B型烟粉虱为害后烟草的生理响应
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摘要
B型烟粉虱是一种世界性的入侵害虫。该虫自入侵我国后,已在局部烟区爆发成灾,并且成为了烟草上的重要害虫。B型烟粉虱为害能够造成多种的寄主生理紊乱,如西葫芦银叶,莴苣叶茎变白,部分观赏植物叶片失绿等现象,并引起某些植物光合速率的下降。而我们的前期研究发现B型烟粉虱为害后烟草叶片出现明显的白脉现象,并对烟草的生长发育产生明显不利影响。同时发现,B型烟粉虱为害造成烟草叶片中防御酶活性增加,诱导了烟草防御的水杨酸信号途径,而抑制了茉莉酸信号途径。本文在上述研究的基础上,深入研究了B型烟粉虱为害烟草后对光合作用的影响和机制,通过测定诱导的烟草植株韧皮部汁液中相关防御物质的变化,探讨B型烟粉虱取食后烟草生理生化变化在对蚜虫等其它刺吸式口器昆虫抗虫性中的的作用。本文对于阐明B型烟粉虱对烟草的为害机制,科学评价其生态风险性具有重要的理论意义。本研究主要结果如下：
1.B型烟粉虱若虫为害对烟草叶片光合作用造成不利影响,并具有明显的空间和时间效应。随着B型烟粉虱若虫为害时间的推移,虫体叶光合速率呈现逐渐降低的趋势,在B型烟粉虱取食15d,20d时相对于对照明显降低了38.22%和54.28%；而系统白脉叶的光合速率呈现先降低后升高的趋势,在烟粉虱若虫取食15d和20d时相对于对照明显降低了40.98%和18.48%,表明B型烟粉虱为害不仅对虫体叶的光合作用造成不利影响,而系统白脉叶的光合作用同样受到抑制。B型烟粉虱为害烟草10d后才对烟草叶片的光合作用造成影响,表明B型烟粉虱为害对烟草光合作用的影响具有时间效应。B型烟粉虱为害后烟草叶片叶绿素a,b和类胡萝卜含量降低,尤其是是叶绿素a在空间和时间上降低最为明显。
2.B型烟粉虱取食对烟草光系统Ⅱ(PS Ⅱ)结构和功能造成了不利影响,进而影响烟草的光合效率。植物的光系统Ⅱ是植物光合机构中对环境变化最敏感的部位,也是光合作用的重要部位,与植物光合作用过程中光能的吸收、转换和光合电子传递的有关。B型烟粉虱为害烟草后虫体叶和系统白脉叶的单位面积反应中心数量(RC/CS)均明显降低,而PS Ⅱ反应中心的关闭程度(1-qP)却明显升高,同时使供体侧的放氧复合体(OEC)受到严重的伤害,对光系统Ⅱ的结构造成较大伤害。PS Ⅱ反应中心的失活和关闭限制了PS Ⅱ的电子向下游的传递速率,使PS Ⅱ吸收的光能转变成化学能的比例降低。同时B型烟粉虱为害使烟草PS Ⅱ电子传递受到抑制,这些电子传递中受抑制的位点包括了放氧复合体和QA到QB间的电子传递过程。B型烟粉虱为害影响了烟草虫体叶和系统白脉叶光系统的能量流动,B型烟粉虱为害导致虫体叶和系统白脉叶热耗散掉的能量(DIo/RC)的增加和能量利用效率(φpo,Ψo和qφE0)下降,导致过剩激发能增加,活性氧过氧化氢积累增加,并且这种影响具有明显的系统传导性,可能与B型烟粉虱诱导烟草的SA信号途径防御反应有关。
3.通过SDS-PAGE电泳发现,B型烟粉虱3龄若虫为害烟草后,白脉叶相对于对照有9种蛋白被诱导上调表达,其中4种蛋白是特异性表达,而其它5种蛋白被诱导上调；而虫体叶中只有两种蛋白被诱导上调。这表明B型烟粉虱可能诱导了相关的防御酶的表达,并且在白脉叶上较虫体叶被诱导表达更强烈。
4.通过测定烟草韧皮部汁液中防御物质,明确了B型烟粉虱为害后烟草韧皮部汁液中相关防御蛋白以及酚类物质的时空变化。B型烟粉虱为害后虫体叶位和系统白脉叶位韧皮部汁液中多酚氧化酶(PPO),β-1,3-葡聚糖酶,几丁质酶的活性均较对照明显增加,但是在系统白脉叶位的升高幅度明显高于虫体叶位,特别是在15d和20d的时候。B型烟粉虱为害后韧皮部汁液的过氧化物酶(POD)和苯丙氨酸解氨酶(PAL)在虫体叶的升高幅度明显高于系统白脉叶位。过氧化氢酶(CAT)在系统白脉叶位韧皮部汁液被B型烟粉虱取食诱导增加,而在虫体叶位中却被抑制。虫体叶韧皮部汁液中抗坏血酸过氧化物酶(APX)和总酚含量与对照相比没有明显差异,在系统白脉叶中也只是在B型烟粉虱为害20d,25d时才有一定的差异。由此结合前期的蚜虫生测试验分析,发现B型烟粉虱为害后诱导烟草韧皮部汁液中PPO,β-1-3-葡聚糖酶,几丁质酶及CAT与烟草对烟蚜的防御有一定的关系。而抗坏血酸过氧化物酶与酚类物质与烟草防御烟蚜的关系不大。
Bemisia tabaci (Gennadius) biotype B is a worldwide invasive pest. It has been a key pest of tobacco, and broke out in parts of tobacco-growing areas when it invades in China. B. tabaci biotype B feeding cause plant systemic or physiological disorders include silverleaf of squash, stem blanching of lettuce, white streaking of Brassica, and chlorosis of new foliage in numerous ornamental plants. Furthermore B. tabaci biotype B feeding caused reduction of net photosynthetic rate. Our previous studies showed that the feeding of B. tabaci biotype B directly inhibited the growth of tobacco and induced obviously white vein of tobacco leaves. It is indicated that the physiological response of tobacco was affected by B. tabaci biotype B. Photosynthesis is the most basic the physiological response of the plant. However, the influence mechanism of B. tabaci biotype B on the net photosynthetic rate (Pn) is also unknown. Meanwhile, the feeding of B. tabaci biotype B caused increase the activity of defense enzymes of tobacco leaves。Furthermore B. tabaci biotype B induces salicylic acid defenses and suppresses effectual jasmonic acid defenses. According to the above study results, we chose the model plant-tobacco as the host plant to investigate the effect and possible mechanism of Bemisia tabaci (Gennadius) B-biotype infestation on photosynthesis of tobacco plants. We also studied the temporal and spatial change of defense enzyme of phloem sap of tobacco plants induced by B. tabaci biotype B. We discussed the role of physiological and biochemical response of plants in the defense responses to other phloem feeder of tobacco plants induced by B. tabaci biotype B. This research had important academic meaning for showing the damage mechanism of B. tabaci biotype B to tobacco plants and scientifically assessing the ecological risk of B. tabaci biotype B. Major results as follows:
1. There was temporal and spatial disadvantaged influence on photosynthesis of tobacco plants damaged by B. tabaci biotype B. While net photosynthetic rate (Pn) in infested leaves exhibited a dramatic decrease as time progressed, especially on15d and20d, the decrease was38.22%and54.2%; The Pn in systemic leaves of treatment displayed a decrease first and then an increase, but the level decreased by41.0%and18.5%than the control on15d and20d, respectively. This indicated that the inhibition to the photosynthesis caused by B. tabaci infestation is not only in the infested leaves, but also in the systemic leaves. The photosynthesis of tobacco leaves was affected only10days after B. tabaci infestation. The results show that the adverse effect of photosynthesis caused by B. tabaci infestation with time effect. B. tabaci infestation altered the content of chlorophyll, especially the chlorophyll a.
2. B. tabaci infestation damaged the photosystem II (PS II) of tobacco leaves. Photosystem II is an important part of the photosynthese. It is the most sensitive part of photosynthetic apparatus to environmental change. The PS II is associated with the energy of light, absorption, energy conversion, electron transfer. B. tabaci infestation reduced the density of active reaction centers per excited cross-section (RC/CS) both in infested and systemic leaves (P<0.05). B. tabaci infestation caused the closure degree of the PS II reaction centers (1-qP) increase in infested and systemic leaves. B. tabaci infestation severely damaged the oxygenevolving complex (OEC). This indicated the structure of photosystem II was damaged. The reaction centers were damaged which decrease the rate of electron transporting. The inhibited points of the electron transporting included the OEC and the electron transporting from Qa to QB.Energy flux of PS Ⅱ was also affected by the B. tabaci infestation. B. tabaci infestation increased the energy dissipation per active reaction centers (DIo/RC) and decreased the energy utilization efficiency (cppo, Ψo and φpEo). It increased the excess excitation energy and the production of ROS, which could improve the salicylic acid defenses induced by B. tabaci. The effects caused by B. tabaci in tobacco leaves were systematically conductive.
3. The proteins of tobacco leaves were assayed by sodium dodecyl sulfate-polyacrylamide gel electrophoresis (SDS-PAGE). The3rd instars of B. tabaci biotype B feeding induced the expression of nine proteins increase in systemic leaves. Four proteins of these were specific induced; and five of them were just induced. In infested leaves, there were only two proteins were just induced
4. We investigated the temporal and spatial change of main defense enzyme activity and nutrition content of phloem sap of tobacco plants infested by B. tabaci. The phloem was the feeding sites for the phloem feeder. There are many proteins and secondary metabolisms that are related to defense response in the phloem sap. There was difference between the changes of the activity of polyphenol oxidase (PPO), β-1-3-glucanase and chitinase of phloem sap in local and systemic leaves. However, the increase rate of their activity of phloem sap in local leaves was lower than that in systemic leaves. The increase rate of their activity of peroxidase (POD) and phenylalanine ammonialyase (PAL) of phloem sap in systemic leaves was lower than that in local leaves. The activity of catalase (CAT) of phloem sap in systemic leaves was induced increase by B. tabaci infestation, however, in local leaves it decreased. The activity of ascorbate peroxidase (APX) and content of total phenol of phloem sap were not significantly changed by B. tabaci infestation in local leaves, which rarely increased in systemic leaves on20d and25d. This indicated that PPO, β-1-3-glucanase, chitinase and CAT of phloem sap are related to the defense response of tobacco plant to aphid.
1.B型烟粉虱若虫为害对烟草叶片光合作用造成不利影响,并具有明显的空间和时间效应。随着B型烟粉虱若虫为害时间的推移,虫体叶光合速率呈现逐渐降低的趋势,在B型烟粉虱取食15d,20d时相对于对照明显降低了38.22%和54.28%；而系统白脉叶的光合速率呈现先降低后升高的趋势,在烟粉虱若虫取食15d和20d时相对于对照明显降低了40.98%和18.48%,表明B型烟粉虱为害不仅对虫体叶的光合作用造成不利影响,而系统白脉叶的光合作用同样受到抑制。B型烟粉虱为害烟草10d后才对烟草叶片的光合作用造成影响,表明B型烟粉虱为害对烟草光合作用的影响具有时间效应。B型烟粉虱为害后烟草叶片叶绿素a,b和类胡萝卜含量降低,尤其是是叶绿素a在空间和时间上降低最为明显。
2.B型烟粉虱取食对烟草光系统Ⅱ(PS Ⅱ)结构和功能造成了不利影响,进而影响烟草的光合效率。植物的光系统Ⅱ是植物光合机构中对环境变化最敏感的部位,也是光合作用的重要部位,与植物光合作用过程中光能的吸收、转换和光合电子传递的有关。B型烟粉虱为害烟草后虫体叶和系统白脉叶的单位面积反应中心数量(RC/CS)均明显降低,而PS Ⅱ反应中心的关闭程度(1-qP)却明显升高,同时使供体侧的放氧复合体(OEC)受到严重的伤害,对光系统Ⅱ的结构造成较大伤害。PS Ⅱ反应中心的失活和关闭限制了PS Ⅱ的电子向下游的传递速率,使PS Ⅱ吸收的光能转变成化学能的比例降低。同时B型烟粉虱为害使烟草PS Ⅱ电子传递受到抑制,这些电子传递中受抑制的位点包括了放氧复合体和QA到QB间的电子传递过程。B型烟粉虱为害影响了烟草虫体叶和系统白脉叶光系统的能量流动,B型烟粉虱为害导致虫体叶和系统白脉叶热耗散掉的能量(DIo/RC)的增加和能量利用效率(φpo,Ψo和qφE0)下降,导致过剩激发能增加,活性氧过氧化氢积累增加,并且这种影响具有明显的系统传导性,可能与B型烟粉虱诱导烟草的SA信号途径防御反应有关。
3.通过SDS-PAGE电泳发现,B型烟粉虱3龄若虫为害烟草后,白脉叶相对于对照有9种蛋白被诱导上调表达,其中4种蛋白是特异性表达,而其它5种蛋白被诱导上调；而虫体叶中只有两种蛋白被诱导上调。这表明B型烟粉虱可能诱导了相关的防御酶的表达,并且在白脉叶上较虫体叶被诱导表达更强烈。
4.通过测定烟草韧皮部汁液中防御物质,明确了B型烟粉虱为害后烟草韧皮部汁液中相关防御蛋白以及酚类物质的时空变化。B型烟粉虱为害后虫体叶位和系统白脉叶位韧皮部汁液中多酚氧化酶(PPO),β-1,3-葡聚糖酶,几丁质酶的活性均较对照明显增加,但是在系统白脉叶位的升高幅度明显高于虫体叶位,特别是在15d和20d的时候。B型烟粉虱为害后韧皮部汁液的过氧化物酶(POD)和苯丙氨酸解氨酶(PAL)在虫体叶的升高幅度明显高于系统白脉叶位。过氧化氢酶(CAT)在系统白脉叶位韧皮部汁液被B型烟粉虱取食诱导增加,而在虫体叶位中却被抑制。虫体叶韧皮部汁液中抗坏血酸过氧化物酶(APX)和总酚含量与对照相比没有明显差异,在系统白脉叶中也只是在B型烟粉虱为害20d,25d时才有一定的差异。由此结合前期的蚜虫生测试验分析,发现B型烟粉虱为害后诱导烟草韧皮部汁液中PPO,β-1-3-葡聚糖酶,几丁质酶及CAT与烟草对烟蚜的防御有一定的关系。而抗坏血酸过氧化物酶与酚类物质与烟草防御烟蚜的关系不大。
Bemisia tabaci (Gennadius) biotype B is a worldwide invasive pest. It has been a key pest of tobacco, and broke out in parts of tobacco-growing areas when it invades in China. B. tabaci biotype B feeding cause plant systemic or physiological disorders include silverleaf of squash, stem blanching of lettuce, white streaking of Brassica, and chlorosis of new foliage in numerous ornamental plants. Furthermore B. tabaci biotype B feeding caused reduction of net photosynthetic rate. Our previous studies showed that the feeding of B. tabaci biotype B directly inhibited the growth of tobacco and induced obviously white vein of tobacco leaves. It is indicated that the physiological response of tobacco was affected by B. tabaci biotype B. Photosynthesis is the most basic the physiological response of the plant. However, the influence mechanism of B. tabaci biotype B on the net photosynthetic rate (Pn) is also unknown. Meanwhile, the feeding of B. tabaci biotype B caused increase the activity of defense enzymes of tobacco leaves。Furthermore B. tabaci biotype B induces salicylic acid defenses and suppresses effectual jasmonic acid defenses. According to the above study results, we chose the model plant-tobacco as the host plant to investigate the effect and possible mechanism of Bemisia tabaci (Gennadius) B-biotype infestation on photosynthesis of tobacco plants. We also studied the temporal and spatial change of defense enzyme of phloem sap of tobacco plants induced by B. tabaci biotype B. We discussed the role of physiological and biochemical response of plants in the defense responses to other phloem feeder of tobacco plants induced by B. tabaci biotype B. This research had important academic meaning for showing the damage mechanism of B. tabaci biotype B to tobacco plants and scientifically assessing the ecological risk of B. tabaci biotype B. Major results as follows:
1. There was temporal and spatial disadvantaged influence on photosynthesis of tobacco plants damaged by B. tabaci biotype B. While net photosynthetic rate (Pn) in infested leaves exhibited a dramatic decrease as time progressed, especially on15d and20d, the decrease was38.22%and54.2%; The Pn in systemic leaves of treatment displayed a decrease first and then an increase, but the level decreased by41.0%and18.5%than the control on15d and20d, respectively. This indicated that the inhibition to the photosynthesis caused by B. tabaci infestation is not only in the infested leaves, but also in the systemic leaves. The photosynthesis of tobacco leaves was affected only10days after B. tabaci infestation. The results show that the adverse effect of photosynthesis caused by B. tabaci infestation with time effect. B. tabaci infestation altered the content of chlorophyll, especially the chlorophyll a.
2. B. tabaci infestation damaged the photosystem II (PS II) of tobacco leaves. Photosystem II is an important part of the photosynthese. It is the most sensitive part of photosynthetic apparatus to environmental change. The PS II is associated with the energy of light, absorption, energy conversion, electron transfer. B. tabaci infestation reduced the density of active reaction centers per excited cross-section (RC/CS) both in infested and systemic leaves (P<0.05). B. tabaci infestation caused the closure degree of the PS II reaction centers (1-qP) increase in infested and systemic leaves. B. tabaci infestation severely damaged the oxygenevolving complex (OEC). This indicated the structure of photosystem II was damaged. The reaction centers were damaged which decrease the rate of electron transporting. The inhibited points of the electron transporting included the OEC and the electron transporting from Qa to QB.Energy flux of PS Ⅱ was also affected by the B. tabaci infestation. B. tabaci infestation increased the energy dissipation per active reaction centers (DIo/RC) and decreased the energy utilization efficiency (cppo, Ψo and φpEo). It increased the excess excitation energy and the production of ROS, which could improve the salicylic acid defenses induced by B. tabaci. The effects caused by B. tabaci in tobacco leaves were systematically conductive.
3. The proteins of tobacco leaves were assayed by sodium dodecyl sulfate-polyacrylamide gel electrophoresis (SDS-PAGE). The3rd instars of B. tabaci biotype B feeding induced the expression of nine proteins increase in systemic leaves. Four proteins of these were specific induced; and five of them were just induced. In infested leaves, there were only two proteins were just induced
4. We investigated the temporal and spatial change of main defense enzyme activity and nutrition content of phloem sap of tobacco plants infested by B. tabaci. The phloem was the feeding sites for the phloem feeder. There are many proteins and secondary metabolisms that are related to defense response in the phloem sap. There was difference between the changes of the activity of polyphenol oxidase (PPO), β-1-3-glucanase and chitinase of phloem sap in local and systemic leaves. However, the increase rate of their activity of phloem sap in local leaves was lower than that in systemic leaves. The increase rate of their activity of peroxidase (POD) and phenylalanine ammonialyase (PAL) of phloem sap in systemic leaves was lower than that in local leaves. The activity of catalase (CAT) of phloem sap in systemic leaves was induced increase by B. tabaci infestation, however, in local leaves it decreased. The activity of ascorbate peroxidase (APX) and content of total phenol of phloem sap were not significantly changed by B. tabaci infestation in local leaves, which rarely increased in systemic leaves on20d and25d. This indicated that PPO, β-1-3-glucanase, chitinase and CAT of phloem sap are related to the defense response of tobacco plant to aphid.
引文
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