苹果果实发育中类黄酮含量变化及相关基因的研究
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摘要
类黄酮是苹果最重要的次生代谢产物，在苹果营养品质、生长发育、抗逆抗病以及苹果分类学等研究中有重要作用，本文以类黄酮含量较高的几种野生型苹果和类黄酮含量较低的栽培型苹果为试验材料，通过对不同基因型、不同部位以及不同发育时期的苹果果实进行研究，分析苹果果实中类黄酮含量的变化规律及相关调控基因的表达规律，并对转录因子MdMYB12进行了基因克隆分析及表达，为今后利用分子生物学方法培育果肉高类黄酮含量的苹果品种的育种工作奠定基础。全文研究分为以下内容：
(1)以‘新疆野生苹果’、‘八棱海棠’、‘津轻’和‘国庆’的果实为试材，用反相高效液相色谱法和分光光度法测定果皮、果肉在幼果期和膨大期绿原酸和总黄酮的含量差别和变化趋势，发现类黄酮含量变化与绿原酸变化没有必然联系，类黄酮在果皮中含量较高，而绿原酸在果肉中含量较高，二者在幼果期含量均高于膨大期，在野生型苹果中含量均高于栽培型苹果。用反相高效液相色谱法测定四种苹果果皮、果肉在幼果期和膨大期12种类黄酮化合物的含量差别和变化趋势的实验中发现，野生型苹果‘新疆野生苹果’中黄烷醇和根皮柑含量较高，栽培型苹果‘津轻’中黄酮醇含量较高，四种苹果的果皮果肉中，类黄酮物质含量均随着果实的成熟而降低。
(2)为了进一步探明类黄酮含量变化与果实发育进程的关系，以栽培型苹果‘津轻’和野生型苹果‘丽江山荆子’5个不同发育时期的果皮果肉为试验材料，对果实外观测定中发现，随着果实的发育成熟，‘津轻’果实明显膨大，而‘丽江山荆子’果实大小无明显变化。‘丽江山荆子’果实发育各个时期中绿原酸和总黄酮含量均远高于‘津轻’，苹果果实中绿原酸含量变化与果实膨大无直接关系，而总黄酮含量随着果实的膨大而下降。对果实中12种类黄酮含量变化的分析中发现，虽然‘津轻’总黄酮含量远远低于‘丽江山荆子’，但其果皮中黄酮醇类物质含量却高于‘丽江山荆子’；黄烷醇含量在‘丽江山荆子’中随着果实的发育无显著变化，且果肉和果皮中含量无显著差异，在‘津轻’果实发育前期黄烷醇含量迅速下降，后期含量无显著变化。‘津轻’果皮中根皮柑含量随着果实发育呈缓慢下降趋势，而在果肉中变化不显著，‘丽江山荆子’果皮果肉中的根皮柑含量变化趋势相同，均在果实发育前期有所下降，随着果实的发育成熟含量无显著变化。
(3)应用实时荧光定量PCR技术，分析5个传统内参基因/&SWW丄ACTB、GAPDH、UBQ和TUB在苹果不同基因型、不同组织以及果实不同发育时期的mRNA表达差异情况。经geNorm程序分析发现，5个内参基因的表达稳定性各异，在果实不同基因型以及不同发育时期的基因表达分析中最稳定；JCar和在6种不同组织中表达均稳定。本研究证明了在所有材料中均表达稳定，是研究苹果基因表达分析中首选的内参基因。
(4)对‘津轻’和‘丽江山荆子’中调控类黄酮合成代谢的11个结构基因的荧光定量PCR分析中发现，类黄酮含量较高的‘丽江山荆子’中PAL,CHS,CHI,FHT,FLS,DFR,LAR,ANS,ANR和朽丑这10个基因表达量均显著高于‘津轻’，FGT在两种苹果中表达量无明显差异。随着果实的发育进行，基因表达量存在差异，且与部分类黄酮含量变化呈正相关关系，其中CHI基因与类黄酮含量变化趋势相关性最显著。
(5)利用对已公布的‘金冠’苹果基因组的生物信息学分析，以‘丽江山荆子’果皮为实验材料克隆了 转录因子，对在‘津轻’和‘丽江山荆子’的荧光定量分析中发现，其转录与多种类黄酮物质存在正相关。
Flavonoids are the most important secondary metabolites in apple. They play an important role inapple nutritional quality, growth and development and disease resistance, as well as in apple taxonomyresearch. This study utilizes several types of higher flavonoid content in wild-type apples and lowerflavonoid content in cultivated-type apples as experimental mateirals, and takes the different genotypes,different parts and different developmental periods of apple fruits as research objects to study thechange rules of flavonoid content, and the molecular regulatory network and regulatory gene expressionin apple fruit. Additionally, we conducted gene cloning analysis and analysis of the expression oftranscription factor MdMYB12, and have thus laid the foundation for different apple variety breedingwork using molecular biological breeding to cultivate higher flavonoid content in apple fruit. This paperis divided into the following content;
(1) W‘e utilized the fruit of ‘Malus sieversii\Malus robusta (Cam) R''ehd，.，Tsugaru and‘G’uoqing as experimental materials to determine the content difference and change trend ofchlorogenic acid and the total flavonoid content in peel and fruit in the young fruit period and in theincreasing stage by RP-HPLC and spectrophotometry analysis. In doing so, we discovered thatflavonoid content rules have no connection with the chlorogenic acid of apple polyphenols, thatflavonoid content is higher in the peel and that chlorogenic acid content is higher in fruit, that bothflavonoid and chlorogenic acid content are higher in the young fruit period than in the increasing period,and that their content is higher in wild-type apples than in cultivated-type apples. Through theexperiment, which used RP-HPLC to determine the total flavonoids and12kinds of flavonoid'compounds content difference and change trend in four types of apple peels and fruits in the young fruitstage and in the increasing stage, we found out that flavan3-ol and phloridzin are hi‘gher in Malussieversii\and that flavonol content is higher in the cultivated-type apple Jinqing. In the four varietiesof apple peels and fruits, the content of flavonoids decreases with the maturity of the fruit.
(2) In order to further prove that flavonoid content changes in relation to fruit development, we‘chose the fruit peel ''and fruit pulp of the cultivation type apple Tsugaru and the wild type Malus rockiiS’chneid. as experimental materials in five different developmental periods, and found that the fruit sizeof Tsugaru increases with the maturity of fruit enlargement, while th‘at of Malus rockii Schneid.‘exhibits no significant change with maturity. The contents of chlorogenic acid and flavonoids in Malusrockii Schneid: were far higher th‘an those in T，sugaru，and the change of chlorogenic acid content inapple fruit had no direct relation to fruit inlfation, while flavonoid content decreased with fruit inflation.Based on the analysis of content changes of12flavonoids in two varieties of apple, we found that thet''otal flavonoid content in Tsugaru was much lower than that in ‘Malus rockii Schneid.\but that theflavonoid content in the fruit peel was higher than that in ‘Malus rockii Schneid'.. The change of flavan3-ol content in ‘Malus rockii Schneid: was not obvious with fruit development, and showed no obviousdifference in the fruit pulp and peel, but the content decreased obviously in the early stage of fruit d''evelopment and had no significant change in later developmental stages in Tsugaru. The phloridzincontent in th'e 'Tsugaru peel decreased slowly with fruit growth, while there was no obvious change infruit pulp. The change in both the ‘Malus rockii Schneid,. fruit peel and pulp was the same; that is,phloridzin content decreased in the early fruit developmental stages and showed no obvious change asfruit matured.
(3) With the application of real-time fluorescent quantitative PCR，we analyzed the mRNAexpression differences of five traditional reference genes18SrRNA, ACTB, GAPDH, UBQ and TUB indifferent apple genotypes, different tissues and different developmental periods. Through geNormprogram analysis, we found that the expression stability of the five reference genes is different. UBQ isthe most stable in different genotypes and in gene expression analysis of different developmentalperiods. ACTB expression and UBQ expression are both stable in six different tissues. This study provesthat UBQ gene expression is relatively stable in experimental samples, and the UBQ gene is the firstchoice in apple gene expression analysis.
(4) We conducted RT-PCR analysis of11structural genes that regulate the anabolism offl'‘avonoids in 'Tsugaru and Mal'us rockii Schneid.. From this, we found that the expression quantity of10genes (PAL’ CHS，CHI，FHT，FLS，DFR，LAR，ANS，ANR，F3H) in ‘Malus rockii S，chneid.，whichh‘as higher levels of flavonoids than T，‘，sugaru，is more than in Tsugaru，while FGT expressionquantity showed a significant difference between the two kinds of apples. The gene expression quantitychanges with the growth of fruit, and is related to the changes in flavonoid content. The variation trendof CHI gene transcription is the most positively correlated with that of flavonoid content.
(5) With the bioinformatics analysis of the publi‘shed apple G'olden Delicious genome, wecloned the mdMYB12transcription factor f‘rom the Malus rockii Schneid,. fruit peel and found that thetranscription is correlated with several kinds of flavonoid substances based on fluorescence quantitativeanalysis of mdMYB12i'‘n 'Tsugaru and Malus rockii Schneid,.
(1)以‘新疆野生苹果’、‘八棱海棠’、‘津轻’和‘国庆’的果实为试材，用反相高效液相色谱法和分光光度法测定果皮、果肉在幼果期和膨大期绿原酸和总黄酮的含量差别和变化趋势，发现类黄酮含量变化与绿原酸变化没有必然联系，类黄酮在果皮中含量较高，而绿原酸在果肉中含量较高，二者在幼果期含量均高于膨大期，在野生型苹果中含量均高于栽培型苹果。用反相高效液相色谱法测定四种苹果果皮、果肉在幼果期和膨大期12种类黄酮化合物的含量差别和变化趋势的实验中发现，野生型苹果‘新疆野生苹果’中黄烷醇和根皮柑含量较高，栽培型苹果‘津轻’中黄酮醇含量较高，四种苹果的果皮果肉中，类黄酮物质含量均随着果实的成熟而降低。
(2)为了进一步探明类黄酮含量变化与果实发育进程的关系，以栽培型苹果‘津轻’和野生型苹果‘丽江山荆子’5个不同发育时期的果皮果肉为试验材料，对果实外观测定中发现，随着果实的发育成熟，‘津轻’果实明显膨大，而‘丽江山荆子’果实大小无明显变化。‘丽江山荆子’果实发育各个时期中绿原酸和总黄酮含量均远高于‘津轻’，苹果果实中绿原酸含量变化与果实膨大无直接关系，而总黄酮含量随着果实的膨大而下降。对果实中12种类黄酮含量变化的分析中发现，虽然‘津轻’总黄酮含量远远低于‘丽江山荆子’，但其果皮中黄酮醇类物质含量却高于‘丽江山荆子’；黄烷醇含量在‘丽江山荆子’中随着果实的发育无显著变化，且果肉和果皮中含量无显著差异，在‘津轻’果实发育前期黄烷醇含量迅速下降，后期含量无显著变化。‘津轻’果皮中根皮柑含量随着果实发育呈缓慢下降趋势，而在果肉中变化不显著，‘丽江山荆子’果皮果肉中的根皮柑含量变化趋势相同，均在果实发育前期有所下降，随着果实的发育成熟含量无显著变化。
(3)应用实时荧光定量PCR技术，分析5个传统内参基因/&SWW丄ACTB、GAPDH、UBQ和TUB在苹果不同基因型、不同组织以及果实不同发育时期的mRNA表达差异情况。经geNorm程序分析发现，5个内参基因的表达稳定性各异，在果实不同基因型以及不同发育时期的基因表达分析中最稳定；JCar和在6种不同组织中表达均稳定。本研究证明了在所有材料中均表达稳定，是研究苹果基因表达分析中首选的内参基因。
(4)对‘津轻’和‘丽江山荆子’中调控类黄酮合成代谢的11个结构基因的荧光定量PCR分析中发现，类黄酮含量较高的‘丽江山荆子’中PAL,CHS,CHI,FHT,FLS,DFR,LAR,ANS,ANR和朽丑这10个基因表达量均显著高于‘津轻’，FGT在两种苹果中表达量无明显差异。随着果实的发育进行，基因表达量存在差异，且与部分类黄酮含量变化呈正相关关系，其中CHI基因与类黄酮含量变化趋势相关性最显著。
(5)利用对已公布的‘金冠’苹果基因组的生物信息学分析，以‘丽江山荆子’果皮为实验材料克隆了 转录因子，对在‘津轻’和‘丽江山荆子’的荧光定量分析中发现，其转录与多种类黄酮物质存在正相关。
Flavonoids are the most important secondary metabolites in apple. They play an important role inapple nutritional quality, growth and development and disease resistance, as well as in apple taxonomyresearch. This study utilizes several types of higher flavonoid content in wild-type apples and lowerflavonoid content in cultivated-type apples as experimental mateirals, and takes the different genotypes,different parts and different developmental periods of apple fruits as research objects to study thechange rules of flavonoid content, and the molecular regulatory network and regulatory gene expressionin apple fruit. Additionally, we conducted gene cloning analysis and analysis of the expression oftranscription factor MdMYB12, and have thus laid the foundation for different apple variety breedingwork using molecular biological breeding to cultivate higher flavonoid content in apple fruit. This paperis divided into the following content;
(1) W‘e utilized the fruit of ‘Malus sieversii\Malus robusta (Cam) R''ehd，.，Tsugaru and‘G’uoqing as experimental materials to determine the content difference and change trend ofchlorogenic acid and the total flavonoid content in peel and fruit in the young fruit period and in theincreasing stage by RP-HPLC and spectrophotometry analysis. In doing so, we discovered thatflavonoid content rules have no connection with the chlorogenic acid of apple polyphenols, thatflavonoid content is higher in the peel and that chlorogenic acid content is higher in fruit, that bothflavonoid and chlorogenic acid content are higher in the young fruit period than in the increasing period,and that their content is higher in wild-type apples than in cultivated-type apples. Through theexperiment, which used RP-HPLC to determine the total flavonoids and12kinds of flavonoid'compounds content difference and change trend in four types of apple peels and fruits in the young fruitstage and in the increasing stage, we found out that flavan3-ol and phloridzin are hi‘gher in Malussieversii\and that flavonol content is higher in the cultivated-type apple Jinqing. In the four varietiesof apple peels and fruits, the content of flavonoids decreases with the maturity of the fruit.
(2) In order to further prove that flavonoid content changes in relation to fruit development, we‘chose the fruit peel ''and fruit pulp of the cultivation type apple Tsugaru and the wild type Malus rockiiS’chneid. as experimental materials in five different developmental periods, and found that the fruit sizeof Tsugaru increases with the maturity of fruit enlargement, while th‘at of Malus rockii Schneid.‘exhibits no significant change with maturity. The contents of chlorogenic acid and flavonoids in Malusrockii Schneid: were far higher th‘an those in T，sugaru，and the change of chlorogenic acid content inapple fruit had no direct relation to fruit inlfation, while flavonoid content decreased with fruit inflation.Based on the analysis of content changes of12flavonoids in two varieties of apple, we found that thet''otal flavonoid content in Tsugaru was much lower than that in ‘Malus rockii Schneid.\but that theflavonoid content in the fruit peel was higher than that in ‘Malus rockii Schneid'.. The change of flavan3-ol content in ‘Malus rockii Schneid: was not obvious with fruit development, and showed no obviousdifference in the fruit pulp and peel, but the content decreased obviously in the early stage of fruit d''evelopment and had no significant change in later developmental stages in Tsugaru. The phloridzincontent in th'e 'Tsugaru peel decreased slowly with fruit growth, while there was no obvious change infruit pulp. The change in both the ‘Malus rockii Schneid,. fruit peel and pulp was the same; that is,phloridzin content decreased in the early fruit developmental stages and showed no obvious change asfruit matured.
(3) With the application of real-time fluorescent quantitative PCR，we analyzed the mRNAexpression differences of five traditional reference genes18SrRNA, ACTB, GAPDH, UBQ and TUB indifferent apple genotypes, different tissues and different developmental periods. Through geNormprogram analysis, we found that the expression stability of the five reference genes is different. UBQ isthe most stable in different genotypes and in gene expression analysis of different developmentalperiods. ACTB expression and UBQ expression are both stable in six different tissues. This study provesthat UBQ gene expression is relatively stable in experimental samples, and the UBQ gene is the firstchoice in apple gene expression analysis.
(4) We conducted RT-PCR analysis of11structural genes that regulate the anabolism offl'‘avonoids in 'Tsugaru and Mal'us rockii Schneid.. From this, we found that the expression quantity of10genes (PAL’ CHS，CHI，FHT，FLS，DFR，LAR，ANS，ANR，F3H) in ‘Malus rockii S，chneid.，whichh‘as higher levels of flavonoids than T，‘，sugaru，is more than in Tsugaru，while FGT expressionquantity showed a significant difference between the two kinds of apples. The gene expression quantitychanges with the growth of fruit, and is related to the changes in flavonoid content. The variation trendof CHI gene transcription is the most positively correlated with that of flavonoid content.
(5) With the bioinformatics analysis of the publi‘shed apple G'olden Delicious genome, wecloned the mdMYB12transcription factor f‘rom the Malus rockii Schneid,. fruit peel and found that thetranscription is correlated with several kinds of flavonoid substances based on fluorescence quantitativeanalysis of mdMYB12i'‘n 'Tsugaru and Malus rockii Schneid,.
引文
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