6-BA处理对玉米胚乳发育及细胞程序性死亡调控效应的研究
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摘要
本试验以郑单958(Z)、农大108(N)、特爆2号(T)为材料,研究授粉后喷施6-苄基腺嘌呤(6-BA)对玉米胚乳发育及胚乳细胞程序性死亡(Programmed cell death,PCD)的影响。采用生理生化方法,从胚乳细胞数目、内源激素、籽粒营养物质等方面探讨了6-BA处理对玉米胚乳发育的影响;从显微构造、超微构造和分子生物学等方面观察了玉米胚乳PCD过程的特征及6-BA处理对胚乳细胞PCD过程的调控作用,主要研究结果如下:
1 6-BA处理对胚乳细胞数目的影响
对玉米籽粒胚乳细胞数目的观察发现,6-BA处理提高了3个品种的胚乳细胞数目,且胚乳细胞数目随6-BA浓度的提高而增加。但6-BA处理对3个品种胚乳细胞数目的影响程度不同,与对照相比,郑单958、特爆2号的胚乳细胞增殖达到显著水平,而农大108未达到显著水平。施用6-BA引起胚乳细胞数目的增多,可能是6-BA处理后玉米粒重提高的原因之一。
2 6-BA处理对胚乳内源激素含量的影响
6-BA处理后,郑单958、特爆2号的Z+ZR、GA含量较对照显著提高,随6-BA浓度的提高而增加。而农大108 Z+ZR、GA含量的提高,未达到显著水平。6-BA处理显著提高了3个品种灌浆前期的iPA含量,同时抑制了玉米灌浆后期胚乳内的ABA的积累,显著提高了郑单958灌浆前期和后期、特爆2号灌浆前中期、农大108灌浆后期的IAA含量。施用6-BA引起籽粒内源激素含量的改变,可能是导致籽粒产量和品质性状改变的原因之一。
3 6-BA处理对籽粒营养物质的影响
6-BA处理明显提高了3个品种灌浆前中期的蔗糖含量、整个灌浆期的蛋白质含量、籽粒灌浆后期的支链淀粉、总淀粉含量,为形成较高的粒重提供了基础。
4玉米胚乳细胞PCD的特点
超微结构观察发现,玉米胚乳细胞在发育过程中出现细胞核变形、解体,形成核残体和染色质凝集的现象;在核解体过程中,线粒体出现伴随变化,如线粒体嵴消失呈中央空泡化;琼脂糖凝胶电泳发现,胚乳DNA显示以180-200bp倍增的梯状带。这些现象表明玉米胚乳细胞的死亡是PCD过程。用Evans Blue对胚乳的染色图表明,玉米胚乳细胞死亡始于胚乳上端并高度有序地向下、向外扩展。超微结构与Evans Blue染色结果说明,细胞核消亡后,细胞仍存活,胞质中仍可正常合成和积累淀粉、蛋白质,在胚乳细胞完成了合成与积累营养物质的使命后,即细胞被淀粉充满后,胚乳细胞死亡并形成无生命的营养物质储藏库。
5 6-BA处理对胚乳细胞PCD的影响
6-BA处理的胚乳出现梯状DNA带的时间与对照一致,但DNA电泳带的亮度比对照明显弱,且随6-BA浓度的增大,减弱程度增强。经Evans Blue染色证明喷施6-BA使得蓝色反应区域面积减小,颜色变浅。超微结构观察亦发现,经6-BA处理的胚乳细胞核降解程度明显轻于对照,以上研究结果均说明,6-BA处理可在一定程度上减缓PCD的过程,从而改善籽粒品质,提高玉米粒重。
Three maize cultivars(Z,N,T),were used in the present study .The effects of 6-BA on the development of maize endosperm after pollination were investigated by the physiological and biochemical method with the characteristic of hormone, the number of endosperm, the kernel nutriment content. The characteristic of different maize of endosperm programmed cell death and the effects of 6-BA on the endosperm programmed cell death with the optical microscope, transmission electro microscope, molecular biology, respectively. The main results were follows:
1 The effects of exogenous 6-BA on the number of endosperm cell
The endosperm cell numbers were determined. The numbers of different maize endosperm were increased by the different 6-BA concentration. But the effects degree of 6-BA were different on three maize cultivars. Compared to the contrast, the endosperm cell numbers of Z and T. were prominently increased but N. It was probably one of the important causes to improve grain yield by spraying this regulator.
2 The effects of exogenous 6-BA on endogenous hormones contents
Endogenous hormones contents in endosperm were measured for each treatment. The results indicated that 6-BA had different effects on endogenous hormones at different grain filling stages. Compared to the control , discrepancy between the treatments of Z and T were prominent but N, the content of Z+ZR, GA were increased by the increased concentration; the iPA content in the early grain filling stage of three maize cultivars was prominently enhanced; notably restrained the accumulation of ABA; the IAA content was notably increased in the early and late stage grain filling of Z, in the late grain filling stage of N, in the early grain filling stage and decreased in the late grain filling stage of T. Under this experiment the changes of endogenous hormones were probably one of the important causes led to variations of yield and quality.
3 The effects of exogenous 6-BA on the content of nutritious substance
The content of nutritious substance were significantly increased in different grain filling stage, including the content of sucrose in the early and middle grain filling stage, the content of protein in the entire grain filling stage, the content of amylopection, total starch of a grain in the late filling stage.
4 The characteristic of maize endosperm PCD
Conventional ultrastructural observation showed that the starchy endosperm cells of maize underwent nucleus deformation, nucleus disintegration, nucleus residue formation and chromatin condensation. In the process of nucleus disintegration, mitochondrion also showed a series of structural changes, such as the cristae of mitochondrion disappeared, resulting in central hollowed mitochondrion. The above results proved that the cell death of starchy endosperm in maize was a PCD process. Evans Blue staining proved that death of endosperm cell started the top of endosperm and expanded toward adown and forth of endosperm. The results of ultrastructual observation and Evans Blue staining proved that nucleus disintegration, the starchy endosperm cell still remained alive, and the reserves including starch and storage proteins were still synthesized and accumulated in the cell as normal. It was the date of cell death until the cell was filled with starch endosperm cells in maize.
5 The effects of exogenous 6-BA on the PCD of Endosperm
Treatment with 6-BA reduced the extent of DNA fragmentation in developing starchy endosperm cells and the reduced degree increased with the concentration of 6-BA. Evans Blue staining proved that the blue area lessen compared to control. The results of ultrastructual observation proved the degree of nucleus disintegration lessen compared to control. It proved that the 6-BA may lessen the endosperm cell PCD, probably one of the important causes leading to variations of yield and quality.
1 6-BA处理对胚乳细胞数目的影响
对玉米籽粒胚乳细胞数目的观察发现,6-BA处理提高了3个品种的胚乳细胞数目,且胚乳细胞数目随6-BA浓度的提高而增加。但6-BA处理对3个品种胚乳细胞数目的影响程度不同,与对照相比,郑单958、特爆2号的胚乳细胞增殖达到显著水平,而农大108未达到显著水平。施用6-BA引起胚乳细胞数目的增多,可能是6-BA处理后玉米粒重提高的原因之一。
2 6-BA处理对胚乳内源激素含量的影响
6-BA处理后,郑单958、特爆2号的Z+ZR、GA含量较对照显著提高,随6-BA浓度的提高而增加。而农大108 Z+ZR、GA含量的提高,未达到显著水平。6-BA处理显著提高了3个品种灌浆前期的iPA含量,同时抑制了玉米灌浆后期胚乳内的ABA的积累,显著提高了郑单958灌浆前期和后期、特爆2号灌浆前中期、农大108灌浆后期的IAA含量。施用6-BA引起籽粒内源激素含量的改变,可能是导致籽粒产量和品质性状改变的原因之一。
3 6-BA处理对籽粒营养物质的影响
6-BA处理明显提高了3个品种灌浆前中期的蔗糖含量、整个灌浆期的蛋白质含量、籽粒灌浆后期的支链淀粉、总淀粉含量,为形成较高的粒重提供了基础。
4玉米胚乳细胞PCD的特点
超微结构观察发现,玉米胚乳细胞在发育过程中出现细胞核变形、解体,形成核残体和染色质凝集的现象;在核解体过程中,线粒体出现伴随变化,如线粒体嵴消失呈中央空泡化;琼脂糖凝胶电泳发现,胚乳DNA显示以180-200bp倍增的梯状带。这些现象表明玉米胚乳细胞的死亡是PCD过程。用Evans Blue对胚乳的染色图表明,玉米胚乳细胞死亡始于胚乳上端并高度有序地向下、向外扩展。超微结构与Evans Blue染色结果说明,细胞核消亡后,细胞仍存活,胞质中仍可正常合成和积累淀粉、蛋白质,在胚乳细胞完成了合成与积累营养物质的使命后,即细胞被淀粉充满后,胚乳细胞死亡并形成无生命的营养物质储藏库。
5 6-BA处理对胚乳细胞PCD的影响
6-BA处理的胚乳出现梯状DNA带的时间与对照一致,但DNA电泳带的亮度比对照明显弱,且随6-BA浓度的增大,减弱程度增强。经Evans Blue染色证明喷施6-BA使得蓝色反应区域面积减小,颜色变浅。超微结构观察亦发现,经6-BA处理的胚乳细胞核降解程度明显轻于对照,以上研究结果均说明,6-BA处理可在一定程度上减缓PCD的过程,从而改善籽粒品质,提高玉米粒重。
Three maize cultivars(Z,N,T),were used in the present study .The effects of 6-BA on the development of maize endosperm after pollination were investigated by the physiological and biochemical method with the characteristic of hormone, the number of endosperm, the kernel nutriment content. The characteristic of different maize of endosperm programmed cell death and the effects of 6-BA on the endosperm programmed cell death with the optical microscope, transmission electro microscope, molecular biology, respectively. The main results were follows:
1 The effects of exogenous 6-BA on the number of endosperm cell
The endosperm cell numbers were determined. The numbers of different maize endosperm were increased by the different 6-BA concentration. But the effects degree of 6-BA were different on three maize cultivars. Compared to the contrast, the endosperm cell numbers of Z and T. were prominently increased but N. It was probably one of the important causes to improve grain yield by spraying this regulator.
2 The effects of exogenous 6-BA on endogenous hormones contents
Endogenous hormones contents in endosperm were measured for each treatment. The results indicated that 6-BA had different effects on endogenous hormones at different grain filling stages. Compared to the control , discrepancy between the treatments of Z and T were prominent but N, the content of Z+ZR, GA were increased by the increased concentration; the iPA content in the early grain filling stage of three maize cultivars was prominently enhanced; notably restrained the accumulation of ABA; the IAA content was notably increased in the early and late stage grain filling of Z, in the late grain filling stage of N, in the early grain filling stage and decreased in the late grain filling stage of T. Under this experiment the changes of endogenous hormones were probably one of the important causes led to variations of yield and quality.
3 The effects of exogenous 6-BA on the content of nutritious substance
The content of nutritious substance were significantly increased in different grain filling stage, including the content of sucrose in the early and middle grain filling stage, the content of protein in the entire grain filling stage, the content of amylopection, total starch of a grain in the late filling stage.
4 The characteristic of maize endosperm PCD
Conventional ultrastructural observation showed that the starchy endosperm cells of maize underwent nucleus deformation, nucleus disintegration, nucleus residue formation and chromatin condensation. In the process of nucleus disintegration, mitochondrion also showed a series of structural changes, such as the cristae of mitochondrion disappeared, resulting in central hollowed mitochondrion. The above results proved that the cell death of starchy endosperm in maize was a PCD process. Evans Blue staining proved that death of endosperm cell started the top of endosperm and expanded toward adown and forth of endosperm. The results of ultrastructual observation and Evans Blue staining proved that nucleus disintegration, the starchy endosperm cell still remained alive, and the reserves including starch and storage proteins were still synthesized and accumulated in the cell as normal. It was the date of cell death until the cell was filled with starch endosperm cells in maize.
5 The effects of exogenous 6-BA on the PCD of Endosperm
Treatment with 6-BA reduced the extent of DNA fragmentation in developing starchy endosperm cells and the reduced degree increased with the concentration of 6-BA. Evans Blue staining proved that the blue area lessen compared to control. The results of ultrastructual observation proved the degree of nucleus disintegration lessen compared to control. It proved that the 6-BA may lessen the endosperm cell PCD, probably one of the important causes leading to variations of yield and quality.
引文
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