.
Rolland F,Sheen J.2005.Sugar sensing and signalling networks in plants.Biochemical Society Transactions,33(1):269-271.
Rosa M,Hilal M,Gonzalez J A,Prado F E.2009.Low-temperature effect on enzyme activities involved in sucrose-starch partitioning in salt-sressed and salt-acclimated cotyledons of quinoa(Chenopodium quinoa Willd.)seedlings.Plant Physiology and Biochemistry,47(4):300-307.
Tang Zhang-cheng.1999.Modern plant physiology experiment guide.Beijing:Science Press.(in Chinese)汤章城.1999.现代植物生理学实验指南.北京:科学出版社.
Tao R,Uratsu S L,Dandekar A M.1995.Sorbitol synthesis in transgenic tobacco with apple cDNA encoding NADP-dependent sorbitol-6-phosphate dehydrogenase.Plant&Cell Physiology,36(3):525-532.
Wahl V,Ponnu J,Schlereth A,Arrivault S,Langenecker T,Franke A,Feil R,Lunn J E,Stitt M,Schmid M.2013.Regulation of flowering by trehalose-6-phosphate signaling in Arabidopsis thaliana.Science,339(6120):704-707.
Xing L B,Zhang D,Li Y M,Shen Y W,Zhao C P,Ma J J,An Na,Han M Y.2015.Transcription profiles reveal sugar and hormone signaling pathways mediating flower induction in apple(Malus domestica Borkh.).Plant and Cell Physiology,56(10):2052-2068.
Yamaki S.1986.Roles of four sorbitol related enzymes and invertase in the seasonal alteration of sugar metabolism in apple tissue.J Amer Soc Hort Sci,111:134-137.
Zeng Xian-bin,Cao Xian-zu,Liang Jian-sheng.1993.Dynamic characteristics of invertase of sugarbeet roots(Beta vulgaris L.).Journal of Jiangsu Agricultural College,14(1):21-25.(in Chinese)曾宪彬,曹显祖,梁建生.1993.甜菜块根中蔗糖酶的若干动力学性质.江苏农学院学报,14(1):21-25.
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