Bukhov N, Egorova E, Krendeleva T, Rubin A, Wiese C, Heber U
Timiriazev Institute of Plant Physiology Russian Academy of Science, Botanicheskaya 35, 127276, Moscow, Russia,
Photosynth Res. 2001;70(2):155-66. doi: 10.1023/A:1017950307360.
Kinetics of the dark relaxation of variable chlorophyll fluorescence, Fv, were studied after brief illumination of dark-adapted barley leaves in order to understand the rapid reversibility of pulse-induced fluorescence increases, which is observed even when fast linear electron transport to an external electron acceptor is not possible. Four kinetically distinct components were observed which reveal complexity in the oxidation of the reduced primary quinone acceptor of Photosystem II, Q (A) (-) : the slowest component accounted for 4-5% of maximal Fv and had a life-time of several seconds. It is suggested to represent a minor population of inactive Photosystem II centers. The other three components displayed first-order kinetics with half-time of 6-8 ms ('fast' component), 60-80 ms ('middle' component) and 650-680 ms ('slow' component). The fast component dominated Fv when methyl viologen or far-red light accelerated oxidation of plastohydroquinone. It shows rapid oxidation of Q (A) (-) during electron flow to plastoquinone commensurate with maximum linear electron flow through the electron transport chain. The other two components were observed under conditions of restricted electron flow and excessive reduction of electron carriers. Unexpectedly, the slow component, which is interpreted to reflect the recombination between Q (A) (-) and an intermediate on the oxidizing side of Photosystem II, saturated already at low irradiances of actinic light when plastoquinone was not yet strongly reduced suggesting that dark-adaptation of leaves results not only in the loss of activity of light-regulated enzymes of the carbon cycle but affects also electron flow from Q(A) (-) to plastoquinone. KCN poisoning or high temperature treatment of leaves produced a nonexponential pattern of slow Fv relaxation. This effect was largely (heat treatment) or even completely (KCN) abolished by far-red light.
为了理解脉冲诱导的荧光增加的快速可逆性,对暗适应的大麦叶片进行短暂光照后,研究了可变叶绿素荧光Fv的暗弛豫动力学,即使在无法快速线性电子传递到外部电子受体时也能观察到这种可逆性。观察到四个动力学上不同的组分,这揭示了光系统II的还原初级醌受体Q(A)(-)氧化过程的复杂性:最慢的组分占最大Fv的4 - 5%,寿命为几秒。它被认为代表少量无活性的光系统II中心。其他三个组分呈现一级动力学,半衰期分别为6 - 8毫秒(“快速”组分)、60 - 80毫秒(“中间”组分)和650 - 680毫秒(“慢速”组分)。当甲基紫精或远红光加速质体氢醌氧化时,快速组分主导Fv。它显示在电子流向质体醌的过程中Q(A)(-)的快速氧化,与通过电子传递链的最大线性电子流相当。另外两个组分是在电子流受限和电子载体过度还原的条件下观察到的。出乎意料的是,慢速组分被解释为反映Q(A)(-)与光系统II氧化侧中间体之间的复合,当质体醌尚未被强烈还原时,在低光强的光化光照射下就已饱和,这表明叶片的暗适应不仅导致碳循环中光调节酶活性的丧失,还影响从Q(A)(-)到质体醌的电子流。KCN中毒或叶片高温处理产生了Fv慢速弛豫的非指数模式。这种效应在很大程度上(热处理)甚至完全(KCN)被远红光消除。