植物对斑潜蝇防御的多层次研究
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摘要
植物与植食性昆虫长期的相互作用中形成了多种防御机制。健康状态下,植物存在组成型防御;在受到病虫害或者环境中其他因素胁迫时,植物会产生诱导性防御。诱导性防御不仅能在被损伤的叶片处发生(局部防御),也可以在其它部位产生(系统防御)。组成型防御和诱导性防御反映了植物防御时间分布上的多样性,而局部防御和系统防御则反映了植物防御空间分布上的多样性。不仅如此,这些防御机制又主要以多种方式对植食性昆虫产生影响:⑴植物利用本身含有的、能够对植食性昆虫产生不利影响的物质进行防御(直接防御);⑵植物在受到伤害后与周边的健康植物进行“交流”,健康植物在收到报警信号后表达抗性物质,共同进行防御(植物-植物相互交流);⑶植物通过释放挥发物来吸引植食性昆虫的天敌,从而达到防御植食性昆虫的目的(间接防御)。
本文正是利用实验室建立的“植物-斑潜蝇-寄生蜂”三级营养系统为模型,针对植物防御中的关键问题,从以下四个方面进行了研究。
1.植物与植食性昆虫相互作用——拟南芥对斑潜蝇的防御策略
潜食性昆虫生活在植物叶片内部,具有特殊的取食方式,因此对植物的伤害与咀嚼式昆虫或刺吸式昆虫有很大的区别。本文利用affymetrix拟南芥全基因组芯片检测拟南芥对斑潜蝇危害后的防御反应,结果发现斑潜蝇危害引起的防御反应与细菌相似,却和其他取食方式的昆虫相差很大,这些结果表明植物防御反应与昆虫的危害模式有关。
2.植物-自身防御机制——利马豆新叶与老叶系统防御能力的差异
植物的不同部位往往具有不同的生物学特征,并且新叶更易受到昆虫的危害。那么,植物的不同部位,尤其是新叶与老叶的系统防御能力是否存在差异?我们利用定量PCR和挥发物鉴定技术检测了利马豆新叶与老叶的系统防御能力;结果表明,利马豆新叶的系统防御能力强于老叶,这也说明植物在受到危害后自身存在着不同的防御能力,加强防御受保护的部位。
3.植物-植物共同防御机制——“植物-植物相互交流”的系统性研究
“植物-植物相互交流”是近三十年前才被发现,此后就成为了一个热门的话题。但是大多数研究都集中在了现象的确认上,对机理的认识还很少。本文对这个问题进行了系统性的研究。首先,我们确认了“演讲者”和“听众”之间的交流和它们的亲缘关系无关,而主要取决于“演讲者”挥发物的性质;随后,我们分析了“演讲者”释放的不同种类的挥发物对“听众”的效果,以及作用时间和浓度的影响;然后,我们利用全基因组芯片对对“听众”的反应进行了全面的检测,详细分析其信号通路和差异基因功能;最后,我们利用突变体植物证明了乙烯(ET)信号通路是“演讲者“和”“听众”植物相互交流所不可或缺的。
4.植物-天敌间接防御机制——“利马豆-斑潜蝇-寄生蜂”三级营养系统活动节律的相互关系
节律是生命的基本特征,它可以由体内生物钟控制,也可以受到外界信号的影响。但是不同生物的活动节律之间是否存在相互关系还知之甚少。我们利用“利马豆-斑潜蝇-寄生蜂”这样一个具有密切关系的系统来研究这个问题。结果发现,利马豆、斑潜蝇、寄生蜂的活动节律具有强烈的同步性,同时这种同步性会受到光照的影响,是三种生物在自然界正常光周期下共同进化的结果。不仅如此,没有产过卵的处女寄生蜂和具有产卵经验的寄生蜂还对不同时间段的挥发物具有不同的敏感性,从而在整个生命周期中能够利用植物挥发物的节律信息来定位寄主昆虫。
以上这四方面研究以植物为主体,分别针对植物防御中的四个关键问题⑴植物与植食性昆虫相互作用的复杂性⑵植物自身的系统防御机制⑶植物-植物共同防御机制⑷植物-天敌间接防御机制进行了深入的探讨,以期能够加深人们对植物防御机制的认识,为害虫防治提供理论基础以及方法参考。
During the long time interactions with insects, plants have developed a sophisticated defense system. Healthy plants usually utilize constitutive defenses which always exist in plants, while they will express the inducible defense which reacts efficiently after being damaged by attackers. These two kinds of plant defense reflect the diversity of time course. Induced defense can be express at the attacked leaves (local defense) or undamaged leaves (systemic defense) of the plants; and they reflect the diversity of defense position. Not only that, all these defenses can be achieved by multiple pathways: Firstly, plants can defense the insects with noxious materials always existing in the plant directly (direct defense); secondly, damaged plants can also“communicate”with their healthy neighbors, and the healthy neighbors will prepare for damage and defense insects together (plant-plant communication); Thirdly, there are many enemies of herbivory insects in the nature, plants can attract the natural enemies that may help defense the insects (indirect defense).
We used the“plant-leafminer-parasitoid”tritrophic systems to study 4 key points about plant defenses;
1. Interactions between plants and insects——Transcription defense responses of Arabidopsis thaliana to leafminer damage.
Leafminers, a kind of special insects which live inside the plants, have a different damage patterns from other insects. We use an affimetrix ATH1 arabisopsis microarray to test the transcription response pattern of Arabidopsis thaliana to pea leafminers damages, both locally infected leaves (LI) and systemically infected leaves (SI) were tested. The data proved that Arabidopsis thaliana response to pea leafminer was more similar with that of pathogens than other insects. The results support our hypothesis that the plant response patterns were greatly related to the attacker’s damages patterns.
2. Systemic defenses of new secondary leaves are stranger than old primary leaves in lima bean (Phaseolus lunatus).
New secondary leaves often have different biological characters from the old primary leaves, and the new leaves are more vulnerability to herbivores. Whether the systemic defense abilities in secondary and primary leaves are different? Our researches showed that the new secondary leaves had stronger systemic defense ability then the old primary leaves, both in direct defense and in indirect defense. Thus, diverse parts plant has diverse defense abilities; the plant prefers to protect the new leaves when attacked.
3. Systemic researches of“plant-plant communication”.
“Plant-plant communication”is a new and hot topic. Many studies have focused on this interesting topic. However, the mechanism of this phenomenon was not clearly known. Firstly, we demonstrated that communication of plants was nonrelated to the phylogenetic relationship of different species, while the character of volatiles was a crucial factor; secondly, the VOCs that emitters released is the“words”between plants, and different kind of volatiles have different characters; thirdly, we analyzed the response of Arabidopsis thaliana signal pathway in detail; finally, Arabidopsis thaliana mutant was used to disclose that ethylene response signal of plant was indispensable in“plant-plant communication”. Our results gave a whole and clear picture of plant-plant communication between lima bean and Arabidopsis thaliana.
4. Rhythm interactions in tritrophic system.
Diurnal rhythm is a basic character of organisms, which can be controlled by inherent circadian clock or affected by other factors. However, whether the rhythms of different organisms can interact with each other is poorly known. Here, we used the“lima bean-leafminer-parasitoid”tritrophic systems to answer this question. The results showed that the rhythms in the tritrophic system were synchronized with each other, but this can be disturbed by dark, which indicated that the rhythm interactions in the tritrophic system was the result of co-evolution under natural light-dark cycle. We also found that naive and oviposition-experienced parasitoids were able to respectively utilize the volatile rhythm information from the damaged plant to locate host leafminers.
We hope that our results can help us understand more about plant defense phenomenon and mechanisms, which may guide pest controlling in the agriculture.
本文正是利用实验室建立的“植物-斑潜蝇-寄生蜂”三级营养系统为模型,针对植物防御中的关键问题,从以下四个方面进行了研究。
1.植物与植食性昆虫相互作用——拟南芥对斑潜蝇的防御策略
潜食性昆虫生活在植物叶片内部,具有特殊的取食方式,因此对植物的伤害与咀嚼式昆虫或刺吸式昆虫有很大的区别。本文利用affymetrix拟南芥全基因组芯片检测拟南芥对斑潜蝇危害后的防御反应,结果发现斑潜蝇危害引起的防御反应与细菌相似,却和其他取食方式的昆虫相差很大,这些结果表明植物防御反应与昆虫的危害模式有关。
2.植物-自身防御机制——利马豆新叶与老叶系统防御能力的差异
植物的不同部位往往具有不同的生物学特征,并且新叶更易受到昆虫的危害。那么,植物的不同部位,尤其是新叶与老叶的系统防御能力是否存在差异?我们利用定量PCR和挥发物鉴定技术检测了利马豆新叶与老叶的系统防御能力;结果表明,利马豆新叶的系统防御能力强于老叶,这也说明植物在受到危害后自身存在着不同的防御能力,加强防御受保护的部位。
3.植物-植物共同防御机制——“植物-植物相互交流”的系统性研究
“植物-植物相互交流”是近三十年前才被发现,此后就成为了一个热门的话题。但是大多数研究都集中在了现象的确认上,对机理的认识还很少。本文对这个问题进行了系统性的研究。首先,我们确认了“演讲者”和“听众”之间的交流和它们的亲缘关系无关,而主要取决于“演讲者”挥发物的性质;随后,我们分析了“演讲者”释放的不同种类的挥发物对“听众”的效果,以及作用时间和浓度的影响;然后,我们利用全基因组芯片对对“听众”的反应进行了全面的检测,详细分析其信号通路和差异基因功能;最后,我们利用突变体植物证明了乙烯(ET)信号通路是“演讲者“和”“听众”植物相互交流所不可或缺的。
4.植物-天敌间接防御机制——“利马豆-斑潜蝇-寄生蜂”三级营养系统活动节律的相互关系
节律是生命的基本特征,它可以由体内生物钟控制,也可以受到外界信号的影响。但是不同生物的活动节律之间是否存在相互关系还知之甚少。我们利用“利马豆-斑潜蝇-寄生蜂”这样一个具有密切关系的系统来研究这个问题。结果发现,利马豆、斑潜蝇、寄生蜂的活动节律具有强烈的同步性,同时这种同步性会受到光照的影响,是三种生物在自然界正常光周期下共同进化的结果。不仅如此,没有产过卵的处女寄生蜂和具有产卵经验的寄生蜂还对不同时间段的挥发物具有不同的敏感性,从而在整个生命周期中能够利用植物挥发物的节律信息来定位寄主昆虫。
以上这四方面研究以植物为主体,分别针对植物防御中的四个关键问题⑴植物与植食性昆虫相互作用的复杂性⑵植物自身的系统防御机制⑶植物-植物共同防御机制⑷植物-天敌间接防御机制进行了深入的探讨,以期能够加深人们对植物防御机制的认识,为害虫防治提供理论基础以及方法参考。
During the long time interactions with insects, plants have developed a sophisticated defense system. Healthy plants usually utilize constitutive defenses which always exist in plants, while they will express the inducible defense which reacts efficiently after being damaged by attackers. These two kinds of plant defense reflect the diversity of time course. Induced defense can be express at the attacked leaves (local defense) or undamaged leaves (systemic defense) of the plants; and they reflect the diversity of defense position. Not only that, all these defenses can be achieved by multiple pathways: Firstly, plants can defense the insects with noxious materials always existing in the plant directly (direct defense); secondly, damaged plants can also“communicate”with their healthy neighbors, and the healthy neighbors will prepare for damage and defense insects together (plant-plant communication); Thirdly, there are many enemies of herbivory insects in the nature, plants can attract the natural enemies that may help defense the insects (indirect defense).
We used the“plant-leafminer-parasitoid”tritrophic systems to study 4 key points about plant defenses;
1. Interactions between plants and insects——Transcription defense responses of Arabidopsis thaliana to leafminer damage.
Leafminers, a kind of special insects which live inside the plants, have a different damage patterns from other insects. We use an affimetrix ATH1 arabisopsis microarray to test the transcription response pattern of Arabidopsis thaliana to pea leafminers damages, both locally infected leaves (LI) and systemically infected leaves (SI) were tested. The data proved that Arabidopsis thaliana response to pea leafminer was more similar with that of pathogens than other insects. The results support our hypothesis that the plant response patterns were greatly related to the attacker’s damages patterns.
2. Systemic defenses of new secondary leaves are stranger than old primary leaves in lima bean (Phaseolus lunatus).
New secondary leaves often have different biological characters from the old primary leaves, and the new leaves are more vulnerability to herbivores. Whether the systemic defense abilities in secondary and primary leaves are different? Our researches showed that the new secondary leaves had stronger systemic defense ability then the old primary leaves, both in direct defense and in indirect defense. Thus, diverse parts plant has diverse defense abilities; the plant prefers to protect the new leaves when attacked.
3. Systemic researches of“plant-plant communication”.
“Plant-plant communication”is a new and hot topic. Many studies have focused on this interesting topic. However, the mechanism of this phenomenon was not clearly known. Firstly, we demonstrated that communication of plants was nonrelated to the phylogenetic relationship of different species, while the character of volatiles was a crucial factor; secondly, the VOCs that emitters released is the“words”between plants, and different kind of volatiles have different characters; thirdly, we analyzed the response of Arabidopsis thaliana signal pathway in detail; finally, Arabidopsis thaliana mutant was used to disclose that ethylene response signal of plant was indispensable in“plant-plant communication”. Our results gave a whole and clear picture of plant-plant communication between lima bean and Arabidopsis thaliana.
4. Rhythm interactions in tritrophic system.
Diurnal rhythm is a basic character of organisms, which can be controlled by inherent circadian clock or affected by other factors. However, whether the rhythms of different organisms can interact with each other is poorly known. Here, we used the“lima bean-leafminer-parasitoid”tritrophic systems to answer this question. The results showed that the rhythms in the tritrophic system were synchronized with each other, but this can be disturbed by dark, which indicated that the rhythm interactions in the tritrophic system was the result of co-evolution under natural light-dark cycle. We also found that naive and oviposition-experienced parasitoids were able to respectively utilize the volatile rhythm information from the damaged plant to locate host leafminers.
We hope that our results can help us understand more about plant defense phenomenon and mechanisms, which may guide pest controlling in the agriculture.
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