甜菜抗丛根病细胞生物学特征研究
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摘要
甜菜是我国乃至世界上重要的糖料作物和经济作物之一,由于长期以来受到甜菜各种病害的袭击,不仅给甜菜生产造成巨大的经济损失,并且制约制糖业的发展。甜菜丛根病(Rhizomania)是危害最为严重的病害之一,该病以多粘菌(Polymyxa betae)为传播介体,由甜菜坏死黄脉病毒(Beet necrotic yellow vein virus, BN YVV)引起的土传病害。近年来,国内外主要从病原基因组学、分子生物学及寄主生理生化角度来探讨病毒致病机理和甜菜抗丛根病机理。本研究以甜菜抗、感丛根病品种与BNYVV互作体系为研究对象,对不同互作体系中的形态学特征和细胞学特征进行了比较研究,利用生理生化方法明确不同互作体系间活性氧(H2O2和02-·)及保护酶系(POD和SOD)产生的时间变化特征,采用电镜细胞化学标记技术在亚细胞水平上对活性氧(H2O2和O2-·)及保护酶系(POD和SOD)的空间分布进行了定位,并检测了不同互作体系间木质素组织定位的差异及细胞壁糖蛋白在寄主细胞内积累的规律,以明确活性氧代谢及细胞壁成分在甜菜抗丛根病性中的作用,从而为进一步揭示甜菜抗丛根病信号传导机制奠定理论基础。通过研究取得的主要结果如下：
1.利用光学显微技术和透射电镜技术研究了甜菜抗、感丛根病品种与BNYVV互作过程中的形态学和超微结构特征。结果表明,不同寄主与病毒互作体系的甜菜块根形态学和超微结构上均表现出明显的差异,而甜菜叶脉受该病毒侵染后其形态学解剖结构无差异,但超微结构受到影响。形态学水平上,抗、感病品种表现为表皮破坏,皮层薄壁细胞组织产生裂隙,大多细胞都脱落,木质部附近的木薄壁细胞破损,完全丧失了对根部的保护作用。其中感病品种比抗病品种表现被伤害程度更为严重,其皮层薄壁细胞几乎全部脱落,生长速度缓慢,根部发育受阻。而叶脉的形态解剖结构与对照比较,未发现异常变化。另外抗病品种比感病品种有更发达的维管束结构。在亚细胞水平上,BNYVV对感病品种的细胞超微结构破坏严重,整个细胞变形,空虚化,细胞核结构发生紊乱,线粒体和高尔基体明显增多,细胞质中小液泡增多,在液泡膜边缘可见到一些纤细丝状物质的圆形或卵圆形小泡突入液泡中。叶脉细胞叶绿体完全瓦解,出现许多嗜锇颗粒。而抗病品种细胞超微结构破坏较轻,寄主细胞产生一系列显著的结构防卫反应：形成细胞壁沉积物及液泡膜上显示出黑色颗粒状沉积物等。说明组织形态结构变化和细胞器病理变化与甜菜抗丛根病性有关。
2.利用生理生化方法和电镜细胞化学标记技术研究了甜菜-BNYVV互作过程中H2O2和O2-·产生的时间变化特征和空间分布定位。研究结果表明,甜菜抗、感丛根病体系均在侵染早期出现大量H2O2和O2-·,其中抗病体系的H2O2和O2-·产生量明显高于感病体系。H2O2在抗病和感病体系中的分布位置基本相似,多分布在块根、叶脉细胞的液泡膜和质膜上,叶脉细胞间隙也有H2O2的分布,而O2-·在抗病品种块根与叶脉细胞的质膜上定位,感病品种的则在液泡膜上被发现。而且感病体系H202和O2-·沉积量明显弱于抗病体系。H2O2和O2-·产生量和分布与甜菜抗丛根病性有密切联系,不同部位定位的H2O2和O2-·作为信号分子,参与了甜菜对病毒侵染的防御反应。
3.采用生化检测及酶细胞化学方法研究了甜菜与BNYVV互作过程中POD和SOD的活性及其在细胞内分布特征。结果表明,病毒感染前POD主要定位在甜菜块根细胞细胞壁及叶脉细胞线粒体和细胞间隙,当病毒感染后,抗、感病甜菜品种块根皮层细胞的细胞壁、质膜和液泡膜上的POD及叶脉薄壁细胞的细胞壁、质膜、液泡膜、线粒体、细胞间隙上的POD活性均比其对照显著升高,抗、感病甜菜品种在POD分布部位上没有区别,但两者沉积量有所差异,抗病品种明显高于感病品种；SOD在未感染病毒的抗、感病甜菜块根皮层薄壁细胞质膜与液泡膜,叶脉厚角组织薄壁细胞质膜及叶脉细胞线粒体、细胞间隙上被发现,病毒侵染后甜菜抗、感病品种块根SOD在细胞内分布位置相同都分布在质膜和液泡膜,但抗病品种以质膜上分布为主,感病品种以液泡膜上分布为主。抗病品种SOD活性不仅明显高于其对照,也明显高于病毒侵染后的感病品种。甜菜抗、感病品种POD及SOD细胞内分布的沉积量与生理生化检测的酶活性结果一致,从亚细胞学水平说明高活性的POD及SOD是甜菜抗丛根病性生化标记的重要生理机制之一。
4.通过组织化学染色法及电镜细胞化学方法研究BNYVV对甜菜细胞壁中木质素和HRGP的影响。结果显示,BNYVV侵染前后在甜菜抗、感病品种的块根与叶脉横切面的所有导管上均有木质素和HRGP的沉积,块根薄壁组织的细胞壁由外向内也逐渐显示出由深至浅的木质素沉积；抗病品种木质素和HRGP的积累量较感病品种和其对照增加更为显著；说明在甜菜与BNYVV互作体系中,HRGP参与木质素的合成,木质素和HRGP在甜菜组织内快速积累是甜菜抗丛根病性的表现之一。
Sugar beet (Beta vulgaris L.) is important sugar and cash crops in our country even in the world. Under the various attacked of sugar beet diseases not only have causing huge economic losses in the sugar beet production industry but also restricted the development of the sugar industry. Rhizomania caused by Beet necrotic yellow vein virus (BNYVV) and mediated by Polymyxa betae which was one of the most serious soil-born diseases. Recently, numerous researchs had studied the mechanism of rhizomania at domestic and abroad were main under the views of genomics, molecular biology, physiological and biochemical. No research studying the mechanism of rhizomania and its resistance signal transduction pathway from the cytological at present. Therefore, in this study, sugar beet varieties and BNYVV interaction system were the research objects, the morphological and cytological features of the two objects were studied. Also, to make sure the time-varying characteristics and located the spatial distribution in the sub-cellular level of the production of the active oxygen species (H2O2and O2-·) and the protection of enzymes (POD and SOD) in the interaction system, the physiological and biochemical methods and electron microscopic cytochemistry labeling technique were used, And we have detected the locating differences within organization of the lignin and the law of cell wall glycoprotein accumulated in the host cell in the interactions system to observed the roles of the active oxygen metabolism and cell wall components in the sugar beet resistance to rhizomania disease resistance, then lay a theoretical foundation for the further study of the signal conduction mechanism of sugar beet resistance to rhizomania disease. The main results obtained through the study were as follows:
1. We have utilized optical microscopy and transmission electron microscopy to reveal the structural features of the morphology and ultrastructure between the interaction of the resistant and susceptible beet varieties and the BNYVV. The results showed that the morphology and ultrastructure of the sugar beet roots of the resistant and susceptible interaction system, infected by BNYVV, showed significant differences, the ultrastructure of the sugar beet vein have been seriously affected too, but the morphological anatomical structure of the sugar beet vein was no difference. The performance of the Morphology happened in the resistant and susceptible sugar beet included the epidermal destruction, the fracture of the cortical parenchyma tissue, the shed in most cell and the damage of the wood parenchyma cells near the xylem, the protection for the roots loss completely. The performance of the susceptible beet is more serious than the resistant beet. In the susceptible beet, the cortical parenchyma cells were almost all off, growing slowly and the root's development was blocked. However, there are no abnormal changes in the morphological anatomical structure of the beet vein. In addition, the resistant varieties have a well developed vascular bundle structure than susceptible varieties. When observed from the sub-cellular level, we found that BNYVV resulted in a series of damaged on cell ultrastructure of the susceptible beet, such as the deformation, empty and the structure of the nuclear is disorder, the mitochondria and the Golgi apparatus was significantly increased, small and medium-sized vacuoles were increased in the cytoplasmic. We observed some round or oval vesicles containing thin filaments broke into the vacuole at the edge of the vacuolar membrane. In the Vein cells the chloroplast disintegrated completely and a abundance of Osmiophilic globule emerged. Destruction of the cell ultrastructure of resistant varieties was lighter, a series of significant structural defense reaction produced in:the host cells:there was sediments in the cell wall and black granular sediment showed in the vacuolar membrane. These indicated that the description of the morphological changes of the tissue and the pathological changes of the organelle were relative to the resistance to rhizomania.
2. We have utilized Physiological and biochemical methods and electron microscopy cytochemistry labeling technique to study the time-varying characteristics and the orientation of spatial distribution of the H2O2and O2-·generated between the process of the beet-BNYVV interaction. A large number of H2O2and O2·detected both in the resistant and susceptible beet, but the production in the resistant beet was higher than the susceptible beet significantly. H2O2whose position of distribution was similar in the resistant and susceptible beet, which was located in the roots, the tonoplast and plasma membrane of the veins cell, and exist in the gap of the veins cell also. The O2-·is detected in the vein and the plasma membran of the veins cell in the resistant, but found that in the tonoplast in the susceptible beet. However, the content of H2O2and O2-·was distinctly different in the different beet systems, the content in the susceptible system was significantly lower than the resistant system. The results suggested that the production and distribution of the H2O2and O2-·were significantly different in the different interactive system, these difference were close with the beet's resistance to the rhizomania. As a signaling molecule, the H2O2and O2-·located in different positions involved in the defense response of the beet to virus infection.
3. Research was carried on POD and SOD activity and their intracellular distribution characteristics in sugar beet and BNYVV interaction processes by using biochemical detection and enzyme cytochemical methods. The results showed that before the viral infection, the POD was mainly localized in the root cell wall and mitochondria of the vein cells and the gaps between the cells. after the virus infection, in the resistant and susceptible sugar beet varieties, the POD activity in the cell wall, plasma membrane and tonoplast of the tuberous root cortex cells, and in the cell wall, plasma membrane, tonoplast, mitochondria and the gap of the veins cells is significantly increased than that of the control group. Although the distribution of POD was almost same in both beet system, but the deposition in the former was significantly higher in the latter. When the resitant and susceptcible were not infected virus, SOD was defected in plasma membrane and tonoplast of the beet root cortical parenchyma cell, cytoplasmic membrane of vein collenchyma thin-walled, mitochondria of veins and the gaps of the cells. The distribution of SOD in susceptible varieties and resistant sugar beet root is the same after the infection by virus, which was in the cells, and the same position are located in the plasma membrane and tonoplast, But superoxide dismutas is mainly distributed on the plasma membrane for resistant varieties, while it was distributed on the'vacuole membrane for susceptible ones. Resistant varieties SOD activity was not only significantly higher than the control group, but also significantly higher than susceptible varieties, which were infected by virus. The results of POD and SOD intracellular distribution and physiological and biochemical detection in Resistant sugar beet varieties were the same with susceptible varieties, higher activity of POD and SOD in resistant sugar beet plexus root disease was one of the important mechanisms of biochemical markers at the subcellular level.
4. By the methods of histochemical staining and electron microscope cytochemical, We have Studied the Influence of BNYVV on lignin and HRGP, which were in sugar beet cell wall, clearly to see the relationships between different resistance beet cell wall lignin as well as different HRGP accumulation and distribution and the nature of anti plexus root disease in sugar beet roots. The results showed that no matter it was the resistant sugar beet varieties or not, no matter the beet varieties were infected by BNYVV before or after, there are lignin and HRGP deposition in their roots and all catheters on vein transverse. In addition, lignin deposition was shown gradually from deep to shallow, which was throughout the parenchyma cell walls from the outside to the inside in roots; lignin and HRGP accumulation in resistant varieties is more significant than susceptible varieties and its control group; it was notable that it also showed black granular sediment on sugar beet cell vacuoie membrane when determinated the cell wall surface glycoprotein, and vacuole membrane deposition was deeper·in disease-resistant varieties than susceptible varieties and the control group. It explains that HRGP was involved in the synthesis of lignin between sugar beet and BNYVV interaction system, the rapid accumulation of Lignin and HRGP in Sugar beet tissue was one of the manifestations of rhizomania.
1.利用光学显微技术和透射电镜技术研究了甜菜抗、感丛根病品种与BNYVV互作过程中的形态学和超微结构特征。结果表明,不同寄主与病毒互作体系的甜菜块根形态学和超微结构上均表现出明显的差异,而甜菜叶脉受该病毒侵染后其形态学解剖结构无差异,但超微结构受到影响。形态学水平上,抗、感病品种表现为表皮破坏,皮层薄壁细胞组织产生裂隙,大多细胞都脱落,木质部附近的木薄壁细胞破损,完全丧失了对根部的保护作用。其中感病品种比抗病品种表现被伤害程度更为严重,其皮层薄壁细胞几乎全部脱落,生长速度缓慢,根部发育受阻。而叶脉的形态解剖结构与对照比较,未发现异常变化。另外抗病品种比感病品种有更发达的维管束结构。在亚细胞水平上,BNYVV对感病品种的细胞超微结构破坏严重,整个细胞变形,空虚化,细胞核结构发生紊乱,线粒体和高尔基体明显增多,细胞质中小液泡增多,在液泡膜边缘可见到一些纤细丝状物质的圆形或卵圆形小泡突入液泡中。叶脉细胞叶绿体完全瓦解,出现许多嗜锇颗粒。而抗病品种细胞超微结构破坏较轻,寄主细胞产生一系列显著的结构防卫反应：形成细胞壁沉积物及液泡膜上显示出黑色颗粒状沉积物等。说明组织形态结构变化和细胞器病理变化与甜菜抗丛根病性有关。
2.利用生理生化方法和电镜细胞化学标记技术研究了甜菜-BNYVV互作过程中H2O2和O2-·产生的时间变化特征和空间分布定位。研究结果表明,甜菜抗、感丛根病体系均在侵染早期出现大量H2O2和O2-·,其中抗病体系的H2O2和O2-·产生量明显高于感病体系。H2O2在抗病和感病体系中的分布位置基本相似,多分布在块根、叶脉细胞的液泡膜和质膜上,叶脉细胞间隙也有H2O2的分布,而O2-·在抗病品种块根与叶脉细胞的质膜上定位,感病品种的则在液泡膜上被发现。而且感病体系H202和O2-·沉积量明显弱于抗病体系。H2O2和O2-·产生量和分布与甜菜抗丛根病性有密切联系,不同部位定位的H2O2和O2-·作为信号分子,参与了甜菜对病毒侵染的防御反应。
3.采用生化检测及酶细胞化学方法研究了甜菜与BNYVV互作过程中POD和SOD的活性及其在细胞内分布特征。结果表明,病毒感染前POD主要定位在甜菜块根细胞细胞壁及叶脉细胞线粒体和细胞间隙,当病毒感染后,抗、感病甜菜品种块根皮层细胞的细胞壁、质膜和液泡膜上的POD及叶脉薄壁细胞的细胞壁、质膜、液泡膜、线粒体、细胞间隙上的POD活性均比其对照显著升高,抗、感病甜菜品种在POD分布部位上没有区别,但两者沉积量有所差异,抗病品种明显高于感病品种；SOD在未感染病毒的抗、感病甜菜块根皮层薄壁细胞质膜与液泡膜,叶脉厚角组织薄壁细胞质膜及叶脉细胞线粒体、细胞间隙上被发现,病毒侵染后甜菜抗、感病品种块根SOD在细胞内分布位置相同都分布在质膜和液泡膜,但抗病品种以质膜上分布为主,感病品种以液泡膜上分布为主。抗病品种SOD活性不仅明显高于其对照,也明显高于病毒侵染后的感病品种。甜菜抗、感病品种POD及SOD细胞内分布的沉积量与生理生化检测的酶活性结果一致,从亚细胞学水平说明高活性的POD及SOD是甜菜抗丛根病性生化标记的重要生理机制之一。
4.通过组织化学染色法及电镜细胞化学方法研究BNYVV对甜菜细胞壁中木质素和HRGP的影响。结果显示,BNYVV侵染前后在甜菜抗、感病品种的块根与叶脉横切面的所有导管上均有木质素和HRGP的沉积,块根薄壁组织的细胞壁由外向内也逐渐显示出由深至浅的木质素沉积；抗病品种木质素和HRGP的积累量较感病品种和其对照增加更为显著；说明在甜菜与BNYVV互作体系中,HRGP参与木质素的合成,木质素和HRGP在甜菜组织内快速积累是甜菜抗丛根病性的表现之一。
Sugar beet (Beta vulgaris L.) is important sugar and cash crops in our country even in the world. Under the various attacked of sugar beet diseases not only have causing huge economic losses in the sugar beet production industry but also restricted the development of the sugar industry. Rhizomania caused by Beet necrotic yellow vein virus (BNYVV) and mediated by Polymyxa betae which was one of the most serious soil-born diseases. Recently, numerous researchs had studied the mechanism of rhizomania at domestic and abroad were main under the views of genomics, molecular biology, physiological and biochemical. No research studying the mechanism of rhizomania and its resistance signal transduction pathway from the cytological at present. Therefore, in this study, sugar beet varieties and BNYVV interaction system were the research objects, the morphological and cytological features of the two objects were studied. Also, to make sure the time-varying characteristics and located the spatial distribution in the sub-cellular level of the production of the active oxygen species (H2O2and O2-·) and the protection of enzymes (POD and SOD) in the interaction system, the physiological and biochemical methods and electron microscopic cytochemistry labeling technique were used, And we have detected the locating differences within organization of the lignin and the law of cell wall glycoprotein accumulated in the host cell in the interactions system to observed the roles of the active oxygen metabolism and cell wall components in the sugar beet resistance to rhizomania disease resistance, then lay a theoretical foundation for the further study of the signal conduction mechanism of sugar beet resistance to rhizomania disease. The main results obtained through the study were as follows:
1. We have utilized optical microscopy and transmission electron microscopy to reveal the structural features of the morphology and ultrastructure between the interaction of the resistant and susceptible beet varieties and the BNYVV. The results showed that the morphology and ultrastructure of the sugar beet roots of the resistant and susceptible interaction system, infected by BNYVV, showed significant differences, the ultrastructure of the sugar beet vein have been seriously affected too, but the morphological anatomical structure of the sugar beet vein was no difference. The performance of the Morphology happened in the resistant and susceptible sugar beet included the epidermal destruction, the fracture of the cortical parenchyma tissue, the shed in most cell and the damage of the wood parenchyma cells near the xylem, the protection for the roots loss completely. The performance of the susceptible beet is more serious than the resistant beet. In the susceptible beet, the cortical parenchyma cells were almost all off, growing slowly and the root's development was blocked. However, there are no abnormal changes in the morphological anatomical structure of the beet vein. In addition, the resistant varieties have a well developed vascular bundle structure than susceptible varieties. When observed from the sub-cellular level, we found that BNYVV resulted in a series of damaged on cell ultrastructure of the susceptible beet, such as the deformation, empty and the structure of the nuclear is disorder, the mitochondria and the Golgi apparatus was significantly increased, small and medium-sized vacuoles were increased in the cytoplasmic. We observed some round or oval vesicles containing thin filaments broke into the vacuole at the edge of the vacuolar membrane. In the Vein cells the chloroplast disintegrated completely and a abundance of Osmiophilic globule emerged. Destruction of the cell ultrastructure of resistant varieties was lighter, a series of significant structural defense reaction produced in:the host cells:there was sediments in the cell wall and black granular sediment showed in the vacuolar membrane. These indicated that the description of the morphological changes of the tissue and the pathological changes of the organelle were relative to the resistance to rhizomania.
2. We have utilized Physiological and biochemical methods and electron microscopy cytochemistry labeling technique to study the time-varying characteristics and the orientation of spatial distribution of the H2O2and O2-·generated between the process of the beet-BNYVV interaction. A large number of H2O2and O2·detected both in the resistant and susceptible beet, but the production in the resistant beet was higher than the susceptible beet significantly. H2O2whose position of distribution was similar in the resistant and susceptible beet, which was located in the roots, the tonoplast and plasma membrane of the veins cell, and exist in the gap of the veins cell also. The O2-·is detected in the vein and the plasma membran of the veins cell in the resistant, but found that in the tonoplast in the susceptible beet. However, the content of H2O2and O2-·was distinctly different in the different beet systems, the content in the susceptible system was significantly lower than the resistant system. The results suggested that the production and distribution of the H2O2and O2-·were significantly different in the different interactive system, these difference were close with the beet's resistance to the rhizomania. As a signaling molecule, the H2O2and O2-·located in different positions involved in the defense response of the beet to virus infection.
3. Research was carried on POD and SOD activity and their intracellular distribution characteristics in sugar beet and BNYVV interaction processes by using biochemical detection and enzyme cytochemical methods. The results showed that before the viral infection, the POD was mainly localized in the root cell wall and mitochondria of the vein cells and the gaps between the cells. after the virus infection, in the resistant and susceptible sugar beet varieties, the POD activity in the cell wall, plasma membrane and tonoplast of the tuberous root cortex cells, and in the cell wall, plasma membrane, tonoplast, mitochondria and the gap of the veins cells is significantly increased than that of the control group. Although the distribution of POD was almost same in both beet system, but the deposition in the former was significantly higher in the latter. When the resitant and susceptcible were not infected virus, SOD was defected in plasma membrane and tonoplast of the beet root cortical parenchyma cell, cytoplasmic membrane of vein collenchyma thin-walled, mitochondria of veins and the gaps of the cells. The distribution of SOD in susceptible varieties and resistant sugar beet root is the same after the infection by virus, which was in the cells, and the same position are located in the plasma membrane and tonoplast, But superoxide dismutas is mainly distributed on the plasma membrane for resistant varieties, while it was distributed on the'vacuole membrane for susceptible ones. Resistant varieties SOD activity was not only significantly higher than the control group, but also significantly higher than susceptible varieties, which were infected by virus. The results of POD and SOD intracellular distribution and physiological and biochemical detection in Resistant sugar beet varieties were the same with susceptible varieties, higher activity of POD and SOD in resistant sugar beet plexus root disease was one of the important mechanisms of biochemical markers at the subcellular level.
4. By the methods of histochemical staining and electron microscope cytochemical, We have Studied the Influence of BNYVV on lignin and HRGP, which were in sugar beet cell wall, clearly to see the relationships between different resistance beet cell wall lignin as well as different HRGP accumulation and distribution and the nature of anti plexus root disease in sugar beet roots. The results showed that no matter it was the resistant sugar beet varieties or not, no matter the beet varieties were infected by BNYVV before or after, there are lignin and HRGP deposition in their roots and all catheters on vein transverse. In addition, lignin deposition was shown gradually from deep to shallow, which was throughout the parenchyma cell walls from the outside to the inside in roots; lignin and HRGP accumulation in resistant varieties is more significant than susceptible varieties and its control group; it was notable that it also showed black granular sediment on sugar beet cell vacuoie membrane when determinated the cell wall surface glycoprotein, and vacuole membrane deposition was deeper·in disease-resistant varieties than susceptible varieties and the control group. It explains that HRGP was involved in the synthesis of lignin between sugar beet and BNYVV interaction system, the rapid accumulation of Lignin and HRGP in Sugar beet tissue was one of the manifestations of rhizomania.
引文
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