外源植物生长调节物质对小白菜硫代葡萄糖苷的影响及相关合成基因表达研究
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摘要
硫代葡萄糖苷(β-thioglucoside N-hydroxysulfates)是十字花科植物中一组重要的富含氮硫的阴离子次生代谢物质。硫代葡萄糖苷已被证实与十字花科蔬菜的风味及营养成分、植物自我保护机制以及人类的身体健康有着密切的关系。植物中硫代葡萄糖苷的组分和含量主要由基因型决定,但同时受到植株生长环境条件及各种外源植物生长调节剂的影响。近年来,利用基因工程手段和外源物质调控植物体内硫代葡萄糖苷组分和含量已成为这方面研究的一个热点。调控十字花科蔬菜中硫代葡萄糖苷组分和含量从而提高其对人体的营养和保健作用,或提高植株应对病虫等外界胁迫的能力,对提高蔬菜的营养品质、产量及抗性具有重要的理论和实际意义。
本文以起源于我国的重要十字花科蔬菜小白菜(Brassica rapa L.ssp. chinensis var.communis)为试验材料,克隆了小白菜硫代葡萄糖苷合成关键基因并分析了其在不同品种及部位的表达水平,同时研究了油菜素内酯、多胺、杀虫剂、除草剂、氧化氮(NO)、硫化氢(H2S)及二氧化硫(SO2)等新型植物生长调节物质对小白菜硫代葡萄糖苷组分和含量及合成关键基因表达水平的影响,从生理层面和基因表达层面探讨了外源植物生长调节物质对小白菜硫代葡萄糖苷的调控作用。主要结果如下：
(1)研究了叶面喷施油菜素内酯和多胺对小白菜地上部硫苷组分和含量的影响。结果表明外源油菜素内酯处理增加了小白菜地上部脂肪族硫苷和总硫苷含量,减少了吲哚族硫苷含量,对芳香族硫苷的影响不大。油菜素内酯处理还影响了不同组分硫苷的相对含量。油菜素内酯对小白菜脂肪族硫苷含量的提高可能是通过提高植株光合作用促进了植株生长和氨基酸、葡萄糖等硫苷合成底物的积累所致。而油菜素内酯减少吲哚族硫苷积累的机制可能和油菜素内酯影响生长素运输及植株防御系统有关。外源多胺影响了小白菜地上部硫苷含量,其中0.1mM精胺(spm)和1.0mM亚精胺(spd)叶面喷施处理都增加了小白菜地上部脂肪族、芳香族和总硫苷含量。多胺提高小白菜中脂肪族硫苷含量可能和外源多胺影响了植物内源多胺和乙烯合成有关。外源油菜素内酯和多胺处理对小白菜地上部硫苷含量的影响都存在短期效应。
(2)研究了4种不同种类杀虫剂和5种不同种类除草剂对小白菜地上部硫苷组分和含量的影响。结果表明4种杀虫剂处理均大量增加了小白菜地上部脂肪族、吲哚族、芳香族和总硫苷含量。4种杀虫剂对小白菜地上部硫苷含量影响基本一致。杀虫剂处理后小白菜硫苷含量的增加可能是由于杀虫剂对小白菜植株造成了胁迫,从而诱导了硫苷-黑芥子酶系统启动,合成更多抗性物质来应对杀虫剂胁迫。5种除草剂处理均显著减少了小白菜地上部脂肪族硫苷含量,这可能与其抑制了脂肪族氨基酸合成有关。精禾草克处理虽然减少了脂肪族硫苷含量,但增加了吲哚族硫苷和芳香族硫苷含量,从而导致总硫苷含量的增加。除草剂处理对小白菜硫苷含量的影响存在短期效应。
(3)研究了叶面喷施外源供体形式的NO和H2G及SO2处理对小白菜硫苷组分和含量的影响。结果表明适当浓度的NO处理可以短期(5天)提高小白菜地上部硫苷含量,但对小白菜地上部硫苷的长期(10天)积累有负面作用。这可能是因为NO启动了植株体内抗氧化酶系统、防御基因表达等抗胁迫进程,从而导致硫苷等防御物质的短期积累。同时,NO处理也影响了小白菜根中的硫苷组分和含量。不同浓度和处理的NO对小白菜地上部和根中硫苷组分及含量的影响不同。H2S和Na2SO4对照处理均促进了小白菜地上部各族硫苷和总硫苷含量的积累,其中H2S存在着浓度效应,而Na2SO4处理不同浓度间没有差异。这表明叶面喷施MaHS形式的H2S和Na2SO4均能被小白菜快速吸收并转化成内源含硫代谢物。叶面喷施1.0mM和10mMNaHSO3两个浓度的SO2不仅不能作为小白菜生长所需的硫的供体,反而对其生长有毒害作用,减少了其硫苷积累。S02处理短时间内(5天)对小白菜地上部硫苷含量影响不大,但具有长期影响。NO和H2S处理对小白菜硫苷含量的影响随着时间延长而减弱或消失。
(4)运用同源克隆技术从小白菜中克隆到两条控制脂肪族硫苷合成的基因BcCYP83A1-1和BcCYP83A1-2,以及一条控制吲哚族和芳香族硫苷合成的基因BcCYP83B1,其编码区和编码氨基酸序列与拟南芥和大白菜同源基因具有高度的保守性,为通过基因工程改变硫苷组分和含量打下基础。
(5)运用实时荧光定量PCR (qRT-PCR)技术分析了小白菜硫苷合成关键基因(BcCYP83A1-1、BcCYP83A1-2和BcCYP83B1)在不同品种小白菜和小白菜不同部位的表达水平,以及外源多胺、杀虫剂和除草剂处理对其表达水平的影响。结果表明,BcCYP83A1主要在叶片和叶柄中表达,叶片和叶柄中BcCYP83A1表达量高于BcCYP83B1表达量；BcCYP83B1主要在根中表达,根中BcCYP83B1表达量高于BcCYP83A1表达量。小白菜硫苷合成关键基因在7个品种小白菜和小白菜3个部位中的表达水平与其相应的硫苷组分及含量一致。多胺、杀虫剂和除草剂等外源植物生长调节物质均诱导了小白菜叶片中硫苷合成基因的表达,且其表达水平和相应的硫苷组分和含量基本一致。
Glucosinolates (β-thioglucoside N-hydroxysulfates, GSs) are nitrogen-and sulfur-containing anionic natural products, one of the important secondary metabolites found in Brassicaceae plants. Glucosinolates have been proved to have strong relations with Brassica vegetable's flavors, tastes, nutrition and plant defense system. Glucosinolates also have beneficial effects on human health. The GSs in plant are mostly determined by plant genotypes, as well as affected by plant growth conditions, environment factors and plant growth regulators. Modifying the GS profiles and concentrations by genetic engineering and other factors, to get more beneficial GSs for human health, or to promote the capacity of plant defense to environment stress, has become a hot research topic in recent years. For the development of good quality vegetable crops and sustainable agriculture, it is of great importance to study the effects of plant growth regulator on plant GS metabolism.
This study uses the traditional and important Chinese cruciferous vegetable pak choi (Brassica rapa L. ssp. Chinensis var. communis) to study the effects of brassinosteroid (BR), polyamine (PA), insecticide, herbicide, nitric dioxide (NO), hydrogen sulfide (H2S) and sulfur dioxide (SO2) on plant GS accumulation. We also isolated the GSs biosynthetic gene BcCYP83A1and BcCYP83B1from pak choi and analyzed the gene expression pattern in leaves of seven pak choi cultivars and in three organs of pak choi. By analyzing the GS accumulation and gene expression, we explicated the effects and mechanisms of plant regulators on plant GSs synthesis. The following are the major results:
(1) The effects of exogenous BR and PA on plant growth and GS contents were studied in pak choi plants grown in nutrient solution. The results show BR increased the shoot aliphatic and total GSs contents, decreased the indole GS contents of pak choi. The increased aliphatic GSs by BR may due to the enhanced amino acid and glucose accumulation. And the decreasing of indole GSs by BR may have involvements with auxin metabolism and plant defense system.0.1mM supermine (spm) and1.0mM supermidine (spd) both increased the aliphatic GSs, aromatic GSs and total GSs. The effects of exogenous PA on GS contents of pak choi may involve in the inner PA and ethylene metabolisom.
(2) The effects of four insecticides and five herbicides on plant growth and GS contents were investigated in pak choi plants grown in nutrient solution. All the insecticides increased the shoot aliphatic GSs, aromatic GSs, indole GSs and total GSs of pak choi. It is speculated that the increasing GS contents were the strategy of plant response to insecticide stress. All herbicides decreased the aliphatic GS contents in shoots of pak choi, and these may due to the decreasing effects of herbicides on aliphatic amino acid. Besides of the decreasing of aliphatic GSs, quizalofop-p-ethyl also increased the aromatic GSs and indole GSs contents which leading to the increasing of total GSs. The influences of herbicides on pak choi GS contents were short term effects.
(3) The effects of exogenous NO, H2S and SO2on plant growth and GS contents were studied in pak choi plants grown in nutrient solution. The results show proper concentration of exogenous NO have short term (5day) positive effects on GS accumulation in shoots of pak choi, but have negative effects on GS contents in long term (10day). Exogenous NO also affected the profile and contents of GSs in roots of pak choi. Both H2S and Na2SO4treatments increased the aliphatic GSs, aromatic GSs, indole GSs and total GSs in shoots of pak choi. The GS contents were increased followed by the increasing of H2S concentrations. However, the effects of different concentration Na2SO4treatments on GSs had no difference. The decreased GS contents treated by1.0mM and10mM NaHSO3show SO2can not be used as sulfur source by pak choi, but have negative effects on plant growth and GS accumulation. The positive effects of H2S on GSs accumulation were short term effects.
(4) The aliphatic and indole GS biosynthetic gene BcCYP83Al and BcCYP83B1were isolated from pak choi. Two copies of BcCYP83A1, named BcCYP83A1-1and BcCYP83A1-2and one copy of BcCYP83B1were isolated. The predicted amino acid sequences of BcCYP83Al-1, BcCYP83A1-2and BcCYP83Bl shared high sequence identity of87.65,86.48and95.59%to the corresponding ones in Arabidopsis, and98.80,98.61and98.80%to the corresponding ones in Brassica pekinensis (Chinese cabbage), respectively. The cloning of the GS biosynthetic genes would have beneficial to modify the GS profiles and contents using genetic methods.
(5) Quantitative real-time PCR (qRT-PCR) were used to analysis the expression levels of GS biosynthetic genes in different cultivars. different organs and as affected by plant growth regulators. The results indicated that both BcCYP83A1and BcCYP83B1expressed in roots, leaves and petioles of pak choi, while the transcript abundances of BcCYP83A1were higher in leaves than in petioles and roots, whereas BcCYP83B1 showed higher abundances in roots. The expression levels of GS biosynthetic genes were consistent with the GS profile accumulation in leaves of seven cultivars and three organs. All the examined plant growth regulators affected the GS biosynthetic genes expression and the expression levels of GS biosynthetic genes were consistent with the GS profile accumulation as affected by plant growth regulators.
本文以起源于我国的重要十字花科蔬菜小白菜(Brassica rapa L.ssp. chinensis var.communis)为试验材料,克隆了小白菜硫代葡萄糖苷合成关键基因并分析了其在不同品种及部位的表达水平,同时研究了油菜素内酯、多胺、杀虫剂、除草剂、氧化氮(NO)、硫化氢(H2S)及二氧化硫(SO2)等新型植物生长调节物质对小白菜硫代葡萄糖苷组分和含量及合成关键基因表达水平的影响,从生理层面和基因表达层面探讨了外源植物生长调节物质对小白菜硫代葡萄糖苷的调控作用。主要结果如下：
(1)研究了叶面喷施油菜素内酯和多胺对小白菜地上部硫苷组分和含量的影响。结果表明外源油菜素内酯处理增加了小白菜地上部脂肪族硫苷和总硫苷含量,减少了吲哚族硫苷含量,对芳香族硫苷的影响不大。油菜素内酯处理还影响了不同组分硫苷的相对含量。油菜素内酯对小白菜脂肪族硫苷含量的提高可能是通过提高植株光合作用促进了植株生长和氨基酸、葡萄糖等硫苷合成底物的积累所致。而油菜素内酯减少吲哚族硫苷积累的机制可能和油菜素内酯影响生长素运输及植株防御系统有关。外源多胺影响了小白菜地上部硫苷含量,其中0.1mM精胺(spm)和1.0mM亚精胺(spd)叶面喷施处理都增加了小白菜地上部脂肪族、芳香族和总硫苷含量。多胺提高小白菜中脂肪族硫苷含量可能和外源多胺影响了植物内源多胺和乙烯合成有关。外源油菜素内酯和多胺处理对小白菜地上部硫苷含量的影响都存在短期效应。
(2)研究了4种不同种类杀虫剂和5种不同种类除草剂对小白菜地上部硫苷组分和含量的影响。结果表明4种杀虫剂处理均大量增加了小白菜地上部脂肪族、吲哚族、芳香族和总硫苷含量。4种杀虫剂对小白菜地上部硫苷含量影响基本一致。杀虫剂处理后小白菜硫苷含量的增加可能是由于杀虫剂对小白菜植株造成了胁迫,从而诱导了硫苷-黑芥子酶系统启动,合成更多抗性物质来应对杀虫剂胁迫。5种除草剂处理均显著减少了小白菜地上部脂肪族硫苷含量,这可能与其抑制了脂肪族氨基酸合成有关。精禾草克处理虽然减少了脂肪族硫苷含量,但增加了吲哚族硫苷和芳香族硫苷含量,从而导致总硫苷含量的增加。除草剂处理对小白菜硫苷含量的影响存在短期效应。
(3)研究了叶面喷施外源供体形式的NO和H2G及SO2处理对小白菜硫苷组分和含量的影响。结果表明适当浓度的NO处理可以短期(5天)提高小白菜地上部硫苷含量,但对小白菜地上部硫苷的长期(10天)积累有负面作用。这可能是因为NO启动了植株体内抗氧化酶系统、防御基因表达等抗胁迫进程,从而导致硫苷等防御物质的短期积累。同时,NO处理也影响了小白菜根中的硫苷组分和含量。不同浓度和处理的NO对小白菜地上部和根中硫苷组分及含量的影响不同。H2S和Na2SO4对照处理均促进了小白菜地上部各族硫苷和总硫苷含量的积累,其中H2S存在着浓度效应,而Na2SO4处理不同浓度间没有差异。这表明叶面喷施MaHS形式的H2S和Na2SO4均能被小白菜快速吸收并转化成内源含硫代谢物。叶面喷施1.0mM和10mMNaHSO3两个浓度的SO2不仅不能作为小白菜生长所需的硫的供体,反而对其生长有毒害作用,减少了其硫苷积累。S02处理短时间内(5天)对小白菜地上部硫苷含量影响不大,但具有长期影响。NO和H2S处理对小白菜硫苷含量的影响随着时间延长而减弱或消失。
(4)运用同源克隆技术从小白菜中克隆到两条控制脂肪族硫苷合成的基因BcCYP83A1-1和BcCYP83A1-2,以及一条控制吲哚族和芳香族硫苷合成的基因BcCYP83B1,其编码区和编码氨基酸序列与拟南芥和大白菜同源基因具有高度的保守性,为通过基因工程改变硫苷组分和含量打下基础。
(5)运用实时荧光定量PCR (qRT-PCR)技术分析了小白菜硫苷合成关键基因(BcCYP83A1-1、BcCYP83A1-2和BcCYP83B1)在不同品种小白菜和小白菜不同部位的表达水平,以及外源多胺、杀虫剂和除草剂处理对其表达水平的影响。结果表明,BcCYP83A1主要在叶片和叶柄中表达,叶片和叶柄中BcCYP83A1表达量高于BcCYP83B1表达量；BcCYP83B1主要在根中表达,根中BcCYP83B1表达量高于BcCYP83A1表达量。小白菜硫苷合成关键基因在7个品种小白菜和小白菜3个部位中的表达水平与其相应的硫苷组分及含量一致。多胺、杀虫剂和除草剂等外源植物生长调节物质均诱导了小白菜叶片中硫苷合成基因的表达,且其表达水平和相应的硫苷组分和含量基本一致。
Glucosinolates (β-thioglucoside N-hydroxysulfates, GSs) are nitrogen-and sulfur-containing anionic natural products, one of the important secondary metabolites found in Brassicaceae plants. Glucosinolates have been proved to have strong relations with Brassica vegetable's flavors, tastes, nutrition and plant defense system. Glucosinolates also have beneficial effects on human health. The GSs in plant are mostly determined by plant genotypes, as well as affected by plant growth conditions, environment factors and plant growth regulators. Modifying the GS profiles and concentrations by genetic engineering and other factors, to get more beneficial GSs for human health, or to promote the capacity of plant defense to environment stress, has become a hot research topic in recent years. For the development of good quality vegetable crops and sustainable agriculture, it is of great importance to study the effects of plant growth regulator on plant GS metabolism.
This study uses the traditional and important Chinese cruciferous vegetable pak choi (Brassica rapa L. ssp. Chinensis var. communis) to study the effects of brassinosteroid (BR), polyamine (PA), insecticide, herbicide, nitric dioxide (NO), hydrogen sulfide (H2S) and sulfur dioxide (SO2) on plant GS accumulation. We also isolated the GSs biosynthetic gene BcCYP83A1and BcCYP83B1from pak choi and analyzed the gene expression pattern in leaves of seven pak choi cultivars and in three organs of pak choi. By analyzing the GS accumulation and gene expression, we explicated the effects and mechanisms of plant regulators on plant GSs synthesis. The following are the major results:
(1) The effects of exogenous BR and PA on plant growth and GS contents were studied in pak choi plants grown in nutrient solution. The results show BR increased the shoot aliphatic and total GSs contents, decreased the indole GS contents of pak choi. The increased aliphatic GSs by BR may due to the enhanced amino acid and glucose accumulation. And the decreasing of indole GSs by BR may have involvements with auxin metabolism and plant defense system.0.1mM supermine (spm) and1.0mM supermidine (spd) both increased the aliphatic GSs, aromatic GSs and total GSs. The effects of exogenous PA on GS contents of pak choi may involve in the inner PA and ethylene metabolisom.
(2) The effects of four insecticides and five herbicides on plant growth and GS contents were investigated in pak choi plants grown in nutrient solution. All the insecticides increased the shoot aliphatic GSs, aromatic GSs, indole GSs and total GSs of pak choi. It is speculated that the increasing GS contents were the strategy of plant response to insecticide stress. All herbicides decreased the aliphatic GS contents in shoots of pak choi, and these may due to the decreasing effects of herbicides on aliphatic amino acid. Besides of the decreasing of aliphatic GSs, quizalofop-p-ethyl also increased the aromatic GSs and indole GSs contents which leading to the increasing of total GSs. The influences of herbicides on pak choi GS contents were short term effects.
(3) The effects of exogenous NO, H2S and SO2on plant growth and GS contents were studied in pak choi plants grown in nutrient solution. The results show proper concentration of exogenous NO have short term (5day) positive effects on GS accumulation in shoots of pak choi, but have negative effects on GS contents in long term (10day). Exogenous NO also affected the profile and contents of GSs in roots of pak choi. Both H2S and Na2SO4treatments increased the aliphatic GSs, aromatic GSs, indole GSs and total GSs in shoots of pak choi. The GS contents were increased followed by the increasing of H2S concentrations. However, the effects of different concentration Na2SO4treatments on GSs had no difference. The decreased GS contents treated by1.0mM and10mM NaHSO3show SO2can not be used as sulfur source by pak choi, but have negative effects on plant growth and GS accumulation. The positive effects of H2S on GSs accumulation were short term effects.
(4) The aliphatic and indole GS biosynthetic gene BcCYP83Al and BcCYP83B1were isolated from pak choi. Two copies of BcCYP83A1, named BcCYP83A1-1and BcCYP83A1-2and one copy of BcCYP83B1were isolated. The predicted amino acid sequences of BcCYP83Al-1, BcCYP83A1-2and BcCYP83Bl shared high sequence identity of87.65,86.48and95.59%to the corresponding ones in Arabidopsis, and98.80,98.61and98.80%to the corresponding ones in Brassica pekinensis (Chinese cabbage), respectively. The cloning of the GS biosynthetic genes would have beneficial to modify the GS profiles and contents using genetic methods.
(5) Quantitative real-time PCR (qRT-PCR) were used to analysis the expression levels of GS biosynthetic genes in different cultivars. different organs and as affected by plant growth regulators. The results indicated that both BcCYP83A1and BcCYP83B1expressed in roots, leaves and petioles of pak choi, while the transcript abundances of BcCYP83A1were higher in leaves than in petioles and roots, whereas BcCYP83B1 showed higher abundances in roots. The expression levels of GS biosynthetic genes were consistent with the GS profile accumulation in leaves of seven cultivars and three organs. All the examined plant growth regulators affected the GS biosynthetic genes expression and the expression levels of GS biosynthetic genes were consistent with the GS profile accumulation as affected by plant growth regulators.
引文
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