快速叶绿素荧光(OJIP)可作为监测植物在非生物胁迫下光合生理状态的有效工具
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摘要
在自然条件下生活的植物会受到许多干扰光合作用过程的不利因素的影响,导致生长、发育和产量的下降。叶绿素a荧光光谱(ChlF)的研究为叶片光化学效率研究提供了一条新的途径。具体地说,对荧光信号的分析可获取PSII反应中心、捕光天线复合体以及PSII供体侧/受体侧的状态和功能的详细信息。在这里,我们回顾了快速ChlF技术(OJIP & JIP-test)分析光合反应对环境胁迫的相关成果,并讨论了这一创新方法的潜在科学和实际应用。最近便携式设备(Handy PEA & M-PEA, Hansatech Instruments)的出现,特别是在作物表型分型和监测方面,大大扩展了ChlF技术的潜在应用。
关键词 Chlorophyll fluorescence、JIP-test、Photosynthesis、Photosystem II、Quantum efficiency、Stress detection
缩写
Absorption flux | 吸收通量 | |
Chlorophyll | 叶绿素 | |
Chlorophyll fluorescence | 叶绿素荧光 | |
Cross section of the sample | 样品横截面 | |
Cytochrome b6f | 细胞色素b6f | |
Delayed (chlorophyll) fluorescence | 延迟(叶绿素)荧光 | |
Drought factor index | 干旱因子指数 | |
Light-harvesting complex (of PSII) | PSII捕光色素复合体 | |
Oxygen-evolving complex | 放氧复合体 | |
Excited PSII reaction center | 激发的PSII反应中心 | |
PSI reaction center | PSI反应中心 | |
Photosynthetically active radiation | 光合有效辐射 | |
Plastocyanin | 质体蓝素 | |
Principal component analysis | 主成分分析 | |
Prompt (chlorophyll) fluorescence | 瞬时(叶绿素)荧光 | |
Pheophytin | 去镁叶绿素 | |
Plastoquinone | 质体醌 | |
Photosystem I, II | 光系统I, II | |
Primary plastoquinone electron acceptor of PSII | PSII初级质体醌电子受体 | |
Secondary plastoquinone electron acceptor | 次级质体醌电子受体 | |
Reaction center | 反应中心 | |
Reactive oxygen species | 活性氧 |
在自然条件下,植物受到许多不利的环境胁迫因子的影响。这些会破坏光合器官,导致植物生产力和总产量下降。光合作用对环境胁迫特别敏感(Kalaji et al. 2012),使光合测量成为植物胁迫研究的重要组成部分。然而,传统的方法,甚至是技术上先进的方法,如通过气体交换(CO2、H2O和O2)测量光合速率,需要耗费大量时间和人力,且提供的有关整体光合功能的信息并不完整。相比之下,ChlF测量是一种简单、无损、廉价和快速的工具,可用于分析光依赖性光合反应和间接评估同一样本组织中的叶绿素含量(Govindjee 1995; Papageorgiou & Govindjee 2011; Stirbet & Govindjee 2011, 2012)。ChlF方法的这些技术优势使其成为植物育种家(例如作物表型和监测)、生物技术学家、植物生理学家、林业工作者、生态学家和环境学家的流行技术。
关键的是,从植物胁迫研究的角度来看,ChlF测量还提供了有关植物生理状况的间接信息。通过分析叶绿素荧光(ChlF)诱导曲线,可以评估光系统II(PSII)和光合电子传递链的生理状况。它还提供了光依赖的光化学反应和光无关的生化反应的相关信息。总的来说,ChlF测量直接或间接地与依赖光的光合反应的所有阶段有关,包括水的光解、电子传递、类囊体膜上pH梯度的形成、ATP合成以及光合机构的一般生物能条件等(Bernát et al. 2012)。
暗适应叶片照光后可获得多相叶绿素荧光诱导曲线(O–J–I–P-瞬变)(图1)。曲线的轨迹提供了有关光合机构结构和功能的大量信息(Kautsky & Hirsch 1931; Schreiber et al. 1994)。
JIP-test是基于多相快速叶绿素荧光的上升阶段,用于研究光依赖性反应与ChlF的相关性。它基于类囊体膜的“能量流”理论(Strasser et al. 2000)。这个理论可以用简单的代数方程来计算,代表每一个被检测的捕光复合体的总能量流入和流出之间的平衡,并提供关于吸收能量的可能分配的信息。利用这些方程,可以描述PSII复合体之间的能量通信(也称为“聚集grouping”或“连通性connectivity”和“总体分组概率overall grouping probability”)(Stirbet 2013)。
JIP-test(OJIP)的名称来源于ChlF信号形成的感应曲线上的特定位点(图1):这些位点对应于PSII原初电子受体(Pheo)和QA的逐渐还原。诱导曲线的形状取决于PSII各组分间的聚集性(L-band)(Tsimilli-Michaeland Strasser 2013)和电子供体OEC→P680+以及QA-电子的接收之间的平衡(K-band)(Strasser et al. 2005)。
O~J相的荧光上升阶段与部分PSII反应中心的闭合相关,反应了QA的还原水平,其还原程度取决于捕获速率以及QA被QB和其余电子传递链成员氧化的速率。
诱导曲线的J~I相与次级电子受体QB、PQ、Cyt b6f和PC的还原程度相关。诱导曲线的I~P相的上升通常归因于PSI受体侧电子受体(铁氧还原蛋白、中间受体和NADP)的还原。
图1:典型的植物叶绿素荧光多相动力学曲线(主图),曲线以对数时间刻度(10μs~600s)绘制。左上部插图显示了按常规时间标度绘制的相同曲线。右下方插图按常规时间标度绘制的OJIP瞬态(0-30ms)的初始部分。时间标记是指JIP-test用于计算结构和功能参数的选定时间点。
图2:不同胁迫条件下小麦(Triticum sp.L.)叶绿素荧光的O(K)JIP瞬态与非胁迫下的比较。插入显示了O-J相(∆VOJ)、J-I相(∆VJI)、I-P相(∆VIP)的相对可变荧光振幅的变化,以及0.3 ms可变荧光(VK/VJ)与2ms可变荧光比值(VJ)的变化,作为PSII供体侧限制(K-band)的指标。各图显示了相对于非胁迫状态下植物(control,C)的瞬时荧光曲线:a热胁迫(高温胁迫8h,中度光化光照射,叶片温度约40℃);b低温胁迫10d(10/6℃:日间/夜间);c重度干旱胁迫(停止灌溉后12d,叶片含水量约60%);d盐胁迫(NaCl);e氮缺乏胁迫(低氮,LN);f铅胁迫。
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