桃PpmMDH基因克隆及采后激素处理下的表达分析
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摘要
苹果酸脱氢酶在植物的生长发育中发挥重要作用。本研究克隆了桃(Prunus persica)苹果酸脱氢酶(malate dehydrogenase, mMDH)基因(PpmMDH),分析了其序列特征及其在硬肉和软肉桃品种不同桃组织及果实成熟前后的表达情况,并探讨了不同激素处理对桃果实中PpmMDH基因表达的影响。结果表明,克隆得到了PpmMDH基因,GenBank登陆号为KF017594,该cDNA片段长度为1 239bp,具有1个1 020 bp的ORF,编码339个氨基酸,序列分析表明,PpmMDH编码的氨基酸序列与其他植物的mMDH蛋白有较高的相似性,系统进化分析结果表明PpmMDH与梅(Prunus mume)和甜樱桃(Pr.avium)同源关系最近。qRT-PCR结果表明,PpmMDH在雄蕊中表达量较高(P<0.05),其次是花瓣,在叶片、幼果和雌蕊中表达量较低。果实成熟软化后期,PpmMDH在硬肉桃'双久红'果实中的表达量高于软肉桃'川中岛白桃'的。脱落酸(abscisic acid, ABA)、萘乙酸(1-naphthylacetic acid, NAA)、乙烯利(ethephon,ETH)处理桃果实得出,软肉桃果实成熟软化过程中PpmMDH的表达与乙烯有拮抗作用,硬肉桃与软肉桃在果实成熟与贮藏过程中内源激素动态不同,PpmMDH表达水平与酶活动态不同。上述结果表明,PpmMDH不一定是维持果实硬度的上游基因,但是在软肉桃与硬肉桃不同材料间表现出了与生理差异同步的酶活差异。本研究结果为进一步探索PpmMDH在桃果实成熟软化中的作用提供了科学依据。
Malate dehydrogenase plays an important role in plant growth and development. The objective of this study was to clone malate dehydrogenase(PpmMDH), a gene possibly related to fruit ripening and softening in peach(Prunus persica), investigate its sequence characteristics and analyze its expression on different peach tissues and fruits before and after ripening firm-fleshed and soft-fleshed peach varieties. The results indicated that the full-length cDNA of PpmMDH(GenBank No. KF017594) in peach was cloned. The sequence consisted of 1 239 bp with an ORF of 1 020 bp, encoding a polypeptide of 339 amino acids.Homology analysis showed that the deduced PpmMDH protein was highly homologous to other mMDH proteins from different species. Phylogenetic analysis also indicated that PpmMDH was very closely related to mMDH of plum blossom(Prunus mume) 和 cherry(Pr. avium). qRT-PCR results showed that the PpmMDH expression was abundant in stamens, followed by petals and leaves, and lower in young fruits and pistil.During later stage of fruit ripening, the accumulation of PpmMDH was significantly higher in the firm-fleshed mutant 'Shuangjiuhong' than that in the soft-fleshed 'Kawanakajima Hakuto'(P<0.05). Abscisic acid(ABA),1-naphthylacetic acid(NAA) and ethephon(ETH) treatments showed that the expression of PpmMDH was upregulated by ABA, NAA in both cultivars, but the soft-fleshed peach variety 'Kawanakajima Hakuto' was induced in a short period of time, and the firm-fleshed peach variety 'Shuangjiuhong' was upregulated continuously. Different from ABA, NAA, the effect of ETH treatment on the induction of the gene in the softfleshed peach variety 'Kawanakajima Hakuto' was very weak, even negligible, but the gene was continuously upregulated in 'Shuangjiuhong', firm-fleshed peach variety. In the control group, the expression level of PpmMDH in 'Kawanakajima Hakuto' was significantly lower than that in 'Shuangjiuhong'. Infer from this, the expression of PpmMDH was antagonistic to ethylene during ripening and softening of soft-fleshed peach, the endogenous hormone dynamics of firm-fleshed peach and soft-fleshed each were different during fruit ripening and storage, and PpmMDH expression level was different from enzyme activity state. The above results indicated that PpmMDH was not necessarily the upstream gene to maintain fruit firmness, but showed the same enzyme activity difference as physiological difference between different materials of firm-fleshed and soft-fleshed peach. The present study laid a foundation for further exploring the role of PpmMDH in peach fruit ripening and softening. It also provides a scientific basis for the identification of the gene at the molecular level.
Malate dehydrogenase plays an important role in plant growth and development. The objective of this study was to clone malate dehydrogenase(PpmMDH), a gene possibly related to fruit ripening and softening in peach(Prunus persica), investigate its sequence characteristics and analyze its expression on different peach tissues and fruits before and after ripening firm-fleshed and soft-fleshed peach varieties. The results indicated that the full-length cDNA of PpmMDH(GenBank No. KF017594) in peach was cloned. The sequence consisted of 1 239 bp with an ORF of 1 020 bp, encoding a polypeptide of 339 amino acids.Homology analysis showed that the deduced PpmMDH protein was highly homologous to other mMDH proteins from different species. Phylogenetic analysis also indicated that PpmMDH was very closely related to mMDH of plum blossom(Prunus mume) 和 cherry(Pr. avium). qRT-PCR results showed that the PpmMDH expression was abundant in stamens, followed by petals and leaves, and lower in young fruits and pistil.During later stage of fruit ripening, the accumulation of PpmMDH was significantly higher in the firm-fleshed mutant 'Shuangjiuhong' than that in the soft-fleshed 'Kawanakajima Hakuto'(P<0.05). Abscisic acid(ABA),1-naphthylacetic acid(NAA) and ethephon(ETH) treatments showed that the expression of PpmMDH was upregulated by ABA, NAA in both cultivars, but the soft-fleshed peach variety 'Kawanakajima Hakuto' was induced in a short period of time, and the firm-fleshed peach variety 'Shuangjiuhong' was upregulated continuously. Different from ABA, NAA, the effect of ETH treatment on the induction of the gene in the softfleshed peach variety 'Kawanakajima Hakuto' was very weak, even negligible, but the gene was continuously upregulated in 'Shuangjiuhong', firm-fleshed peach variety. In the control group, the expression level of PpmMDH in 'Kawanakajima Hakuto' was significantly lower than that in 'Shuangjiuhong'. Infer from this, the expression of PpmMDH was antagonistic to ethylene during ripening and softening of soft-fleshed peach, the endogenous hormone dynamics of firm-fleshed peach and soft-fleshed each were different during fruit ripening and storage, and PpmMDH expression level was different from enzyme activity state. The above results indicated that PpmMDH was not necessarily the upstream gene to maintain fruit firmness, but showed the same enzyme activity difference as physiological difference between different materials of firm-fleshed and soft-fleshed peach. The present study laid a foundation for further exploring the role of PpmMDH in peach fruit ripening and softening. It also provides a scientific basis for the identification of the gene at the molecular level.
引文
陈露露.2010.两种苹果酸脱氢酶的克隆、表达和酶学分析[D].硕士学位论文,安徽师范大学,导师:朱国萍,pp.9-10.(Chen L L.2010.Cloning,expression and enzymatic characterization of two malate dehydrogenases[D].Thesis for M.S.,Anhui Normal University,Supervisor:Zhu G P,pp.9-10.)
陈露露,翟羽佳,王鹏,等.2010.茶树胞质型苹果酸脱氢酶的原核表达及生物信息学分析[J].云南植物研究,(001):32-40.(Chen L L,Zhai Y J,Wang P,et al.2010.Prokaryotic expression and bioinformatics analysis of cytosolic malate dehydrogenase from Camellia sinensis(Theaceae)[J].Acta Botanica Yunnanica,(001):32-40.)
邓秋菊,刘菊华,金志强,等.2011.香蕉果实采后成熟过程中苹果酸脱氢酶(MDH)及苹果酸含量的变化[J].热带农业科学,31(7):34-38.(Deng Q J,Liu J H,Jin Z Q,et al.2011.Changes of malate dehydrogenase and malic acid content during the ripening of banana fruit[J].Chinese Journal of Tropical Agriculture,31(7):34-38.)
韩英群,郭丹,魏鑫,等.2017.果实采后软化生理机制及调控技术研究进展[J].北方园艺,(05):180-183.(Han YQ,Guo D,Wei X,et al.2017.Advances in physiological mechanism and regulation techniques of postharvest softening of fruits[J].Northern Horticulture,(05):180-183.)
胡留申,李培环,董晓颖,等.2007.长采收期桃果实硬度变化与相关激素含量关系的研究[J].中国农学通报,23(5):281-284.(Hu L S,Li P H,Dong X Y,et al.2007.Studies on the relationship between change in firmness and content of correlative hormone in peach fruits of long-harvested period[J].China Agricultural Science Bulletin,23(5):281-284.)
贾晋,张鲁刚.2009.大白菜苹果酸脱氢酶基因的克隆及序列分析[J].园艺学报,36(9):363-368.(Jia J,Zhang LG.2009.Cloning and sequence analyzing of malate dehydrogenase gene in Chinese cabbage[J].Acta Horticulturae Sinica,36(9):363-368.)
蔺占兵,徐洋,马庆虎,等.2004.小麦中一类依赖于NAD的细胞质型苹果酸脱氢酶cDNA的克隆和进化树分析(英文)[J].农业生物技术学报,12(01):38-42.(Lin Z B,Xu Y,Ma Q H,et al.2004.Cloning and evolutionary analysis of a partial cytosolic malate dehydrogenase cD-NA from wheat[J].Journal of Agricultural Biotechnology,12(01):38-42.
刘炳辉,董晓颖,李志军,等.2008.硬肉桃果实成熟前后几种与果实软化相关的生理指标的变化[J].植物生理学通讯,44(5):887-890.(Liu B H,Dong X Y,Li Z J,et al.2008.Changes in several physiological indexes related to fruit softening of crisp peach fruits before and after ripening[J].Plant Physiology Journal,44(5):887-890.)
罗小英,崔衍波,邓伟,等.2004.超量表达苹果酸脱氢酶基因提高苜蓿对铝毒的耐受性[J].分子植物育种,2(5):621-626.(Luo X Y,Cui Y B,Deng W,et al.2004.Transgenic alfalfa plants overexpressing nodule-enhanced malate dehydrogenase enhances tolerance to aluminum toxicity[J].Molecular Plant Breeding,2(5):621-626.)
吕静神,周影,葛永红,等.2017.1-MCP和乙烯利处理对采后'金冠'苹果常温贮藏过程中生理变化及活性氧代谢的影响[J].食品工业科技,38(5):339-344.(Lv J S,Zhou Y,Ge Y H,et al.2017.Effects of 1-MCP and ethephon treatment on physiological changes and active oxygen metabolism of postharvest golden delicious apples during storage at room temperature[J].Science and Technology of Food Industry,38(5):339-344.)
庞发虎,赵爱玲,余明玉,等.2015.脱落酸和乙烯利与采后梨枣生理及质量指标的相关性分析[J].南方农业学报,46(11):2001-2005.(Pang F H,Zhao A L,Yu M Y,et al.2015.Correlation analysis between abscisic acid and ethephon and physiological and quality indexes of postharvest pear Jujube[J].Journal of Southern Agriculture,46(11):2001-2005.)
宋超,张立军,贾永光,等.2009.植物的苹果酸代谢和转运[J].植物生理学通讯,5:003.(Song C,Zhang L J,Jia YG,et al.2009.Malate metabolism and transport in plants[J].Plant Physiology Journal,5:003.)
汪新颖,王波,侯松涛,等.2009.苹果酸脱氢酶的结构及功能[J].生物学杂志,26(4):69-72.(Wang X Y,Wang B,Hou S T,et al.2009.Structure and function of malate dehydrogenases[J].Journal of biology,26(4):69-72.)
王庆杰.2016.苹果细胞质苹果酸脱氢酶基因的抗逆功能鉴定及机制研究[D].硕士学位论文.山东农业大学,导师:姚玉新,pp.5-6.(Wang Q J.2016.Identification and mechanism of malate dehydrogenase gene in apple cytoplasm[D].Thesis for M.S.,Shandong Agricultural University,Supervisor:Yao Y X,pp.5-6.)
王庆杰,金仲鑫,周李杰.2015.苹果MdcyMDH过量表达对光合、激素和生长的影响[J].中国农业科学,48(14):2868-2875.(Wang Q J,Jin Z X,Zhou L J,et al.2015.Impacts of MdcyMDH overexpression on photosynthesis,hormone and growth in apple[J].Chinese Agricultural Science,48(14):2868-2875.)
王晓云,毕玉芬.2006.植物苹果酸脱氢酶研究进展[J].生物技术通报,(04):44-47,50.(Wang X Y,Bi Y F.2006.Research progress of malate dehydrogenase in plants[J].Biotechnology Bulletin,(04):44-47,50.)
杨方威,段懿菲,冯叙桥.2016.脱落酸的生物合成及对水果成熟的调控研究进展[J].食品科学,37(3):266-272.(Yang F W,Duan Y F,Feng X Q.2016.Advances in biosynthesis and regulation of abscisic acid on fruit maturation[J].Food Science,37(3):266-272.)
姚玉新,李明,由春香,等.2010.苹果果实中苹果酸代谢关键酶与苹果酸和可溶性糖积累的关系[J].园艺学报,37(1):1-8.(Yao Y X,Li M,You C X,et al.2010.Relationship between malic acid metabolism-related key enzymes and accumulation of malic acid as well as the soluble sugars in apple fruit[J].Acta Horticulturae Sinica,37(1):1-8.)
祝军,柴璐,侯柄竹,等.2015.果实中脱落酸的研究进展与展望[J].园艺学报,42(9):1664-1672.(Zhu J,Chai L,Hou B Z,et al.2015.Research progress and prospect of abscisic acid in fruit[J].Acta Horticulturae Sinica,42(9):1664-1672.)
曾文芳,王志强,潘磊,等.2015.生长素对油桃'24-30'果实软化和乙烯生物合成的影响[J].果树学报,(2):200-205.(Zeng W F,Wang Z Q,Pan L,et al.2015.Effects of auxin on fruit softening and ethylene biosynthesis of Nectarine'24-30'[J].Journal of Fruit Science,(2):200-205.)
Christine G.1992.Partitioning of malate dehydrogenase isoenzymes into glyoxysomes,mitochondria,and chloroplasts[J].Plant Physiology,100:557-559.
Christopher R,Goward,David J.1994.Malate dehydrogenase:A model for structure,evolution,and catalysis[J].Protein Science,3:1883-1888.
Christelle E,Annick M,Elisabeth D,et al.2002.Isolation and characterization of six peach cDNAs encoding key proteins in organic acid metabolism and solute accumulation:Involvement in regulating peach fruit acidity[J].Physiologia Plantarum,114:259-270.
Danilo C,Centeno,Sonia Osorio,et al.2011.Malate plays a crucial role in starch metabolism,ripening,and soluble solid content of tomato fruit and affects postharvest softening[J].The Plant Cell,23:162-184.
Hayama H,Shimada T,Fujii H,et al.2006.Ethylene-regulation of fruit softening and softening-related genes in peach[J].Journal of Experimental Botany,57(15):40714-077.
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Lindén P,Keech O,Stenlund H,et al.2016.Reduced mitochondrial malate dehydrogenase activity has a strong effect on photorespiratory metabolism as revealed by 13Clabelling[J].Journal of Experimental Botany,67(10):3123-3135.
Tesfaye M,Temple S J,Allan D L,et al.2001.Overexpression of malate dehydrogenase in transgenic alfalfa enhances organic acid synthesis and confers tolerance to aluminum[J].Plant Physiology,127:1836-1844.
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Zhang M,Yuan B,Leng P.2009.The role of ABA in triggering ethylene biosynthesis and ripening of tomato fruit[J].Journal of Experimental Botany,60(6):1579-1588.
陈露露,翟羽佳,王鹏,等.2010.茶树胞质型苹果酸脱氢酶的原核表达及生物信息学分析[J].云南植物研究,(001):32-40.(Chen L L,Zhai Y J,Wang P,et al.2010.Prokaryotic expression and bioinformatics analysis of cytosolic malate dehydrogenase from Camellia sinensis(Theaceae)[J].Acta Botanica Yunnanica,(001):32-40.)
邓秋菊,刘菊华,金志强,等.2011.香蕉果实采后成熟过程中苹果酸脱氢酶(MDH)及苹果酸含量的变化[J].热带农业科学,31(7):34-38.(Deng Q J,Liu J H,Jin Z Q,et al.2011.Changes of malate dehydrogenase and malic acid content during the ripening of banana fruit[J].Chinese Journal of Tropical Agriculture,31(7):34-38.)
韩英群,郭丹,魏鑫,等.2017.果实采后软化生理机制及调控技术研究进展[J].北方园艺,(05):180-183.(Han YQ,Guo D,Wei X,et al.2017.Advances in physiological mechanism and regulation techniques of postharvest softening of fruits[J].Northern Horticulture,(05):180-183.)
胡留申,李培环,董晓颖,等.2007.长采收期桃果实硬度变化与相关激素含量关系的研究[J].中国农学通报,23(5):281-284.(Hu L S,Li P H,Dong X Y,et al.2007.Studies on the relationship between change in firmness and content of correlative hormone in peach fruits of long-harvested period[J].China Agricultural Science Bulletin,23(5):281-284.)
贾晋,张鲁刚.2009.大白菜苹果酸脱氢酶基因的克隆及序列分析[J].园艺学报,36(9):363-368.(Jia J,Zhang LG.2009.Cloning and sequence analyzing of malate dehydrogenase gene in Chinese cabbage[J].Acta Horticulturae Sinica,36(9):363-368.)
蔺占兵,徐洋,马庆虎,等.2004.小麦中一类依赖于NAD的细胞质型苹果酸脱氢酶cDNA的克隆和进化树分析(英文)[J].农业生物技术学报,12(01):38-42.(Lin Z B,Xu Y,Ma Q H,et al.2004.Cloning and evolutionary analysis of a partial cytosolic malate dehydrogenase cD-NA from wheat[J].Journal of Agricultural Biotechnology,12(01):38-42.
刘炳辉,董晓颖,李志军,等.2008.硬肉桃果实成熟前后几种与果实软化相关的生理指标的变化[J].植物生理学通讯,44(5):887-890.(Liu B H,Dong X Y,Li Z J,et al.2008.Changes in several physiological indexes related to fruit softening of crisp peach fruits before and after ripening[J].Plant Physiology Journal,44(5):887-890.)
罗小英,崔衍波,邓伟,等.2004.超量表达苹果酸脱氢酶基因提高苜蓿对铝毒的耐受性[J].分子植物育种,2(5):621-626.(Luo X Y,Cui Y B,Deng W,et al.2004.Transgenic alfalfa plants overexpressing nodule-enhanced malate dehydrogenase enhances tolerance to aluminum toxicity[J].Molecular Plant Breeding,2(5):621-626.)
吕静神,周影,葛永红,等.2017.1-MCP和乙烯利处理对采后'金冠'苹果常温贮藏过程中生理变化及活性氧代谢的影响[J].食品工业科技,38(5):339-344.(Lv J S,Zhou Y,Ge Y H,et al.2017.Effects of 1-MCP and ethephon treatment on physiological changes and active oxygen metabolism of postharvest golden delicious apples during storage at room temperature[J].Science and Technology of Food Industry,38(5):339-344.)
庞发虎,赵爱玲,余明玉,等.2015.脱落酸和乙烯利与采后梨枣生理及质量指标的相关性分析[J].南方农业学报,46(11):2001-2005.(Pang F H,Zhao A L,Yu M Y,et al.2015.Correlation analysis between abscisic acid and ethephon and physiological and quality indexes of postharvest pear Jujube[J].Journal of Southern Agriculture,46(11):2001-2005.)
宋超,张立军,贾永光,等.2009.植物的苹果酸代谢和转运[J].植物生理学通讯,5:003.(Song C,Zhang L J,Jia YG,et al.2009.Malate metabolism and transport in plants[J].Plant Physiology Journal,5:003.)
汪新颖,王波,侯松涛,等.2009.苹果酸脱氢酶的结构及功能[J].生物学杂志,26(4):69-72.(Wang X Y,Wang B,Hou S T,et al.2009.Structure and function of malate dehydrogenases[J].Journal of biology,26(4):69-72.)
王庆杰.2016.苹果细胞质苹果酸脱氢酶基因的抗逆功能鉴定及机制研究[D].硕士学位论文.山东农业大学,导师:姚玉新,pp.5-6.(Wang Q J.2016.Identification and mechanism of malate dehydrogenase gene in apple cytoplasm[D].Thesis for M.S.,Shandong Agricultural University,Supervisor:Yao Y X,pp.5-6.)
王庆杰,金仲鑫,周李杰.2015.苹果MdcyMDH过量表达对光合、激素和生长的影响[J].中国农业科学,48(14):2868-2875.(Wang Q J,Jin Z X,Zhou L J,et al.2015.Impacts of MdcyMDH overexpression on photosynthesis,hormone and growth in apple[J].Chinese Agricultural Science,48(14):2868-2875.)
王晓云,毕玉芬.2006.植物苹果酸脱氢酶研究进展[J].生物技术通报,(04):44-47,50.(Wang X Y,Bi Y F.2006.Research progress of malate dehydrogenase in plants[J].Biotechnology Bulletin,(04):44-47,50.)
杨方威,段懿菲,冯叙桥.2016.脱落酸的生物合成及对水果成熟的调控研究进展[J].食品科学,37(3):266-272.(Yang F W,Duan Y F,Feng X Q.2016.Advances in biosynthesis and regulation of abscisic acid on fruit maturation[J].Food Science,37(3):266-272.)
姚玉新,李明,由春香,等.2010.苹果果实中苹果酸代谢关键酶与苹果酸和可溶性糖积累的关系[J].园艺学报,37(1):1-8.(Yao Y X,Li M,You C X,et al.2010.Relationship between malic acid metabolism-related key enzymes and accumulation of malic acid as well as the soluble sugars in apple fruit[J].Acta Horticulturae Sinica,37(1):1-8.)
祝军,柴璐,侯柄竹,等.2015.果实中脱落酸的研究进展与展望[J].园艺学报,42(9):1664-1672.(Zhu J,Chai L,Hou B Z,et al.2015.Research progress and prospect of abscisic acid in fruit[J].Acta Horticulturae Sinica,42(9):1664-1672.)
曾文芳,王志强,潘磊,等.2015.生长素对油桃'24-30'果实软化和乙烯生物合成的影响[J].果树学报,(2):200-205.(Zeng W F,Wang Z Q,Pan L,et al.2015.Effects of auxin on fruit softening and ethylene biosynthesis of Nectarine'24-30'[J].Journal of Fruit Science,(2):200-205.)
Christine G.1992.Partitioning of malate dehydrogenase isoenzymes into glyoxysomes,mitochondria,and chloroplasts[J].Plant Physiology,100:557-559.
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