快速葉綠素熒光(OJIP)可作為監(jiān)測植物在非生物脅迫下光合生理狀態(tài)的有效工具
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摘要
在自然條件下生活的植物會受到許多干擾光合作用過程的不利因素的影響，導致生長、發(fā)育和產(chǎn)量的下降。葉綠素a熒光光譜(ChlF)的研究為葉片光化學效率研究提供了一條新的途徑。具體地說，對熒光信號的分析可獲取PSII反應(yīng)中心、捕光天線復合體以及PSII供體側(cè)/受體側(cè)的狀態(tài)和功能的詳細信息。在這里，我們回顧了快速ChlF技術(shù)(OJIP & JIP-test)分析光合反應(yīng)對環(huán)境脅迫的相關(guān)成果，并討論了這一創(chuàng)新方法的潛在科學和實際應(yīng)用。最近便攜式設(shè)備(Handy PEA & M-PEA, Hansatech Instruments)的出現(xiàn)，特別是在作物表型分型和監(jiān)測方面，大大擴展了ChlF技術(shù)的潛在應(yīng)用。
關(guān)鍵詞 Chlorophyll fluorescence、JIP-test、Photosynthesis、Photosystem II、Quantum efficiency、Stress detection
縮寫
Absorption flux | 吸收通量 | |
Chlorophyll | 葉綠素 | |
Chlorophyll fluorescence | 葉綠素熒光 | |
Cross section of the sample | 樣品橫截面 | |
Cytochrome b6f | 細胞色素b6f | |
Delayed (chlorophyll) fluorescence | 延遲(葉綠素)熒光 | |
Drought factor index | 干旱因子指數(shù) | |
Light-harvesting complex (of PSII) | PSII捕光色素復合體 | |
Oxygen-evolving complex | 放氧復合體 | |
Excited PSII reaction center | 激發(fā)的PSII反應(yīng)中心 | |
PSI reaction center | PSI反應(yīng)中心 | |
Photosynthetically active radiation | 光合有效輻射 | |
Plastocyanin | 質(zhì)體藍素 | |
Principal component analysis | 主成分分析 | |
Prompt (chlorophyll) fluorescence | 瞬時(葉綠素)熒光 | |
Pheophytin | 去鎂葉綠素 | |
Plastoquinone | 質(zhì)體醌 | |
Photosystem I, II | 光系統(tǒng)I, II | |
Primary plastoquinone electron acceptor of PSII | PSII初級質(zhì)體醌電子受體 | |
Secondary plastoquinone electron acceptor | 次級質(zhì)體醌電子受體 | |
Reaction center | 反應(yīng)中心 | |
Reactive oxygen species | 活性氧 |
在自然條件下，植物受到許多不利的環(huán)境脅迫因子的影響。這些會破壞光合器官，導致植物生產(chǎn)力和總產(chǎn)量下降。光合作用對環(huán)境脅迫特別敏感(Kalaji et al. 2012)，使光合測量成為植物脅迫研究的重要組成部分。然而，傳統(tǒng)的方法，甚至是技術(shù)上先進的方法，如通過氣體交換(CO2、H2O和O2)測量光合速率，需要耗費大量時間和人力，且提供的有關(guān)整體光合功能的信息并不完整。相比之下，ChlF測量是一種簡單、無損、廉價和快速的工具，可用于分析光依賴性光合反應(yīng)和間接評估同一樣本組織中的葉綠素含量(Govindjee 1995; Papageorgiou & Govindjee 2011; Stirbet & Govindjee 2011, 2012)。ChlF方法的這些技術(shù)優(yōu)勢使其成為植物育種家(例如作物表型和監(jiān)測)、生物技術(shù)學家、植物生理學家、林業(yè)工作者、生態(tài)學家和環(huán)境學家的流行技術(shù)。
關(guān)鍵的是，從植物脅迫研究的角度來看，ChlF測量還提供了有關(guān)植物生理狀況的間接信息。通過分析葉綠素熒光(ChlF)誘導曲線，可以評估光系統(tǒng)II(PSII)和光合電子傳遞鏈的生理狀況。它還提供了光依賴的光化學反應(yīng)和光無關(guān)的生化反應(yīng)的相關(guān)信息�？偟膩碚f，ChlF測量直接或間接地與依賴光的光合反應(yīng)的所有階段有關(guān)，包括水的光解、電子傳遞、類囊體膜上pH梯度的形成、ATP合成以及光合機構(gòu)的一般生物能條件等(Bernát et al. 2012)。
暗適應(yīng)葉片照光后可獲得多相葉綠素熒光誘導曲線(O–J–I–P-瞬變)(圖1)。曲線的軌跡提供了有關(guān)光合機構(gòu)結(jié)構(gòu)和功能的大量信息(Kautsky & Hirsch 1931; Schreiber et al. 1994)。
JIP-test是基于多相快速葉綠素熒光的上升階段，用于研究光依賴性反應(yīng)與ChlF的相關(guān)性。它基于類囊體膜的“能量流”理論(Strasser et al. 2000)。這個理論可以用簡單的代數(shù)方程來計算，代表每一個被檢測的捕光復合體的總能量流入和流出之間的平衡，并提供關(guān)于吸收能量的可能分配的信息。利用這些方程，可以描述PSII復合體之間的能量通信(也稱為“聚集grouping”或“連通性connectivity”和“總體分組概率overall grouping probability”)(Stirbet 2013)。
JIP-test(OJIP)的名稱來源于ChlF信號形成的感應(yīng)曲線上的特定位點(圖1)：這些位點對應(yīng)于PSII原初電子受體(Pheo)和QA的逐漸還原。誘導曲線的形狀取決于PSII各組分間的聚集性(L-band)(Tsimilli-Michaeland Strasser 2013)和電子供體OEC→P680+以及QA-電子的接收之間的平衡(K-band)(Strasser et al. 2005)。
O~J相的熒光上升階段與部分PSII反應(yīng)中心的閉合相關(guān)，反應(yīng)了QA的還原水平，其還原程度取決于捕獲速率以及QA被QB和其余電子傳遞鏈成員氧化的速率。
誘導曲線的J~I相與次級電子受體QB、PQ、Cyt b6f和PC的還原程度相關(guān)。誘導曲線的I~P相的上升通常歸因于PSI受體側(cè)電子受體(鐵氧還原蛋白、中間受體和NADP)的還原。
圖1：典型的植物葉綠素熒光多相動力學曲線(主圖)，曲線以對數(shù)時間刻度(10μs~600s)繪制。左上部插圖顯示了按常規(guī)時間標度繪制的相同曲線。右下方插圖按常規(guī)時間標度繪制的OJIP瞬態(tài)(0-30ms)的初始部分。時間標記是指JIP-test用于計算結(jié)構(gòu)和功能參數(shù)的選定時間點。
圖2：不同脅迫條件下小麥(Triticum sp.L.)葉綠素熒光的O(K)JIP瞬態(tài)與非脅迫下的比較。插入顯示了O-J相(∆VOJ)、J-I相(∆VJI)、I-P相(∆VIP)的相對可變熒光振幅的變化，以及0.3 ms可變熒光(VK/VJ)與2ms可變熒光比值(VJ)的變化，作為PSII供體側(cè)限制(K-band)的指標。各圖顯示了相對于非脅迫狀態(tài)下植物(control，C)的瞬時熒光曲線：a熱脅迫(高溫脅迫8h，中度光化光照射，葉片溫度約40℃)；b低溫脅迫10d(10/6℃：日間/夜間)；c重度干旱脅迫(停止灌溉后12d，葉片含水量約60%)；d鹽脅迫(NaCl)；e氮缺乏脅迫(低氮，LN)；f鉛脅迫。
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