种子保存过程中生活力丧失特性及其机理研究
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摘要
种子成熟后即开始进入衰老过程,即使在种质库低温低湿的保存条件下,种子生活力仍然在缓慢降低。了解种子衰老特性是确保种子长期安全保存的重要基础。本文分析了种子在不同保存条件下生活力丧失特性、不同发芽率更新后种质遗传完整性变化和不同发芽率种子的蛋白质组变化,从发芽能力、遗传和蛋白变化角度深入理解农作物种子的衰老特性,以期为制定种子衰老预警和繁殖更新标准提供理论依据。本研究主要取得以下结果：
(1)低温保存种子生活力监测：通过分析国家库(-18℃)34种作物14706份种子保存20年的生活力监测数据发现,92.9%被监测种子发芽率保持在85%以上,即国家种质库绝大多数种质可安全保存20年以上。但有1.1%种子的发芽率从80%以上降至70%以下,包括小豆、牧草、黄麻等,表明-18℃C保存条件下种子生活力仍在缓慢降低。
(2)种子在不同保存条件下的生活力丧失特性：分析了在中期库(10-4℃)、南昌室温、三亚室温和40℃高温保存的小麦和大豆种子的生活力变化,结果发现各保存条件下的种子寿命均服从正态分布,种子存活曲线均为反S形,不同条件下小麦种子生活力开始快速下降的转折点(Pt)在82.2%至83.6%之间,暗示这些特点均不受保存环境影响,Pt可以作为种子生活力快速下降的预警指标。
(3)种子生活力下降的蛋白质组分析：分析了经50℃加速老化玉米种子干胚(未吸胀)中蛋白质组的变化,分离到40个差异表达蛋白,说明老化可引起干种子蛋白质组变化。蛋白质谱鉴定结果表明老化扰乱了种子的转录和信号转导,导致保护能力的下降,诱导了蛋白酶的上调表达,促进贮藏蛋白的降解,削弱了代谢和能量供应能力,最终导致种子生活力下降。其中与能量供应相关的蛋白所占比例最高,涵盖了从糖酵解、TCA循环、电子传递链到氧化磷酸化各个环节,结合对磷酸戊糖途径(G6PDHase酶活性)、Glc-6-P、丙酮酸和ATP含量的测定,认为贮藏糖类物质的动员和能量供应在玉米种子老化和生活力中具有重要作用。在老化种子中新出现或者消失的蛋白点,可作为衰老预警指标。
(4)种子生活力下降对遗传完整性的影响：分析了10份小麦异质种质在中期库(10-4。C)保存了16-17年繁殖更新后的农艺性状和醇溶蛋白,发现其中4份材料的部分农艺性状(株高、旗叶长和宽)、醇溶蛋白带型频率和一些遗传参数,如I、He发生了明显改变。这4份材料更新时发芽率小于等于66%,而其余更新发芽率大于等于75%的材料在更新后则没有显著变化。因此建议将75%作为种子繁殖更新标准的发芽率下限。
Seeds enters aging phase after maturation. Seed viability keeps on decrease along with the extended storage time, even when being stored at genebanks with low temperature and relative humidty. It is essential to get a better understand of the character of seed aging in order to asure safe long-term storage. In this work, the loss of seed viability under different storage conditions, the changes of genetic integrity brought by regeneration carried ou at various viability levels, and the proteomic changes brought by seed aging were studied by seed viability monitoring, agronomic traits, gliadin SDS-PAGE,2-DE and MALDI-TOF/MS in order to further our understanding of seed aging mechanism, and to provide theoretical basis for the establishment of a germination standard for seed aging and germplasm regeneration. The main results are as follows:
(1) Viability monitoring of seeds being stored at low temperature genebank:The viability of14706seed accessions, belonging to34crops, was detected after being stored at the National Genebank (-18℃) for20to23years. Results showed that the viability of92.9%accessions was above85%, which indicated that most of the collections in the National Genenbank can be safely conserved for more than20years. But the viability of1.1%of the monitored accessions was lower than70%, which suggested that seed still loss their viability even at-18℃storage.
(2) Seed viability loss under different storage conditions:The viability monitoring data of wheat and soybean seeds from a midterm genebank (10～4℃), ambient storage, as well as an accelerated aging experiment were quantified, and results showed that seed longevity of all accessions followed normal distribution, and that seed survival curves showed inverse-S shape, which received no obvious effects from storage condition. The point between the plateau phase and viability sharp decline phase was recorded as the turning point (Pt). Results showed that the Pt under different storage conditions were quite close, which varied between82.2%and83.6%, indicating the possible role of Pt as indicators for seed viability loss during germplasm storage.
(3) Changes of the proteome of aged seeds:The proteomic changes in the dry embryos of maize (Zea mays L. cv. Dabaitou) seeds being artificially aged at50℃were studied using2-DE and MALDI-TOF/MS. A total of40differentially expressed proteins were identified, in which16proteins were up-regulated, indicating that artificial aging affected the proteome of the dry seeds. Identification of these proteins revealed that the signal transduction and transcription was disturbed by artificial aging, which might lead to reduced protection against aging. Artificial aging also increased proteases and broke down stored proteins, impaired metabolism and energy supply, and ultimately resulted in seed deterioration. Proteins involved in metabolism and energy were the largest down-regulated protein group, which concerning glycolysis, TCA cycle, the electron transport chain, and oxidative phosphorylation. The down-regulation of these proteins, together with reduction in the specific activity of G6PDHase, and the content of Glc-6-P, pyruvic acid and ATP in aged seeds, suggested the important roles of the mobilization of stored carbohydrates and energy supply in seed aging and seed vigour. Proteins that dissapered or showed only in aged seeds, could be used as warning indicators for seed aging.
(4) Changes of genetic integrity brought by seed aging:Ten wheat accessions that had been stored at a medium-term genebank (10～4℃) for16to17years were regenerated and16agronomic traits and the gliadin spectrum were compared with seeds stored at a long-term genebank. The results provided evidence that some agronomic traits, such as length and width or the flag leaf, plant height, and the frequency of the gliadin band-type, and/or some of the genetic parameters, such as I, He, of4accessions changed significantly after only one generation. The original germination percentages of these four accessions were equal to and lower than66%. As for those accessions, which were regenerated at viability equal to and higher than75%germination, no significant changes were detected. Therefore,75%germination percentage is recommended as the safe lower limit of regeneration standard for wheat seeds.
(1)低温保存种子生活力监测：通过分析国家库(-18℃)34种作物14706份种子保存20年的生活力监测数据发现,92.9%被监测种子发芽率保持在85%以上,即国家种质库绝大多数种质可安全保存20年以上。但有1.1%种子的发芽率从80%以上降至70%以下,包括小豆、牧草、黄麻等,表明-18℃C保存条件下种子生活力仍在缓慢降低。
(2)种子在不同保存条件下的生活力丧失特性：分析了在中期库(10-4℃)、南昌室温、三亚室温和40℃高温保存的小麦和大豆种子的生活力变化,结果发现各保存条件下的种子寿命均服从正态分布,种子存活曲线均为反S形,不同条件下小麦种子生活力开始快速下降的转折点(Pt)在82.2%至83.6%之间,暗示这些特点均不受保存环境影响,Pt可以作为种子生活力快速下降的预警指标。
(3)种子生活力下降的蛋白质组分析：分析了经50℃加速老化玉米种子干胚(未吸胀)中蛋白质组的变化,分离到40个差异表达蛋白,说明老化可引起干种子蛋白质组变化。蛋白质谱鉴定结果表明老化扰乱了种子的转录和信号转导,导致保护能力的下降,诱导了蛋白酶的上调表达,促进贮藏蛋白的降解,削弱了代谢和能量供应能力,最终导致种子生活力下降。其中与能量供应相关的蛋白所占比例最高,涵盖了从糖酵解、TCA循环、电子传递链到氧化磷酸化各个环节,结合对磷酸戊糖途径(G6PDHase酶活性)、Glc-6-P、丙酮酸和ATP含量的测定,认为贮藏糖类物质的动员和能量供应在玉米种子老化和生活力中具有重要作用。在老化种子中新出现或者消失的蛋白点,可作为衰老预警指标。
(4)种子生活力下降对遗传完整性的影响：分析了10份小麦异质种质在中期库(10-4。C)保存了16-17年繁殖更新后的农艺性状和醇溶蛋白,发现其中4份材料的部分农艺性状(株高、旗叶长和宽)、醇溶蛋白带型频率和一些遗传参数,如I、He发生了明显改变。这4份材料更新时发芽率小于等于66%,而其余更新发芽率大于等于75%的材料在更新后则没有显著变化。因此建议将75%作为种子繁殖更新标准的发芽率下限。
Seeds enters aging phase after maturation. Seed viability keeps on decrease along with the extended storage time, even when being stored at genebanks with low temperature and relative humidty. It is essential to get a better understand of the character of seed aging in order to asure safe long-term storage. In this work, the loss of seed viability under different storage conditions, the changes of genetic integrity brought by regeneration carried ou at various viability levels, and the proteomic changes brought by seed aging were studied by seed viability monitoring, agronomic traits, gliadin SDS-PAGE,2-DE and MALDI-TOF/MS in order to further our understanding of seed aging mechanism, and to provide theoretical basis for the establishment of a germination standard for seed aging and germplasm regeneration. The main results are as follows:
(1) Viability monitoring of seeds being stored at low temperature genebank:The viability of14706seed accessions, belonging to34crops, was detected after being stored at the National Genebank (-18℃) for20to23years. Results showed that the viability of92.9%accessions was above85%, which indicated that most of the collections in the National Genenbank can be safely conserved for more than20years. But the viability of1.1%of the monitored accessions was lower than70%, which suggested that seed still loss their viability even at-18℃storage.
(2) Seed viability loss under different storage conditions:The viability monitoring data of wheat and soybean seeds from a midterm genebank (10～4℃), ambient storage, as well as an accelerated aging experiment were quantified, and results showed that seed longevity of all accessions followed normal distribution, and that seed survival curves showed inverse-S shape, which received no obvious effects from storage condition. The point between the plateau phase and viability sharp decline phase was recorded as the turning point (Pt). Results showed that the Pt under different storage conditions were quite close, which varied between82.2%and83.6%, indicating the possible role of Pt as indicators for seed viability loss during germplasm storage.
(3) Changes of the proteome of aged seeds:The proteomic changes in the dry embryos of maize (Zea mays L. cv. Dabaitou) seeds being artificially aged at50℃were studied using2-DE and MALDI-TOF/MS. A total of40differentially expressed proteins were identified, in which16proteins were up-regulated, indicating that artificial aging affected the proteome of the dry seeds. Identification of these proteins revealed that the signal transduction and transcription was disturbed by artificial aging, which might lead to reduced protection against aging. Artificial aging also increased proteases and broke down stored proteins, impaired metabolism and energy supply, and ultimately resulted in seed deterioration. Proteins involved in metabolism and energy were the largest down-regulated protein group, which concerning glycolysis, TCA cycle, the electron transport chain, and oxidative phosphorylation. The down-regulation of these proteins, together with reduction in the specific activity of G6PDHase, and the content of Glc-6-P, pyruvic acid and ATP in aged seeds, suggested the important roles of the mobilization of stored carbohydrates and energy supply in seed aging and seed vigour. Proteins that dissapered or showed only in aged seeds, could be used as warning indicators for seed aging.
(4) Changes of genetic integrity brought by seed aging:Ten wheat accessions that had been stored at a medium-term genebank (10～4℃) for16to17years were regenerated and16agronomic traits and the gliadin spectrum were compared with seeds stored at a long-term genebank. The results provided evidence that some agronomic traits, such as length and width or the flag leaf, plant height, and the frequency of the gliadin band-type, and/or some of the genetic parameters, such as I, He, of4accessions changed significantly after only one generation. The original germination percentages of these four accessions were equal to and lower than66%. As for those accessions, which were regenerated at viability equal to and higher than75%germination, no significant changes were detected. Therefore,75%germination percentage is recommended as the safe lower limit of regeneration standard for wheat seeds.
引文
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