果树赤霉素代谢与信号途径研究进展
|
摘要
赤霉素作为5大植物激素之一,在果树的花芽分化、花序发育、开花坐果、果实的生长发育及植株的形态建成等方面扮演着重要的角色,但对果树赤霉素的分子生物学研究与其他大田作物相比差距较大。为了在果树生产中能更加合理有效地利用赤霉素调控果树花果发育,研究果树赤霉素的合成及其信号转导途径的分子调控机制十分必要。研究发现GA合成的关键酶KO、GA2ox及GA20ox的表达均与果树矮化呈负相关,而KS的含量则与植株高矮呈正相关,板栗雄性不育现象也与KO、KAO的表达量密切相关。GAMYB基因及LFY基因则在果树的成花诱导和雄蕊发育等生殖生长过程中发挥重要作用。DELLA蛋白在果树的GA信号途径中作为负调控因子可致使矮化植株形成,在果树的细胞周期循环过程、转录调控、花的形成、细胞的信号转导及许多生理过程中,DELLA蛋白泛素化降解均扮演着至关重要的角色。主要从果树赤霉素的合成及赤霉素的信号途径两大方面,着重对果树赤霉素合成过程中的关键酶基因及其定位、果树赤霉素信号途径的重要元件如赤霉素受体GID1、DELLA蛋白等进行了综述,以期为高效利用赤霉素调控果树生长发育提供重要的理论参考。
Gibberellin,as one of five major plant hormones,plays a key role in flower bud differentiation,inflorescence development,flowering and fruit setting,fruit growth and plant morphogenesis,etc. However,the research on molecular biology of gibberellin in fruit cropscompared with field crops differs largely. In order to more effectively and reasonably use gibberellin to regulate fruit and flower developmentof fruit trees in fruit crops production,it is necessary to study the synthesis of gibberellin and the molecular regulation mechanism of signaltransduction pathways. Here,two aspects,the synthesis of gibberellin of fruit crops and gibberellin signaling pathways,are reviewed,focusing on the key enzyme genes in the process of gibberellin synthesis and their localizations,as well as important components of gibberellinsignaling pathway such as gibberellin receptor GID1,DELLA protein and so on. The results showed that the expressions of KO,GA2 ox andGA20 ox in GA synthesis were negatively correlated with the dwarf of fruit trees,while the content of KS was positively correlated with plantheight. The male sterility of chestnut was also closely related to the expression of KO and KAO. GAMYB gene and LFY gene play an importantrole in the reproductive growth of fruit crops,such as flower formation and stamen development. DELLA protein,as a negative regulator inthe GA signaling pathway in fruit trees,lead to the formation of dwarf plants,and DELLA protein ubiquitination degradation plays a vital rolein the cell cycle,transcription regulation,flower formation,cell signal transduction,and many physiological processes of the fruit crops.This paper aims at providing an important theoretical reference for the modulation of fruit crop growth and development by effectively utilizinggibberellin.
Gibberellin,as one of five major plant hormones,plays a key role in flower bud differentiation,inflorescence development,flowering and fruit setting,fruit growth and plant morphogenesis,etc. However,the research on molecular biology of gibberellin in fruit cropscompared with field crops differs largely. In order to more effectively and reasonably use gibberellin to regulate fruit and flower developmentof fruit trees in fruit crops production,it is necessary to study the synthesis of gibberellin and the molecular regulation mechanism of signaltransduction pathways. Here,two aspects,the synthesis of gibberellin of fruit crops and gibberellin signaling pathways,are reviewed,focusing on the key enzyme genes in the process of gibberellin synthesis and their localizations,as well as important components of gibberellinsignaling pathway such as gibberellin receptor GID1,DELLA protein and so on. The results showed that the expressions of KO,GA2 ox andGA20 ox in GA synthesis were negatively correlated with the dwarf of fruit trees,while the content of KS was positively correlated with plantheight. The male sterility of chestnut was also closely related to the expression of KO and KAO. GAMYB gene and LFY gene play an importantrole in the reproductive growth of fruit crops,such as flower formation and stamen development. DELLA protein,as a negative regulator inthe GA signaling pathway in fruit trees,lead to the formation of dwarf plants,and DELLA protein ubiquitination degradation plays a vital rolein the cell cycle,transcription regulation,flower formation,cell signal transduction,and many physiological processes of the fruit crops.This paper aims at providing an important theoretical reference for the modulation of fruit crop growth and development by effectively utilizinggibberellin.
引文
[1]邢超,陈永胜,李跃,等.赤霉素代谢和信息转导及其对植株表型的影响研究进展[J].安徽农业科学,2015(35):225-226.
[2]何昕孺.赤霉素对摩尔多瓦葡萄果实生长发育及品质的影响[D].银川:宁夏大学,2013.
[3]田伟,田义轲,王彩虹,等.苹果内根-贝壳杉烯氧化酶基因的克隆及序列分析[J].果树学报,2011,28(1):1-7.
[4]姜志昂,彭建营,孙建设.苹果Md KS、Md KOA1基因克隆与表达分析[J].植物遗传资源学报,2014,15(2):362-368.
[5]王西成,任国慧,房经贵,等.葡萄赤霉素合成相关基因克隆,亚细胞定位和表达分析[J].中国农业科学,2012,45(11):2224-2231.
[6]田义轲,白牡丹,王彩虹,等.苹果Md CPS基因的克隆,定位及其在柱型/普通型间的表达差异分析[J].华北农学报,2014,29(6):21-27.
[7]邓晓云,戴洪义,梁美霞.苹果内根-贝壳杉烯合成酶基因的克隆及表达分析[J].华北农学报,2013,28(2):46-51.
[8]刘云龙,王彩虹,白牡丹,等.影响苹果赤霉素合成的关键酶基因在新梢和幼嫩种子上的表达分析[J].果树学报,2013,5:732-736.
[9]李文彬.‘红阳’猕猴桃果实发育转录组及花青素累积机理研究[D].北京:中国科学院大学,2015.
[10]欧春青,姜淑苓,王斐,等.梨贝壳杉烯酸氧化酶基因Pc KAO1的克隆与表达分析[J].园艺学报,2013,40(5):849-858.
[11]石琰景.果树GA生物合成基因-贝壳杉烯氧化酶和20-氧化酶的克隆及表达模式的初步研究[D].泰安:山东农业大学,2002.
[12]李节法,田义轲,王彩虹,等.梨贝壳杉烯氧化酶基因Pp KO的克隆及生物信息学分析[J].园艺学报,2010,37(10):1575-1582.
[13]田义轲,王彩虹,白牡丹,等.基于梨贝壳杉烯氧化酶基因Pp KO序列的功能性SNP标记[J].园艺学报,2012,39(10):1876-1884.
[14]田义轲,李节法,王彩虹,等.梨茎尖中Pp KO基因表达量的实时荧光定量PCR分析[J].华北农学报,,2012,27(3):62-66.
[15]程飞飞,欧春青,姜淑苓,等.梨内根-贝壳杉烯合酶基因克隆及表达分析[J].沈阳农业大学学报,2011,42(6):677-682.
[16]郭献平,李兴亮,段续伟,等.板栗野生和芽变雄花序赤霉素合成关键酶基因比较分析[J].中国农业大学学报,2012,17(4):91-95.
[17]郭献平.板栗短雄花序芽变与赤霉素相关的分子机理研究[J].乌鲁木齐:新疆农业大学,2012.
[18]李兴亮,郭献平,沈元月,等.板栗赤霉素缺陷型短雄花序芽变的初步鉴定及Cm GID1基因的表达分析[J].园艺学报,2011,38(7):1251-1258.
[19]刘炳臣.葡萄胚珠败育相关基因Vv ABCG20及活性GA代谢关键基因的初步分析[D].杨凌:西北农林科技大学,2016.
[20]Acheampong AK,Zheng C,Halaly T,et al.Abnormal endogenousrepression of GA signaling in a seedless table grape cultivar withhigh berry growth response to GA application[J].Frontiers inPlant Science,2017,8:850.
[21]程飞飞.矮生梨‘中矮1号’GA2-oxidase基因的克隆与功能分析[D].北京:中国农业科学院,2012.
[22]辛璐.短枝型苹果‘苏帅’赤霉素相关差异基因的筛选与表达分析[D].南京:南京农业大学,2015.
[23]丁伟.缺铁胁迫对梨氮代谢及赤霉素信号转导相关基因表达的影响[D].合肥:安徽农业大学,2015.
[24]王满,金灿,许雪,等.梨极矮化突变体与苹果梨GA20ox基因的克隆及植物表达载体的构建[J].北方园艺,2014(2):89-94.
[25]宣利利,欧春青,王斐,等.梨矮化砧木中矮1号GA20-氧化酶基因克隆与表达分析[J].果树学报,2011(5):883-887.
[26]宣利利,欧春青,王斐,等.梨品种中矮1号GA20-氧化酶基因正义和反义表达载体的构建[J].中国果树,2011(3):11-13.
[27]Wen LH,Zhong WJ,Huo XM,et al.Expression analysis of ABAand GA-related genes during four stages of bud dormancy inJapanese apricot(Prunus mume Sieb.et Zucc)[J].The Journalof Horticultural Science and Biotechnology,2016,91(4):362-369.
[28]刘锴栋,袁长春,黎海利,等.番荔枝GA20氧化酶基因的克隆与表达分析[J].植物生理学报,2015,51(10):1697-1705.
[29]Kotoda N,Matsuo S,Honda I,et al.Isolation and functional analysisof two Gibberellin 20-oxidase genes from Satsuma mandarin(Citrus unshiu Marc.)[J].The Horticulture Journal,2016,85(2):128-140.
[30]宋杨.富士苹果短枝型性状相关基因的分离及表达[D].泰安:山东农业大学,2012.
[31]沙广利,束怀瑞,郝玉金.苹果无融合生殖矮化种质创新[D].泰安:山东农业大学,2007.
[32]姜志昂,孙建设,彭建营,等.苹果Md GA20ox1基因的克隆,亚细胞定位及表达分析[J].园艺学报,2013,40(12):2373-2381.
[33]袁飞荣.转rol ABC基因枳橙砧木的评价[D].长沙:湖南农业大学,2011.
[34]Wang YJ,Deng DX.Molecular basis and evolutionarypattern ofGA-GID1-DELLA regulatory module[J].Molecular Geneticsand Genomics,2014,289(1):1-9.
[35]宋伟,李鼎立,王然,等.蔷薇科植物DELLA蛋白的生物信息学分析[J].中国农学通报,2013,29(19):142-148.
[36]钟翡,沈欣杰,刘芳,等.甜樱桃DELLA蛋白基因Pa GAI的克隆与表达分析[J].园艺学报,2012,39(1):143-150.
[37]徐丽,陈新,张力思,等.核桃GAI基因的克隆和序列分析[J].山东农业科学,2013,45(7):1-5.
[38]翟晓霏,张文,赵旭勉,等.苹果Md RGL1a基因对烟草遗传转化研究[J].中国农业大学学报,2011,16(1):36-41.
[39]刘杭,杨贺忠,李亮,等.梨休眠DELLA蛋白Pp GAI基因的克隆与表达分析[J].分子植物育种,2016(8):1995-2002.
[40]沙广利,束怀瑞,郝玉金.苹果无融合生殖矮化种质创新[D].泰安:山东农业大学,2007.
[41]雷恒久,姚立萍,王爽,等.草莓转录因子基因Fa GAMYB的克隆与表达分析[J].中国农业大学学报,2015,20(5):156-163.
[42]樊胜,雷超,辛明志,等.苹果赤霉素信号转导因子Md GAMYB的克隆和表达分析[J].园艺学报,2017,44(5):817-827.
[43]刘淑芳,陈力耕,陈大明,等.柑橘LEAFY同源基因片段的克隆及分析[J].浙江大学学报:农业与生命科学版,2001,27(3):297-300.
[44]曹秋芬,和田雅人,孟玉平,等.苹果LEAFY同源基因的c DNA克隆与表达分析[J].园艺学报,2003,30(3):267-271.
[45]李宁,陈厚彬,张昭其,等.荔枝LEAFY同源基因克隆及表达分析[J].华南农业大学学报,2013,34(1):57-61.
[46]殷姗姗,李茂福,王华,等.草莓Fa SKP1-1基因的克隆与表达分析[J].中国农业大学学报,2016,21(12):28-34.
[47]张演义,王化坤,宋长年,等.枳2个F-box基因c DNA全长的克隆及其亚细胞定位分析[J].华北农学报,2012,27(5):1-7.
[48]王玉兰,鲁振华,王志强,等.半矮生型桃F-box基因的克隆和表达特征分析[J].果树学报,2013,30(4):543-550.
[49]安振宇,张树伟,何新华,等.柠檬F-box基因的克隆及表达分析[J].分子植物育种,2015(11):2484-2490.
[2]何昕孺.赤霉素对摩尔多瓦葡萄果实生长发育及品质的影响[D].银川:宁夏大学,2013.
[3]田伟,田义轲,王彩虹,等.苹果内根-贝壳杉烯氧化酶基因的克隆及序列分析[J].果树学报,2011,28(1):1-7.
[4]姜志昂,彭建营,孙建设.苹果Md KS、Md KOA1基因克隆与表达分析[J].植物遗传资源学报,2014,15(2):362-368.
[5]王西成,任国慧,房经贵,等.葡萄赤霉素合成相关基因克隆,亚细胞定位和表达分析[J].中国农业科学,2012,45(11):2224-2231.
[6]田义轲,白牡丹,王彩虹,等.苹果Md CPS基因的克隆,定位及其在柱型/普通型间的表达差异分析[J].华北农学报,2014,29(6):21-27.
[7]邓晓云,戴洪义,梁美霞.苹果内根-贝壳杉烯合成酶基因的克隆及表达分析[J].华北农学报,2013,28(2):46-51.
[8]刘云龙,王彩虹,白牡丹,等.影响苹果赤霉素合成的关键酶基因在新梢和幼嫩种子上的表达分析[J].果树学报,2013,5:732-736.
[9]李文彬.‘红阳’猕猴桃果实发育转录组及花青素累积机理研究[D].北京:中国科学院大学,2015.
[10]欧春青,姜淑苓,王斐,等.梨贝壳杉烯酸氧化酶基因Pc KAO1的克隆与表达分析[J].园艺学报,2013,40(5):849-858.
[11]石琰景.果树GA生物合成基因-贝壳杉烯氧化酶和20-氧化酶的克隆及表达模式的初步研究[D].泰安:山东农业大学,2002.
[12]李节法,田义轲,王彩虹,等.梨贝壳杉烯氧化酶基因Pp KO的克隆及生物信息学分析[J].园艺学报,2010,37(10):1575-1582.
[13]田义轲,王彩虹,白牡丹,等.基于梨贝壳杉烯氧化酶基因Pp KO序列的功能性SNP标记[J].园艺学报,2012,39(10):1876-1884.
[14]田义轲,李节法,王彩虹,等.梨茎尖中Pp KO基因表达量的实时荧光定量PCR分析[J].华北农学报,,2012,27(3):62-66.
[15]程飞飞,欧春青,姜淑苓,等.梨内根-贝壳杉烯合酶基因克隆及表达分析[J].沈阳农业大学学报,2011,42(6):677-682.
[16]郭献平,李兴亮,段续伟,等.板栗野生和芽变雄花序赤霉素合成关键酶基因比较分析[J].中国农业大学学报,2012,17(4):91-95.
[17]郭献平.板栗短雄花序芽变与赤霉素相关的分子机理研究[J].乌鲁木齐:新疆农业大学,2012.
[18]李兴亮,郭献平,沈元月,等.板栗赤霉素缺陷型短雄花序芽变的初步鉴定及Cm GID1基因的表达分析[J].园艺学报,2011,38(7):1251-1258.
[19]刘炳臣.葡萄胚珠败育相关基因Vv ABCG20及活性GA代谢关键基因的初步分析[D].杨凌:西北农林科技大学,2016.
[20]Acheampong AK,Zheng C,Halaly T,et al.Abnormal endogenousrepression of GA signaling in a seedless table grape cultivar withhigh berry growth response to GA application[J].Frontiers inPlant Science,2017,8:850.
[21]程飞飞.矮生梨‘中矮1号’GA2-oxidase基因的克隆与功能分析[D].北京:中国农业科学院,2012.
[22]辛璐.短枝型苹果‘苏帅’赤霉素相关差异基因的筛选与表达分析[D].南京:南京农业大学,2015.
[23]丁伟.缺铁胁迫对梨氮代谢及赤霉素信号转导相关基因表达的影响[D].合肥:安徽农业大学,2015.
[24]王满,金灿,许雪,等.梨极矮化突变体与苹果梨GA20ox基因的克隆及植物表达载体的构建[J].北方园艺,2014(2):89-94.
[25]宣利利,欧春青,王斐,等.梨矮化砧木中矮1号GA20-氧化酶基因克隆与表达分析[J].果树学报,2011(5):883-887.
[26]宣利利,欧春青,王斐,等.梨品种中矮1号GA20-氧化酶基因正义和反义表达载体的构建[J].中国果树,2011(3):11-13.
[27]Wen LH,Zhong WJ,Huo XM,et al.Expression analysis of ABAand GA-related genes during four stages of bud dormancy inJapanese apricot(Prunus mume Sieb.et Zucc)[J].The Journalof Horticultural Science and Biotechnology,2016,91(4):362-369.
[28]刘锴栋,袁长春,黎海利,等.番荔枝GA20氧化酶基因的克隆与表达分析[J].植物生理学报,2015,51(10):1697-1705.
[29]Kotoda N,Matsuo S,Honda I,et al.Isolation and functional analysisof two Gibberellin 20-oxidase genes from Satsuma mandarin(Citrus unshiu Marc.)[J].The Horticulture Journal,2016,85(2):128-140.
[30]宋杨.富士苹果短枝型性状相关基因的分离及表达[D].泰安:山东农业大学,2012.
[31]沙广利,束怀瑞,郝玉金.苹果无融合生殖矮化种质创新[D].泰安:山东农业大学,2007.
[32]姜志昂,孙建设,彭建营,等.苹果Md GA20ox1基因的克隆,亚细胞定位及表达分析[J].园艺学报,2013,40(12):2373-2381.
[33]袁飞荣.转rol ABC基因枳橙砧木的评价[D].长沙:湖南农业大学,2011.
[34]Wang YJ,Deng DX.Molecular basis and evolutionarypattern ofGA-GID1-DELLA regulatory module[J].Molecular Geneticsand Genomics,2014,289(1):1-9.
[35]宋伟,李鼎立,王然,等.蔷薇科植物DELLA蛋白的生物信息学分析[J].中国农学通报,2013,29(19):142-148.
[36]钟翡,沈欣杰,刘芳,等.甜樱桃DELLA蛋白基因Pa GAI的克隆与表达分析[J].园艺学报,2012,39(1):143-150.
[37]徐丽,陈新,张力思,等.核桃GAI基因的克隆和序列分析[J].山东农业科学,2013,45(7):1-5.
[38]翟晓霏,张文,赵旭勉,等.苹果Md RGL1a基因对烟草遗传转化研究[J].中国农业大学学报,2011,16(1):36-41.
[39]刘杭,杨贺忠,李亮,等.梨休眠DELLA蛋白Pp GAI基因的克隆与表达分析[J].分子植物育种,2016(8):1995-2002.
[40]沙广利,束怀瑞,郝玉金.苹果无融合生殖矮化种质创新[D].泰安:山东农业大学,2007.
[41]雷恒久,姚立萍,王爽,等.草莓转录因子基因Fa GAMYB的克隆与表达分析[J].中国农业大学学报,2015,20(5):156-163.
[42]樊胜,雷超,辛明志,等.苹果赤霉素信号转导因子Md GAMYB的克隆和表达分析[J].园艺学报,2017,44(5):817-827.
[43]刘淑芳,陈力耕,陈大明,等.柑橘LEAFY同源基因片段的克隆及分析[J].浙江大学学报:农业与生命科学版,2001,27(3):297-300.
[44]曹秋芬,和田雅人,孟玉平,等.苹果LEAFY同源基因的c DNA克隆与表达分析[J].园艺学报,2003,30(3):267-271.
[45]李宁,陈厚彬,张昭其,等.荔枝LEAFY同源基因克隆及表达分析[J].华南农业大学学报,2013,34(1):57-61.
[46]殷姗姗,李茂福,王华,等.草莓Fa SKP1-1基因的克隆与表达分析[J].中国农业大学学报,2016,21(12):28-34.
[47]张演义,王化坤,宋长年,等.枳2个F-box基因c DNA全长的克隆及其亚细胞定位分析[J].华北农学报,2012,27(5):1-7.
[48]王玉兰,鲁振华,王志强,等.半矮生型桃F-box基因的克隆和表达特征分析[J].果树学报,2013,30(4):543-550.
[49]安振宇,张树伟,何新华,等.柠檬F-box基因的克隆及表达分析[J].分子植物育种,2015(11):2484-2490.