类黄酮调控基因AtMYB12的异源表达及其对植物抗病性的贡献
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摘要
生物类黄酮（bioflavonoids）是自然界存在的一大类酚类物质次生代谢产物，在自然界中广泛存在并具有抗氧化、抑制脂质过氧化反应、预防心血管疾病、防癌等作用。当前满足市场需求的途径主要通过依赖化学方法从天然材料如银杏叶、洋葱、柠檬、各种中药材等中提取。
本研究利用番茄果实特异性表达启动子E4、PG、2A12驱动拟南芥中生物类黄酮的特异性调控因子AtMYB12基因在番茄中表达，获得一系列富含类黄酮（芦丁、山奈酚芸香糖苷和柚皮素查尔酮）的番茄转基因株系，进一步验证了拟南芥中生物类黄酮的特异性调控因子AtMYB12基因的功能，比较了不同启动子驱动AtMYB12基因调控类黄酮累积的差异，为进一步利用番茄作为生物反应器生产类黄酮组分物质提供参考。
首先，本研究利用不同果实特异性启动子驱动AtMYB12基因转化番茄品种并研究了其调控合成类黄酮类物质的能力。采用pX6-GFP载体、高保真PCR技术，通过分步克隆策略，分别构建了植物表达载体pX6-E4::AtMYB12、pX6-PG::AtMYB12和pX6-2A12::AtMYB12，并导入农杆菌工程菌株AGL1。通过叶盘法转化番茄品种圣女果或俏美人，分别获得阳性转基因植株。高压液相色谱（HPLC）法检测转基因番茄果实材料，结果显示三个载体pX6-E4::AtMYB12、pX6-PG::AtMYB12和pX6-2A12::AtMYB12的转基因果实果皮中均可检测到类黄酮类物质的提高，其中柚皮素查尔酮在转基因果皮中含量显著提高。三个不同启动子能够导致基因表达量和转基因番茄果实中物质的含量不同，E4启动子能够调控AtMYB12基因在果肉中高效表达，并引起下游参与类黄酮合成基因的上调表达，PG和2A12启动子能够调控AtMYB12基因在果皮中表达，同时引起下游参与类黄酮合成基因的上调表达。
其次，利用烟草遗传转化系统研究了其它预测类黄酮或咖啡酰奎尼酸合成相关基因的功能。将AtMYB12基因的同源基因AtMYB11在转入三生烟草后也起到与AtMYB12基因相同的效果，提高了转基因烟草植株中类黄酮类物质的含量，但是也同样没有激活下游次生代谢产物咖啡酰奎尼酸的合成。转基因烟草植株中，单独超量表达NtHCT、NtHQT、NtC3H均没有引起下游次生代谢产物咖啡酰奎尼酸的合成，但是NtHQT基因的超量表达后的HPLC结果显示转基因烟草中芦丁的含量有所下降，推测可能由于新的未知物质的合成而阻碍或降低了芦丁的合成。
进而分析了类黄酮累积的烟草转基因株系的抗病性状，发现转AtMYB12基因的烟草植株中由于芦丁含量的提高而导致转基因烟草植株对烟草青枯病菌、烟草炭疽病菌、烟草赤星病菌的抗性提高，但转基因植株对供试PVY病毒株系表现更加敏感。转基因烟草植株的抗病反应提高，主要是由于转基因烟草在接种病原后抗病相关基因的迅速高效表达，从而引起植株对多种病害的抗性。而且转基因烟草提取物的平板抑菌试验也进一步证实了芦丁等类黄酮物质对细菌和真菌生长具有一定的抑制效果。转基因的番茄果实由于芦丁含量的提高也对番茄灰霉病表现抗性，同时芦丁的平板抑菌试验进一步证实了芦丁对番茄灰霉病的抗性效果。
生物类黄酮作为植物的一大类次生代谢产物的总称，不仅是植物生长和防御反应中所必须的，同时也对人体有非常显著且良好的效果，我们通过栽培型小果番茄和重要经济作物烟草作为生物反应器生产类黄酮生物制品，得到具有无选择标记的番茄新品种和高含类黄酮和咖啡酰奎尼酸的烟草新品种，为进行品种培育和产业化加工奠定基础。同时芦丁作为一种植物的次生代谢产物，可以作为烟草植株抗病反应的激发子，诱导调控烟草植株对多种病害的抗性，为培育高产优质抗病的烟草新品种奠定了基础。同时，也为结合类黄酮的生物发酵生产和与其它杀菌剂成份混合调制生产生物源农药提供思路。
One subclass of plant polyphenols, bioflavonoids, are ubiquitous in the plant kingdomand have many diverse health-promoting functions including anti-allergic, anti-cancer,antioxidant, anti-inflammatory and anti-viral properties when consumed in foods of plantorigin. Currently, the extract pathway of flavonoids to satisfy consume is dependent onchemical methods to extract from ginkgo biloba leaves, onion and lemon etc.
In this study, by combination fruit specific promoter E4, PG,2A12with Arabidopsisthaliana flavonol-specific transcription factor AtMYB12gene, we got many tomato transgeniclines enriched flavonoids, including rutin, kaempferol rutinoside and naringenin chalcone.The results further confirmed that transcription factor AtMYB12gene could regulate theflavonoids synthesis, and compared the differences of flavonoids accumulation controlled bydifferent fruit specific promoters. The studies of transgenic tomato provide references forproducing flavonoids using tomato as bioreactors.
Firstly, we constructed the plant expression vector pX6-E4::AtMYB12,pX6-PG::AtMYB12and pX6-2A12::AtMYB12by replacing the GFP fragment with DNAfragment either of E4, PG or2A12promoters and AtMYB12in pX6-GFP vector respectively.They were all introduced into Agrobacterium strain AGL1and transformed into two cherrytomato cultivars, Qiaomeiren and Shengnvguo via leaf disc method. After identified by PCR,we got the positive transgenic tomato lines. We detected flavonoids in all three transgenictomato fruits peels by HPLC, and the content of naringenin chalcone was shown todramatically increase among the transgenic tomato fruits peel. Three different tomato fruitspecific promoters regulate AtMYB12gene expression in different positions, and induce thechanges of flavonoids content. The promoter of E4controls the expression of AtMYB12mainly in tomato fresh, and up-regulated expression of related genes involved in flavonoidssynthesis, such as CHS gene (chalcone synthase). The promoters PG and2A12control theexpression of AtMYB12in tomato peel, and induce the up-regulated expression of relatedgenes involved in flavonoids synthesis, such as CHS gene, PAL gene (phenylalanineammonium lyase) and GT gene (flavonol-3-glucosyltransferase).
Second, we have charactered the functions for some other regulator genes for flavonol orcaffeoyl quinic acid synthesis. AtMYB11gene, one of the homologous gene of AtMYB12,was over-expressed in Samsun tobacco, leading to high-level accumulation of polyphenoliccompounds, especially flavonols. The same effect, AtMYB11gene failed to induce theaccumulation of caffeoyl quinic acid (CQA). Over-expression of the genes NtHCT, NtHQTand NtC3H respectively, which prdicated to participate in CQA synthesis, all failed to inducethe accumulation of CQA. To our supprise, over-expression of NtHQT reduced the content ofrutin which presumed to synthesized some novel contents.
Further, a set of pathogenic assay have been tested on transgenic Samsun tobacco carryingAtMYB12dedrived under35S promoter. The results was shown tha the AtMYB12-expressiontransgenic tobacco lines enhance the plant defense responses against pathogens: bacteria suchas Ralstonia solanacearum; fungi such as Colletotrichum destructivum and Alternariaalternate; and insects such as aphid and whitefly because of the high-level accumulation ofthe flavanol rutin, but more sensitivity to potato virus Y (PVY). The enhancement of defenseresponses in transgenic tobacco plants is most attributed to the up-regulated expression ofresistance related genes. The result is further confirmed by the plate culture plused withtransgenic tobacco leaves extracts. The transgenic tomato fruits also enhance the defenseresponse against Botrtytis cinerea because of the accumulation of rutin in tomato fruits, andthe plate culture with different contents of rutin further confirms the result.
Bioflavonoids is a large kind of plant metabolism productions ubiquitous in the plantkingdom, flavonoids play a critical role in preventing human for disease and have evolved asa protective mechanism for a variety of different plants. We use small-fruit-type tomato andmajor economic crop tobacco as bioreactor to product bioflavonoids, to get marker-freetomato and tobacco lines enriched flavonoids and CQA. At the same time, rutin as a proudmember of the flavonoid family, could be an activator to enhance the reaction of plant diseaseresistance. These results are formed a good foundation to support the further study oncultivating high yield and good quality of tobacco variety, especially for disease resistanceplant. Aslo it also open the opportunity to develop the bio-pesticide by combining thebioreactor to product flavonol and adjusted with additive pesticide compounding in industry.
本研究利用番茄果实特异性表达启动子E4、PG、2A12驱动拟南芥中生物类黄酮的特异性调控因子AtMYB12基因在番茄中表达，获得一系列富含类黄酮（芦丁、山奈酚芸香糖苷和柚皮素查尔酮）的番茄转基因株系，进一步验证了拟南芥中生物类黄酮的特异性调控因子AtMYB12基因的功能，比较了不同启动子驱动AtMYB12基因调控类黄酮累积的差异，为进一步利用番茄作为生物反应器生产类黄酮组分物质提供参考。
首先，本研究利用不同果实特异性启动子驱动AtMYB12基因转化番茄品种并研究了其调控合成类黄酮类物质的能力。采用pX6-GFP载体、高保真PCR技术，通过分步克隆策略，分别构建了植物表达载体pX6-E4::AtMYB12、pX6-PG::AtMYB12和pX6-2A12::AtMYB12，并导入农杆菌工程菌株AGL1。通过叶盘法转化番茄品种圣女果或俏美人，分别获得阳性转基因植株。高压液相色谱（HPLC）法检测转基因番茄果实材料，结果显示三个载体pX6-E4::AtMYB12、pX6-PG::AtMYB12和pX6-2A12::AtMYB12的转基因果实果皮中均可检测到类黄酮类物质的提高，其中柚皮素查尔酮在转基因果皮中含量显著提高。三个不同启动子能够导致基因表达量和转基因番茄果实中物质的含量不同，E4启动子能够调控AtMYB12基因在果肉中高效表达，并引起下游参与类黄酮合成基因的上调表达，PG和2A12启动子能够调控AtMYB12基因在果皮中表达，同时引起下游参与类黄酮合成基因的上调表达。
其次，利用烟草遗传转化系统研究了其它预测类黄酮或咖啡酰奎尼酸合成相关基因的功能。将AtMYB12基因的同源基因AtMYB11在转入三生烟草后也起到与AtMYB12基因相同的效果，提高了转基因烟草植株中类黄酮类物质的含量，但是也同样没有激活下游次生代谢产物咖啡酰奎尼酸的合成。转基因烟草植株中，单独超量表达NtHCT、NtHQT、NtC3H均没有引起下游次生代谢产物咖啡酰奎尼酸的合成，但是NtHQT基因的超量表达后的HPLC结果显示转基因烟草中芦丁的含量有所下降，推测可能由于新的未知物质的合成而阻碍或降低了芦丁的合成。
进而分析了类黄酮累积的烟草转基因株系的抗病性状，发现转AtMYB12基因的烟草植株中由于芦丁含量的提高而导致转基因烟草植株对烟草青枯病菌、烟草炭疽病菌、烟草赤星病菌的抗性提高，但转基因植株对供试PVY病毒株系表现更加敏感。转基因烟草植株的抗病反应提高，主要是由于转基因烟草在接种病原后抗病相关基因的迅速高效表达，从而引起植株对多种病害的抗性。而且转基因烟草提取物的平板抑菌试验也进一步证实了芦丁等类黄酮物质对细菌和真菌生长具有一定的抑制效果。转基因的番茄果实由于芦丁含量的提高也对番茄灰霉病表现抗性，同时芦丁的平板抑菌试验进一步证实了芦丁对番茄灰霉病的抗性效果。
生物类黄酮作为植物的一大类次生代谢产物的总称，不仅是植物生长和防御反应中所必须的，同时也对人体有非常显著且良好的效果，我们通过栽培型小果番茄和重要经济作物烟草作为生物反应器生产类黄酮生物制品，得到具有无选择标记的番茄新品种和高含类黄酮和咖啡酰奎尼酸的烟草新品种，为进行品种培育和产业化加工奠定基础。同时芦丁作为一种植物的次生代谢产物，可以作为烟草植株抗病反应的激发子，诱导调控烟草植株对多种病害的抗性，为培育高产优质抗病的烟草新品种奠定了基础。同时，也为结合类黄酮的生物发酵生产和与其它杀菌剂成份混合调制生产生物源农药提供思路。
One subclass of plant polyphenols, bioflavonoids, are ubiquitous in the plant kingdomand have many diverse health-promoting functions including anti-allergic, anti-cancer,antioxidant, anti-inflammatory and anti-viral properties when consumed in foods of plantorigin. Currently, the extract pathway of flavonoids to satisfy consume is dependent onchemical methods to extract from ginkgo biloba leaves, onion and lemon etc.
In this study, by combination fruit specific promoter E4, PG,2A12with Arabidopsisthaliana flavonol-specific transcription factor AtMYB12gene, we got many tomato transgeniclines enriched flavonoids, including rutin, kaempferol rutinoside and naringenin chalcone.The results further confirmed that transcription factor AtMYB12gene could regulate theflavonoids synthesis, and compared the differences of flavonoids accumulation controlled bydifferent fruit specific promoters. The studies of transgenic tomato provide references forproducing flavonoids using tomato as bioreactors.
Firstly, we constructed the plant expression vector pX6-E4::AtMYB12,pX6-PG::AtMYB12and pX6-2A12::AtMYB12by replacing the GFP fragment with DNAfragment either of E4, PG or2A12promoters and AtMYB12in pX6-GFP vector respectively.They were all introduced into Agrobacterium strain AGL1and transformed into two cherrytomato cultivars, Qiaomeiren and Shengnvguo via leaf disc method. After identified by PCR,we got the positive transgenic tomato lines. We detected flavonoids in all three transgenictomato fruits peels by HPLC, and the content of naringenin chalcone was shown todramatically increase among the transgenic tomato fruits peel. Three different tomato fruitspecific promoters regulate AtMYB12gene expression in different positions, and induce thechanges of flavonoids content. The promoter of E4controls the expression of AtMYB12mainly in tomato fresh, and up-regulated expression of related genes involved in flavonoidssynthesis, such as CHS gene (chalcone synthase). The promoters PG and2A12control theexpression of AtMYB12in tomato peel, and induce the up-regulated expression of relatedgenes involved in flavonoids synthesis, such as CHS gene, PAL gene (phenylalanineammonium lyase) and GT gene (flavonol-3-glucosyltransferase).
Second, we have charactered the functions for some other regulator genes for flavonol orcaffeoyl quinic acid synthesis. AtMYB11gene, one of the homologous gene of AtMYB12,was over-expressed in Samsun tobacco, leading to high-level accumulation of polyphenoliccompounds, especially flavonols. The same effect, AtMYB11gene failed to induce theaccumulation of caffeoyl quinic acid (CQA). Over-expression of the genes NtHCT, NtHQTand NtC3H respectively, which prdicated to participate in CQA synthesis, all failed to inducethe accumulation of CQA. To our supprise, over-expression of NtHQT reduced the content ofrutin which presumed to synthesized some novel contents.
Further, a set of pathogenic assay have been tested on transgenic Samsun tobacco carryingAtMYB12dedrived under35S promoter. The results was shown tha the AtMYB12-expressiontransgenic tobacco lines enhance the plant defense responses against pathogens: bacteria suchas Ralstonia solanacearum; fungi such as Colletotrichum destructivum and Alternariaalternate; and insects such as aphid and whitefly because of the high-level accumulation ofthe flavanol rutin, but more sensitivity to potato virus Y (PVY). The enhancement of defenseresponses in transgenic tobacco plants is most attributed to the up-regulated expression ofresistance related genes. The result is further confirmed by the plate culture plused withtransgenic tobacco leaves extracts. The transgenic tomato fruits also enhance the defenseresponse against Botrtytis cinerea because of the accumulation of rutin in tomato fruits, andthe plate culture with different contents of rutin further confirms the result.
Bioflavonoids is a large kind of plant metabolism productions ubiquitous in the plantkingdom, flavonoids play a critical role in preventing human for disease and have evolved asa protective mechanism for a variety of different plants. We use small-fruit-type tomato andmajor economic crop tobacco as bioreactor to product bioflavonoids, to get marker-freetomato and tobacco lines enriched flavonoids and CQA. At the same time, rutin as a proudmember of the flavonoid family, could be an activator to enhance the reaction of plant diseaseresistance. These results are formed a good foundation to support the further study oncultivating high yield and good quality of tobacco variety, especially for disease resistanceplant. Aslo it also open the opportunity to develop the bio-pesticide by combining thebioreactor to product flavonol and adjusted with additive pesticide compounding in industry.
引文
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