春大豆种子田间劣变抗性的评价及抗性机理的研究
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摘要
春大豆种子在发育和成熟过程中常会受到高温高湿胁迫,导致种子发生田间劣变,造成活力和发芽率降低、出苗困难等。春大豆种子劣变问题已成为我国南方春大豆生产发展的重大障碍。目前,国内外有关春大豆种子劣变的研究主要集中在收获期和贮藏期间引起大豆种子劣变的内外诱因,劣变过程中细胞学、遗传学和生理生化等方面,而对春大豆种子田间劣变抗性的分子基础缺乏研究。因此,急需利用我国南方野生大豆和春大豆资源中具不同种子田间劣变抗性程度的种质,深入开展其劣变抗性的分子基础方面的研究。本研究通过温箱蚀化模拟田间高温高湿条件对我国南方92份春大豆种质种子田间劣变抗性进行评价,并对鉴定出的种子田间劣变抗性品种湘豆三号和不抗品种宁镇一号处于生理成熟期(R7期)的种子在田间高温高湿胁迫下的形态、生理生化和蛋白质变化进行了研究,以期揭示春大豆种子劣变性和劣变抗性的基础,为我国南方春大豆种子田间劣变抗性品种的选育提供理论基础和实践指导。获得的主要研究结果如下：
1.对92份我国南方春大豆种质采用温箱蚀化模拟田间高温高湿条件进行胁迫,发现种质间种子田间劣变抗性存在显著差异；鉴定出种子田间劣变抗性种质湘豆三号和不抗种质宁镇一号,并确定了种子发芽率、活力指数和简易活力指数三个指标相结合可以作为种子田间劣变鉴定的标准。
2.高温高湿胁迫,对发育中的湘豆三号种子细胞的超微结构影响不明显；而对宁镇一号种子细胞的超微结构影响较大,表现为：种子细胞中叶绿体和蛋白体的数量不断减少和体积减小,细胞核和核仁的形状变得不规则,液泡和油体所占的面积更大,细胞壁变薄,淀粉粒数目减少,线粒体和光滑内质网数目增多。
3.高温高湿胁迫下,湘豆三号处于发育过程中的种子的相对电导率和MDA显著低于宁镇一号种子的；2个品种种子中ROS皆大量积累,打破了原有的氧化还原平衡体系,但湘豆三号种子中CAT活性和ASA含量显著高于宁镇一号种子的；2个品种种子中的化学成分都受到影响,其中：湘豆三号种子中蛋白和油脂的含量降低的幅度小于宁镇一号种子的；宁镇一号种子中内源乙烯含量比湘豆三号种子中内源乙烯的含量增加得更明显；2个品种叶片和种子中叶绿素含量、光化学效率均显著下降,其中宁镇一号下降的幅度更加明显。湘豆三号种子中NH4+、NO2-、NO3含量降低,而宁镇一号种子中的NH4+、NO3-含量升高,NO2-含量降低。
4.高温高湿胁迫下,采用GC-MS技术对湘豆三号和宁镇一号处于发育过程中的种子进行代谢谱分析,共检测到46种有差异表达的代谢产物,其中从抗性品种湘豆三号种子中检测出差异表达的代谢产物共34种,从不抗品种宁镇一号种子中检测出31种。这些代谢物质共涉及到光合作用、三羧酸循环、氨基酸合成途径、次生代谢产物合成途径、磷酸戊糖途径和光呼吸等6大代谢途径。高温高湿胁迫导致湘豆三号种子中卡尔文循环增强而宁镇一号种子中卡尔文循环减弱；导致宁镇一号种子中磷酸戊糖途径增强,而对湘豆三号种子中磷酸戊糖途径没有明显影响；诱导宁镇一号种子中光呼吸作用增强而对湘豆三号种子中光呼吸作用影响不显著；导致湘豆三号种子中TCA途径减弱,而宁镇一号种子中TCA途径显著增强。此外,高温高湿胁迫会导致湘豆三号和宁镇一号种子处于发育过程中的中氨基酸和次生代谢产物合成增强。
5.通过对高温高湿胁迫处理的宁镇一号和湘豆三号处于发育过程中的种子进行双向电泳及质谱分析,成功鉴定出87个差异表达蛋白点,这些差异蛋白点共涉及到15个代谢途径和细胞过程。在湘豆三号处于发育过程中的种子中共检测到45个差异表达蛋白点,涉及33个差异表达蛋白；其中21个蛋白点上调表达,24个蛋白点下调表达,涉及13个代谢途径和细胞过程,分别为碳素代谢、信号转导、蛋白合成、光合作用、蛋白折叠和组装、能量代谢、细胞修复和防御、脂代谢、氨基酸代谢、转录调控、次生代谢产物合成途径、蛋白降解及转运蛋白；在宁镇一号处于发育过程中的种子中共检测到42个差异表达蛋白点,涉及30个差异表达蛋白；其中22个蛋白点上调表达,20蛋白点下调表达,涉及13个代谢途径和细胞过程,分别为碳素代谢、信号转导、蛋白合成、光合作用、蛋白折叠和组装、能量代谢、细胞修复和防御、脂代谢、氨基酸代谢、氮素代谢、细胞周期、转录调控以及次生代谢产物合成途径。
6.综合形态、生理生化和蛋白质组学研究结果,发现：高温高湿胁迫下,湘豆三号处于发育过程中的种子中增强田间劣变抗性的代谢途径和细胞过程有：①光合作用的加强；②蛋白折叠和组装的加强；③蛋白降解的下降；④次生代谢物合成的增强；⑤能量消耗的减少；⑥ASA合成的增加。导致种子田间劣变性的代谢途径和细胞过程有：①CO2固定能力下降,碳素同化减少；②细胞壁合成降低；③蛋白含量降低；④油脂含量的降低；⑤抗氧化酶类的活性减弱。随着胁迫时间的延长,增强种子田间劣变抗性的因素逐渐增强而导致种子田间劣变性的因素不断减弱,最终前者逐渐占优势,结果发育过程中的湘豆三号种子抗种子劣变性增强。
高温高湿胁迫下,宁镇一号处于发育过程中的种子中增强田间劣变抗性的代谢途径和细胞过程有：①精氨酸含量的增加,②谷氨酰胺合成途径的增强,③CO2吸收的增强,④光呼吸的增强,⑤次生代谢物合成的增强；⑥磷酸戊糖途径的加强；⑦消除ROS能力和DNA修复的水平的提升。导致田间劣变性的代谢途径和细胞过程有：①细胞周期稳定性的降低；②光合作用降低；③蛋白绑定和折叠的降低；④能量的过度消耗；⑤蛋白含量降低；⑥油脂含量的降低；⑦氮素吸收的减弱。随着胁迫时间的延长,导致种子田间劣变性的因素不断增强而增强种子田间劣变抗性的因素不断减弱,最终前者逐渐占优势,结果宁镇一号的大豆种子在发育过程中的发生种子劣变。
High temperature and humidity (HTH) stress during soybean seed development and maturity in the field easily leads seed to pre-harvest deterioration. It often causes reduced vigor and germination rate, et al.This phenomenon has become a major obstacle of spring soybean production.At present, Related with seed deterioration research mainly focuses on the reasons that caused seed deterioration, cytology and genetics in the process of deterioration. The researches on cytological, physiological and molecular basis about seed deterioration and deterioration resistance are lack.
1.92of Spring soybean landraces and cultivars from South China were used to evaluate seed deterioration resistance by incubating weathering treatment. One germplasm (Xiangdou NO.3) identified to resist to pre-harvest seed deterioration and one (Ningzhen NO.1) identified not to resist pre-harvest seed deteriorations were screened out.
2. Transmission electron micrograph analysis indicated that HTH stress resulted in some ultrastructure changes in the cells of the developing seed of cv. Ningzhen No.1and the change was not significant in the cells of the developing seed of cv. Xiangdou No.3.In the cells of the developing seed of cv. Ningzhen No.1, the cells of the developing seed tended to exhibit thinner cell wall, smaller chloroplasts and protein bodies, more irregular cell nucleus and nuclear core, and bigger vacuoles and lipid bodies under HTH stress. Additionally, under HTH stress it was found that the decrease of starch grains in number and the increase of mitochondria and smooth endoplasmic reticula in number in the cell of the seed.
3. It was found that the HTH stress caused heavy accumulated ROS in the cells of the developing seed and breaked the original redox balance system;In the seed of cv. Ningzhen No.1showed higher electrolyte leakage and the content of MDA than cv. Xiangdou No.3.It showed the lower activity of CAT and the content of ASA than cv. Xiangdou No. 3,HTH stress was also caused change of chemical composition of the seed.The content of protein and oil were more decrease of cv. Ningzhen No.1than cv. Xiangdou No.3;the content of Endogenous ethylene of cv. Ningzhen No.1higher than cv. Xiangdou No.3; The chlorophyll content in the leaves and seeds and Fv/Fm were all significantly decreased.; Compared to the control, the content of NO3-and NH4+in HTH-stressed developing seed of cv. Ningzhen No.1were significantly increased while the NO2-content was significantly decreased, the content ofNO3-,NO2-,NH4+in HTH-stressed developing seed of cv. Xiangdou No.3were all decrease.
4. Metabolic profiling based on GC-MS showed46differentially expressed metabolites. These metabolites were classified into6categories.There were8kinds of sugar,4of sugar alcohol and1kinds of sugar acids,28.3%of the total that related to calvin cycle,With the Calvin cycle related substances significantly upregulated expression in cv. Xiangdou No.3and significantly downregulated expression.in cv. Ningzhen No.1;To detect the5kinds of PPP pathway of sugar phosphate material from cv. Ningzhen No.l, in HTH-stressed,they were upregulated expression;Detection of Phosphoglycolate, Ser and Gly3that they were related to photorespiratory patyway and were upregulated; There were6kinds material of TCA pathway,these material were upregulated in cv. Ningzhen No.1and were downregulated.in cv. Xiangdou No.3;14kinds of amino acids were detected, accounting for30.4%of the total number of detected objects in the two species were significantly or very significantly up-regulated expression; Detected7kinds of substances associated with secondary metabolite biosynthesis pathway, significantly up-regulated expression in the two species.
5. Compared the proteome composition of developing seed of cv. Xiangdou NO.3and cv. Ningzhen No.1at different HTH stress time points with their corresponding controls by2-DE.87protein spots were found to be differentially expressed and successfully identified by MALDI-TOF MS:45protein spots were found to be differentially expressed and successfully identified in cv. Xiangdou NO.3. there were21spots upregulated and24spots downregulated,These proteins were involved in13of cellular responses and metabolic processes including carbohydrate metabolism, signal transduction, proteinbiosynthesis, photosynthesis, protein folding and assembly, energy pathway, cell rescue and defense, cell cycle, nitrogen metabolism, lipid metabolism,amino acid metabolism, transcription regulation, and secondary metabolite biosynthesis;42protein spots were found to be differentially expressed and successfully identified in cv. Ningzhen No.1. There were22spots upregulated an20spots downregulated.These proteins were involved in13cellular responses and metabolic processes including carbohydrate metabolism, signal transduction, proteinbiosynthesis, photosynthesis, protein folding and assembly, energy pathway, cell rescue and defense, cell cycle, nitrogen metabolism, lipid metabolism,amino acid metabolism, transcription regulation, and secondary metabolite biosynthesis.
6. According to discussed, the seed deterioration resistant mechanism were found in cv. Ningzhen No.1including:①. Increased synthesis of Arg;②. Glutamine synthesis pathway enhancement;③. CO2absorption enhancement;④. Enhanced photorespiration;⑤. The enhancement of secondary metabolite synthesis;⑥. Enhanced PPP pathy; Ⅶ. Enhanced ROS scavenging and DNA repair. The seed deterioration mechanism in cv. Ningzhen No.1including:①. cell cycle reduced;②. photosynthetic rate down;③. TCA cycle enhanced;④.protein synthesis reduction;⑤. oil synthetic pathy decrease;⑥. protein folding and assembly downregulated;⑦.Nitrogen assimilation.
The seed deterioration resistant mechanism in cv. Xiangdou NO.3including:①. photosynthetic rate upregulated;②. protein folding and assembly upregulated;③. Protein degradation decline;④. Energy consumption reduction;⑤. The enhancement of secondary metabolite synthesis;⑥. The increase of ASA synthesis. The seed deterioration mechanism in cv Xiangdou NO.3including:①. CO2absorption reduction;②. Cell wall synthesis reduction;③. protein synthesis;④. oil synthetic pathy decrease;⑤. Scavenging of reactive oxygen species antioxidase activity weakened.
1.对92份我国南方春大豆种质采用温箱蚀化模拟田间高温高湿条件进行胁迫,发现种质间种子田间劣变抗性存在显著差异；鉴定出种子田间劣变抗性种质湘豆三号和不抗种质宁镇一号,并确定了种子发芽率、活力指数和简易活力指数三个指标相结合可以作为种子田间劣变鉴定的标准。
2.高温高湿胁迫,对发育中的湘豆三号种子细胞的超微结构影响不明显；而对宁镇一号种子细胞的超微结构影响较大,表现为：种子细胞中叶绿体和蛋白体的数量不断减少和体积减小,细胞核和核仁的形状变得不规则,液泡和油体所占的面积更大,细胞壁变薄,淀粉粒数目减少,线粒体和光滑内质网数目增多。
3.高温高湿胁迫下,湘豆三号处于发育过程中的种子的相对电导率和MDA显著低于宁镇一号种子的；2个品种种子中ROS皆大量积累,打破了原有的氧化还原平衡体系,但湘豆三号种子中CAT活性和ASA含量显著高于宁镇一号种子的；2个品种种子中的化学成分都受到影响,其中：湘豆三号种子中蛋白和油脂的含量降低的幅度小于宁镇一号种子的；宁镇一号种子中内源乙烯含量比湘豆三号种子中内源乙烯的含量增加得更明显；2个品种叶片和种子中叶绿素含量、光化学效率均显著下降,其中宁镇一号下降的幅度更加明显。湘豆三号种子中NH4+、NO2-、NO3含量降低,而宁镇一号种子中的NH4+、NO3-含量升高,NO2-含量降低。
4.高温高湿胁迫下,采用GC-MS技术对湘豆三号和宁镇一号处于发育过程中的种子进行代谢谱分析,共检测到46种有差异表达的代谢产物,其中从抗性品种湘豆三号种子中检测出差异表达的代谢产物共34种,从不抗品种宁镇一号种子中检测出31种。这些代谢物质共涉及到光合作用、三羧酸循环、氨基酸合成途径、次生代谢产物合成途径、磷酸戊糖途径和光呼吸等6大代谢途径。高温高湿胁迫导致湘豆三号种子中卡尔文循环增强而宁镇一号种子中卡尔文循环减弱；导致宁镇一号种子中磷酸戊糖途径增强,而对湘豆三号种子中磷酸戊糖途径没有明显影响；诱导宁镇一号种子中光呼吸作用增强而对湘豆三号种子中光呼吸作用影响不显著；导致湘豆三号种子中TCA途径减弱,而宁镇一号种子中TCA途径显著增强。此外,高温高湿胁迫会导致湘豆三号和宁镇一号种子处于发育过程中的中氨基酸和次生代谢产物合成增强。
5.通过对高温高湿胁迫处理的宁镇一号和湘豆三号处于发育过程中的种子进行双向电泳及质谱分析,成功鉴定出87个差异表达蛋白点,这些差异蛋白点共涉及到15个代谢途径和细胞过程。在湘豆三号处于发育过程中的种子中共检测到45个差异表达蛋白点,涉及33个差异表达蛋白；其中21个蛋白点上调表达,24个蛋白点下调表达,涉及13个代谢途径和细胞过程,分别为碳素代谢、信号转导、蛋白合成、光合作用、蛋白折叠和组装、能量代谢、细胞修复和防御、脂代谢、氨基酸代谢、转录调控、次生代谢产物合成途径、蛋白降解及转运蛋白；在宁镇一号处于发育过程中的种子中共检测到42个差异表达蛋白点,涉及30个差异表达蛋白；其中22个蛋白点上调表达,20蛋白点下调表达,涉及13个代谢途径和细胞过程,分别为碳素代谢、信号转导、蛋白合成、光合作用、蛋白折叠和组装、能量代谢、细胞修复和防御、脂代谢、氨基酸代谢、氮素代谢、细胞周期、转录调控以及次生代谢产物合成途径。
6.综合形态、生理生化和蛋白质组学研究结果,发现：高温高湿胁迫下,湘豆三号处于发育过程中的种子中增强田间劣变抗性的代谢途径和细胞过程有：①光合作用的加强；②蛋白折叠和组装的加强；③蛋白降解的下降；④次生代谢物合成的增强；⑤能量消耗的减少；⑥ASA合成的增加。导致种子田间劣变性的代谢途径和细胞过程有：①CO2固定能力下降,碳素同化减少；②细胞壁合成降低；③蛋白含量降低；④油脂含量的降低；⑤抗氧化酶类的活性减弱。随着胁迫时间的延长,增强种子田间劣变抗性的因素逐渐增强而导致种子田间劣变性的因素不断减弱,最终前者逐渐占优势,结果发育过程中的湘豆三号种子抗种子劣变性增强。
高温高湿胁迫下,宁镇一号处于发育过程中的种子中增强田间劣变抗性的代谢途径和细胞过程有：①精氨酸含量的增加,②谷氨酰胺合成途径的增强,③CO2吸收的增强,④光呼吸的增强,⑤次生代谢物合成的增强；⑥磷酸戊糖途径的加强；⑦消除ROS能力和DNA修复的水平的提升。导致田间劣变性的代谢途径和细胞过程有：①细胞周期稳定性的降低；②光合作用降低；③蛋白绑定和折叠的降低；④能量的过度消耗；⑤蛋白含量降低；⑥油脂含量的降低；⑦氮素吸收的减弱。随着胁迫时间的延长,导致种子田间劣变性的因素不断增强而增强种子田间劣变抗性的因素不断减弱,最终前者逐渐占优势,结果宁镇一号的大豆种子在发育过程中的发生种子劣变。
High temperature and humidity (HTH) stress during soybean seed development and maturity in the field easily leads seed to pre-harvest deterioration. It often causes reduced vigor and germination rate, et al.This phenomenon has become a major obstacle of spring soybean production.At present, Related with seed deterioration research mainly focuses on the reasons that caused seed deterioration, cytology and genetics in the process of deterioration. The researches on cytological, physiological and molecular basis about seed deterioration and deterioration resistance are lack.
1.92of Spring soybean landraces and cultivars from South China were used to evaluate seed deterioration resistance by incubating weathering treatment. One germplasm (Xiangdou NO.3) identified to resist to pre-harvest seed deterioration and one (Ningzhen NO.1) identified not to resist pre-harvest seed deteriorations were screened out.
2. Transmission electron micrograph analysis indicated that HTH stress resulted in some ultrastructure changes in the cells of the developing seed of cv. Ningzhen No.1and the change was not significant in the cells of the developing seed of cv. Xiangdou No.3.In the cells of the developing seed of cv. Ningzhen No.1, the cells of the developing seed tended to exhibit thinner cell wall, smaller chloroplasts and protein bodies, more irregular cell nucleus and nuclear core, and bigger vacuoles and lipid bodies under HTH stress. Additionally, under HTH stress it was found that the decrease of starch grains in number and the increase of mitochondria and smooth endoplasmic reticula in number in the cell of the seed.
3. It was found that the HTH stress caused heavy accumulated ROS in the cells of the developing seed and breaked the original redox balance system;In the seed of cv. Ningzhen No.1showed higher electrolyte leakage and the content of MDA than cv. Xiangdou No.3.It showed the lower activity of CAT and the content of ASA than cv. Xiangdou No. 3,HTH stress was also caused change of chemical composition of the seed.The content of protein and oil were more decrease of cv. Ningzhen No.1than cv. Xiangdou No.3;the content of Endogenous ethylene of cv. Ningzhen No.1higher than cv. Xiangdou No.3; The chlorophyll content in the leaves and seeds and Fv/Fm were all significantly decreased.; Compared to the control, the content of NO3-and NH4+in HTH-stressed developing seed of cv. Ningzhen No.1were significantly increased while the NO2-content was significantly decreased, the content ofNO3-,NO2-,NH4+in HTH-stressed developing seed of cv. Xiangdou No.3were all decrease.
4. Metabolic profiling based on GC-MS showed46differentially expressed metabolites. These metabolites were classified into6categories.There were8kinds of sugar,4of sugar alcohol and1kinds of sugar acids,28.3%of the total that related to calvin cycle,With the Calvin cycle related substances significantly upregulated expression in cv. Xiangdou No.3and significantly downregulated expression.in cv. Ningzhen No.1;To detect the5kinds of PPP pathway of sugar phosphate material from cv. Ningzhen No.l, in HTH-stressed,they were upregulated expression;Detection of Phosphoglycolate, Ser and Gly3that they were related to photorespiratory patyway and were upregulated; There were6kinds material of TCA pathway,these material were upregulated in cv. Ningzhen No.1and were downregulated.in cv. Xiangdou No.3;14kinds of amino acids were detected, accounting for30.4%of the total number of detected objects in the two species were significantly or very significantly up-regulated expression; Detected7kinds of substances associated with secondary metabolite biosynthesis pathway, significantly up-regulated expression in the two species.
5. Compared the proteome composition of developing seed of cv. Xiangdou NO.3and cv. Ningzhen No.1at different HTH stress time points with their corresponding controls by2-DE.87protein spots were found to be differentially expressed and successfully identified by MALDI-TOF MS:45protein spots were found to be differentially expressed and successfully identified in cv. Xiangdou NO.3. there were21spots upregulated and24spots downregulated,These proteins were involved in13of cellular responses and metabolic processes including carbohydrate metabolism, signal transduction, proteinbiosynthesis, photosynthesis, protein folding and assembly, energy pathway, cell rescue and defense, cell cycle, nitrogen metabolism, lipid metabolism,amino acid metabolism, transcription regulation, and secondary metabolite biosynthesis;42protein spots were found to be differentially expressed and successfully identified in cv. Ningzhen No.1. There were22spots upregulated an20spots downregulated.These proteins were involved in13cellular responses and metabolic processes including carbohydrate metabolism, signal transduction, proteinbiosynthesis, photosynthesis, protein folding and assembly, energy pathway, cell rescue and defense, cell cycle, nitrogen metabolism, lipid metabolism,amino acid metabolism, transcription regulation, and secondary metabolite biosynthesis.
6. According to discussed, the seed deterioration resistant mechanism were found in cv. Ningzhen No.1including:①. Increased synthesis of Arg;②. Glutamine synthesis pathway enhancement;③. CO2absorption enhancement;④. Enhanced photorespiration;⑤. The enhancement of secondary metabolite synthesis;⑥. Enhanced PPP pathy; Ⅶ. Enhanced ROS scavenging and DNA repair. The seed deterioration mechanism in cv. Ningzhen No.1including:①. cell cycle reduced;②. photosynthetic rate down;③. TCA cycle enhanced;④.protein synthesis reduction;⑤. oil synthetic pathy decrease;⑥. protein folding and assembly downregulated;⑦.Nitrogen assimilation.
The seed deterioration resistant mechanism in cv. Xiangdou NO.3including:①. photosynthetic rate upregulated;②. protein folding and assembly upregulated;③. Protein degradation decline;④. Energy consumption reduction;⑤. The enhancement of secondary metabolite synthesis;⑥. The increase of ASA synthesis. The seed deterioration mechanism in cv Xiangdou NO.3including:①. CO2absorption reduction;②. Cell wall synthesis reduction;③. protein synthesis;④. oil synthetic pathy decrease;⑤. Scavenging of reactive oxygen species antioxidase activity weakened.
引文
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