环式电子传递在植物抗环境胁迫过程中的重要作用
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摘要
光是光合作用的驱动力,但是过剩的光能会导致光损伤。各种环境胁迫会降低光合作用效率,导致叶片吸收的光能大大超过光合作用消耗的光能,进而引起光损伤。植物叶片也进化出各种保护措施来应对过剩光能的存在,其中,环式电子传递是近几年来备受关注的光保护机制之一。但是,关于环式电子传递在抗环境胁迫过程中的重要作用还缺乏了解。本研究通过对热带植物环式电子传递对干旱、低温和强光条件的响应研究,试图阐述环式电子传递在植物适应环境胁迫过程中的重要作用。
零上低温是限制植物生长的一个重要环境因子,以前的研究发现冷敏感植物的光系统Ⅱ对零上低温胁迫不敏感,而光系统Ⅰ对零上低温胁迫敏感。我们的研究发现热带树木的光系统Ⅱ对零上低温十分敏感是导致热带树木难以在亚热带地区过冬的重要原因之一。零上低温导致光系统Ⅱ损伤的机制并不符合传统观念认为的自由基损伤假说,而是新的两步损伤假说：首先是放氧复合体的受损,然后是光系统Ⅱ反应中心的受损,放氧复合体的受损速率决定了光系统Ⅱ反应中心的受损速率。热带树木光系统Ⅰ和Ⅱ对零上低温的敏感程度与环式电子传递在零上低温下的激发程度有关。环式电子传递激发程度高的叶片对零上低温的敏感性较低。经过零上低温处理后,在常温下环式电子传递的激发有助于补充ATP的合成,加快光系统Ⅱ的光损伤的恢复。热带树木在热带北缘地区冬季的适应过程中光系统Ⅰ和Ⅱ活性都保持稳定,这个过程依赖于环式电子传递的激发,因为环式电子传递的激发一方面保护了光系统Ⅰ,另一方面保护了光系统Ⅱ。
干旱胁迫是影响植物生长的重要限制条件。大部分植物都不能适应极端干旱条件,但是复苏植物却能够忍受极度干旱,其中的光合生理机制并不清楚。我们的研究发现一种复苏植物蛛毛苣苔的叶片在失水条件下环式电子传递被强烈激发,一方面减缓其光系统Ⅱ受损的程度,另一方面保护了光系统Ⅰ。但是,长期失水的蛛毛苣苔叶片的光系统Ⅰ和Ⅱ活性都大幅度降低,当复水一天后,光系统Ⅰ和Ⅱ活性都完全恢复,这与模式植物拟南芥缓慢的光系统Ⅰ修复速度相悖。
环式电子传递保护光系统Ⅱ的机制主要有两个,一种是直接缓解光损伤的程度,另一种是防止光损伤的修复受到抑制。目前被研究的较透彻后一种机制,关于前一种机制的了解很少。我们的研究发现当环式电子传递被解除后,放氧复合体的损伤程度以及光系统Ⅱ反应中心的损伤程度都明显加剧,表明强光下环式电子传递的激发保护了放氧复合体。此外,放氧复合体受损的程度决定整个光系统Ⅱ发生损伤的程度,也更好的解释了环式电子传递如何直接缓解光系统Ⅱ的损伤。
叶片在强光下的主动合拢是一个十分有趣的生态现象,我们的研究发现在强光下主动合拢的叶片的环式电子传递和热耗散受强光激发的程度比不会主动合拢的叶片低,表明叶片合拢是为了弥补生理性光保护机制的不足。我们的研究从光保护的角度解释了叶片运动的重要意义。
通过上述四项实验,我们的研究一方面证实了环式电子传递在热带植物抵御低温、干旱和强光胁迫过程中发挥重要的作用,另一方面解释了环式电子传递保护光系统Ⅱ的另外一个重要机制是保护放氧复合体,为认识环式电子传递的重要性提供了新的视角。此外,还从光保护的角度解释了叶片运动的重要意义。
Light is the driving force of photosynthesis. However, excess light energy could induce photoinhibition. Environmental stresses induce the decrease in photosynthetic rate so that leads to the increase in excess light energy and then cause photoinhibition. Plants develop several mechanisms to protect leaves against excess light energy, such as cyclic electron flow (CEF), which has been widely studied in recent years. In order to explore the significant role of CEF in acclimation of plants to environmental stresses, the present study focuses on the response of CEF to chilling temperature, drought stress and high light. The results indicat that:1) The extreme sensitivity of photosystem Ⅱ (PSⅡ) to chilling-and-light stress is an important mechanism for why tropical plants cannot survive the winter in subtropical areas;2) In contrast to the traditional understanding of photoinhition of PSⅡ, the present study gives support to the new two-step scheme responsible for the photoinhibition of PSⅡ in tropical trees illuminated at chilling temperature;3) The extent of stimulation of CEF at chilling temperature determines the extent of photoinhibition of photosystem Ⅰ (PSI) and PSⅡ in tropical trees when illuminated at chilling temperature;4) After chilling-and-light treatment, CEF is stimulated under low light to help the fast recovery of PSⅡ activity at room temperature;5) Cyclic electron flow plays an important role during the acclimation of tropical trees to winter in marginal tropical areas;6) Cyclic electron flow plays an important role in photoprotection for resurrection plant under drought stress;7) Resurrection plant has the ability to rapidly repair PSI reaction centers, which contrasts to the low rate of recovery of PSI activity in non-resurrection plants;8) One role of CEF in alleviating PSⅡ photodamage is to protect oxygen-evolving complex, and the photodamage to oxygen-evolving complex is a limiting step of PSⅡ photoinhibitiom;9) An important mechanism of leaf folding is to remedy the deficiency of CEF and non-photochemical quenching. The present study indicates that CEF plays an impotant role in tropical trees under environmental stresses, such as chilling temperature, drought and high light. Furthermore, we found that a role of CEF in photoprotection for PSⅡ is stabilizing oxygen-evolving complex, which extends our understanding in the role of CEF.
零上低温是限制植物生长的一个重要环境因子,以前的研究发现冷敏感植物的光系统Ⅱ对零上低温胁迫不敏感,而光系统Ⅰ对零上低温胁迫敏感。我们的研究发现热带树木的光系统Ⅱ对零上低温十分敏感是导致热带树木难以在亚热带地区过冬的重要原因之一。零上低温导致光系统Ⅱ损伤的机制并不符合传统观念认为的自由基损伤假说,而是新的两步损伤假说：首先是放氧复合体的受损,然后是光系统Ⅱ反应中心的受损,放氧复合体的受损速率决定了光系统Ⅱ反应中心的受损速率。热带树木光系统Ⅰ和Ⅱ对零上低温的敏感程度与环式电子传递在零上低温下的激发程度有关。环式电子传递激发程度高的叶片对零上低温的敏感性较低。经过零上低温处理后,在常温下环式电子传递的激发有助于补充ATP的合成,加快光系统Ⅱ的光损伤的恢复。热带树木在热带北缘地区冬季的适应过程中光系统Ⅰ和Ⅱ活性都保持稳定,这个过程依赖于环式电子传递的激发,因为环式电子传递的激发一方面保护了光系统Ⅰ,另一方面保护了光系统Ⅱ。
干旱胁迫是影响植物生长的重要限制条件。大部分植物都不能适应极端干旱条件,但是复苏植物却能够忍受极度干旱,其中的光合生理机制并不清楚。我们的研究发现一种复苏植物蛛毛苣苔的叶片在失水条件下环式电子传递被强烈激发,一方面减缓其光系统Ⅱ受损的程度,另一方面保护了光系统Ⅰ。但是,长期失水的蛛毛苣苔叶片的光系统Ⅰ和Ⅱ活性都大幅度降低,当复水一天后,光系统Ⅰ和Ⅱ活性都完全恢复,这与模式植物拟南芥缓慢的光系统Ⅰ修复速度相悖。
环式电子传递保护光系统Ⅱ的机制主要有两个,一种是直接缓解光损伤的程度,另一种是防止光损伤的修复受到抑制。目前被研究的较透彻后一种机制,关于前一种机制的了解很少。我们的研究发现当环式电子传递被解除后,放氧复合体的损伤程度以及光系统Ⅱ反应中心的损伤程度都明显加剧,表明强光下环式电子传递的激发保护了放氧复合体。此外,放氧复合体受损的程度决定整个光系统Ⅱ发生损伤的程度,也更好的解释了环式电子传递如何直接缓解光系统Ⅱ的损伤。
叶片在强光下的主动合拢是一个十分有趣的生态现象,我们的研究发现在强光下主动合拢的叶片的环式电子传递和热耗散受强光激发的程度比不会主动合拢的叶片低,表明叶片合拢是为了弥补生理性光保护机制的不足。我们的研究从光保护的角度解释了叶片运动的重要意义。
通过上述四项实验,我们的研究一方面证实了环式电子传递在热带植物抵御低温、干旱和强光胁迫过程中发挥重要的作用,另一方面解释了环式电子传递保护光系统Ⅱ的另外一个重要机制是保护放氧复合体,为认识环式电子传递的重要性提供了新的视角。此外,还从光保护的角度解释了叶片运动的重要意义。
Light is the driving force of photosynthesis. However, excess light energy could induce photoinhibition. Environmental stresses induce the decrease in photosynthetic rate so that leads to the increase in excess light energy and then cause photoinhibition. Plants develop several mechanisms to protect leaves against excess light energy, such as cyclic electron flow (CEF), which has been widely studied in recent years. In order to explore the significant role of CEF in acclimation of plants to environmental stresses, the present study focuses on the response of CEF to chilling temperature, drought stress and high light. The results indicat that:1) The extreme sensitivity of photosystem Ⅱ (PSⅡ) to chilling-and-light stress is an important mechanism for why tropical plants cannot survive the winter in subtropical areas;2) In contrast to the traditional understanding of photoinhition of PSⅡ, the present study gives support to the new two-step scheme responsible for the photoinhibition of PSⅡ in tropical trees illuminated at chilling temperature;3) The extent of stimulation of CEF at chilling temperature determines the extent of photoinhibition of photosystem Ⅰ (PSI) and PSⅡ in tropical trees when illuminated at chilling temperature;4) After chilling-and-light treatment, CEF is stimulated under low light to help the fast recovery of PSⅡ activity at room temperature;5) Cyclic electron flow plays an important role during the acclimation of tropical trees to winter in marginal tropical areas;6) Cyclic electron flow plays an important role in photoprotection for resurrection plant under drought stress;7) Resurrection plant has the ability to rapidly repair PSI reaction centers, which contrasts to the low rate of recovery of PSI activity in non-resurrection plants;8) One role of CEF in alleviating PSⅡ photodamage is to protect oxygen-evolving complex, and the photodamage to oxygen-evolving complex is a limiting step of PSⅡ photoinhibitiom;9) An important mechanism of leaf folding is to remedy the deficiency of CEF and non-photochemical quenching. The present study indicates that CEF plays an impotant role in tropical trees under environmental stresses, such as chilling temperature, drought and high light. Furthermore, we found that a role of CEF in photoprotection for PSⅡ is stabilizing oxygen-evolving complex, which extends our understanding in the role of CEF.
引文
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