激发子诱导植物防卫反应过程中的信号分子
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摘要
一般将能够诱导寄主防卫反应的生物来源和非生物来源的物质统称为激发子。第六次国际植物组织培养大会规定来自于植物的激发子称为内源激发子，来自于微生物的激发子的称为真激发子。随着植物防卫机理的研究向分子水平的深入，越来越多的实验证明激发子和防卫反应可作为研究植物信号识别及胞内信息传递的良好实验体系。
在本研究工作中，金瓜炭疽细胞壁激发子与脱乙酰几丁质是来源于真菌细胞壁的真激发子，人参细胞壁降解物与寡聚半乳糖醛酸是来源于植物细胞壁的内源激发子。我们也通过昆虫取食与模拟微重力生物学效应来研究植物的防卫反应。
人参是我国传统中草药，具有安神补气，增加免疫的功能，人参皂苷是人参中有效药用成分。我们开始实验的目的是利用激发子提高人参皂苷的含量。经过大量筛选，先后发现金瓜炭疽激发子，脱乙酰几丁质，寡聚半乳糖醛酸都可以明显诱导人参细胞中皂苷含量的提高。为了进一步研究激发子诱导人参皂苷含量提高的机理，我们发现激发子诱导了人参细胞过氧化氢，一氧化氮与茉莉酸的产生，这三种信号分子单独处理人参细胞都可以诱导人参皂苷的合成与防卫反应，抑制激发子诱导的这三种信号分子的产生，可以同时抑制激发子诱导的人参皂苷的积累。这些结果表明过氧化氢，一氧化氮与茉莉酸介导了激发子诱导的人参皂苷的合成。激发子诱导人参细胞皂苷含量升高的过程中，这三种信号分子并不是孤立的，它们之间相互联系，相互影响，共同介导人参皂苷的合成。蛋白激酶与胞内钙离子通过调节信号分子的产生而影响激发子诱导的人参皂苷产生。我们利用激发子在水稻与拟南芥上也进行了相关实验。结果表明一氧化氮介导了真菌激发子诱导的水稻过敏性死亡与抗性相关基因的转录。激发子诱导胞内钙离子的升高以及过氧化氢的产生，对于激发子诱导的抗性相关基因的转录是必须的。
除了上述各类激发子可以诱导植物的防卫反应外，其它刺激因子，如机械伤害，昆虫取食，盐碱，高温，冷害，失重等，都可能诱导植物的防卫反应。棉铃虫取食可以诱导蕃茄小苗过氧化氢的产生，激活水杨酸与茉莉酸代谢途径，并且过氧化氢可能作用于水杨酸与茉莉酸途径的上游。失重或微重力也可以看作为一类刺激因子诱导植物发生反应，在我们的实验中，采用回转方式培养人参细胞，即模拟微重力生物学效应条件，可以诱导人参细胞皂苷含量的升高。这种培养方式也可以诱导人参细胞钙调蛋白基因的波动性表达。抑制钙调蛋白基因的转录，也可以抑制回转方式细胞培养所诱导的人参皂苷含量的升高。这些结果表明钙调蛋白可能介导了模拟微重力生物学效应条件所诱导的人参皂苷的合成。
Endogenous elicitor activity, termed CDW, was prepared from the cell walls of suspension-cultured ginseng (Panax ginseng C.A. Meyer) cells via cellulase degradation. CDW activated in vitro the NADPH oxidase activity of isolated plasma membranes and stimulated in vivo H2O2 generation by ginseng cell suspensions. CDW also increased the activity of phenylalanine ammonia lyase (PAL), expression of a P. ginseng squalene epoxidase (sqe) gene and saponin synthesis. NADPH oxidase inhibitors inhibited both in vitro NADPH oxidase activity and in vivo H2O2 generation. Induction of PAL activity, saponin synthesis and sqe gene expression were all inhibited by such inhibitor treatments and reduced by incubation with catalase and H2O2 scavengers. These data indicate that activation of NADPH oxidase and generation of H2O2 are essential signalling events mediating defence responses induced by the endogenous elicitor(s) present in CDW.
在本研究工作中，金瓜炭疽细胞壁激发子与脱乙酰几丁质是来源于真菌细胞壁的真激发子，人参细胞壁降解物与寡聚半乳糖醛酸是来源于植物细胞壁的内源激发子。我们也通过昆虫取食与模拟微重力生物学效应来研究植物的防卫反应。
人参是我国传统中草药，具有安神补气，增加免疫的功能，人参皂苷是人参中有效药用成分。我们开始实验的目的是利用激发子提高人参皂苷的含量。经过大量筛选，先后发现金瓜炭疽激发子，脱乙酰几丁质，寡聚半乳糖醛酸都可以明显诱导人参细胞中皂苷含量的提高。为了进一步研究激发子诱导人参皂苷含量提高的机理，我们发现激发子诱导了人参细胞过氧化氢，一氧化氮与茉莉酸的产生，这三种信号分子单独处理人参细胞都可以诱导人参皂苷的合成与防卫反应，抑制激发子诱导的这三种信号分子的产生，可以同时抑制激发子诱导的人参皂苷的积累。这些结果表明过氧化氢，一氧化氮与茉莉酸介导了激发子诱导的人参皂苷的合成。激发子诱导人参细胞皂苷含量升高的过程中，这三种信号分子并不是孤立的，它们之间相互联系，相互影响，共同介导人参皂苷的合成。蛋白激酶与胞内钙离子通过调节信号分子的产生而影响激发子诱导的人参皂苷产生。我们利用激发子在水稻与拟南芥上也进行了相关实验。结果表明一氧化氮介导了真菌激发子诱导的水稻过敏性死亡与抗性相关基因的转录。激发子诱导胞内钙离子的升高以及过氧化氢的产生，对于激发子诱导的抗性相关基因的转录是必须的。
除了上述各类激发子可以诱导植物的防卫反应外，其它刺激因子，如机械伤害，昆虫取食，盐碱，高温，冷害，失重等，都可能诱导植物的防卫反应。棉铃虫取食可以诱导蕃茄小苗过氧化氢的产生，激活水杨酸与茉莉酸代谢途径，并且过氧化氢可能作用于水杨酸与茉莉酸途径的上游。失重或微重力也可以看作为一类刺激因子诱导植物发生反应，在我们的实验中，采用回转方式培养人参细胞，即模拟微重力生物学效应条件，可以诱导人参细胞皂苷含量的升高。这种培养方式也可以诱导人参细胞钙调蛋白基因的波动性表达。抑制钙调蛋白基因的转录，也可以抑制回转方式细胞培养所诱导的人参皂苷含量的升高。这些结果表明钙调蛋白可能介导了模拟微重力生物学效应条件所诱导的人参皂苷的合成。
Endogenous elicitor activity, termed CDW, was prepared from the cell walls of suspension-cultured ginseng (Panax ginseng C.A. Meyer) cells via cellulase degradation. CDW activated in vitro the NADPH oxidase activity of isolated plasma membranes and stimulated in vivo H2O2 generation by ginseng cell suspensions. CDW also increased the activity of phenylalanine ammonia lyase (PAL), expression of a P. ginseng squalene epoxidase (sqe) gene and saponin synthesis. NADPH oxidase inhibitors inhibited both in vitro NADPH oxidase activity and in vivo H2O2 generation. Induction of PAL activity, saponin synthesis and sqe gene expression were all inhibited by such inhibitor treatments and reduced by incubation with catalase and H2O2 scavengers. These data indicate that activation of NADPH oxidase and generation of H2O2 are essential signalling events mediating defence responses induced by the endogenous elicitor(s) present in CDW.
引文
1. 王钧 植物抗病反应的分子机理 见 余叔文 ，汤章城主编 植物生理与分子生物学 科学出版社 北京 1998
2. 武维华 主编 植物生理学 科学出版社 北京 2003
3. 张宏明 陈珈 激发子与植物抗病信号转导 植物生理学通讯 1999, 221-226
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