Department of Biophysics, Biology Faculty of the M.V. Lomonosov, Moscow State University, Moscow, Russia, 119234.
Photosynth Res. 2019 Apr;140(1):1-19. doi: 10.1007/s11120-019-00627-8. Epub 2019 Feb 27.
The dark-to-light transitions enable energization of the thylakoid membrane (TM), which is reflected in fast and slow (OJIPSMT or OABCDE) stages of fluorescence induction (FI) and P700 oxidoreduction changes (ΔA). A Thylakoid Membrane model (T-M model), in which special emphasis has been placed on ferredoxin-NADP-oxidoreductase (FNR) activation and energy-dependent qE quenching, was applied for quantifying the kinetics of FI and ΔA. Pea leaves were kept in darkness for 15 min and then the FI and ΔA signals were measured upon actinic illumination, applied either directly or after a 10-s light pulse coupled with a subsequent 10-s dark interval. On the time scale from 40 µs to 30 s, the parallel T-M model fittings to both FI and ΔA signals were obtained. The parameters of FNR activation and the buildup of qE quenching were found to differ for dark-adapted and preilluminated leaves. At the onset of actinic light, photosystem II (PSII) acceptors were oxidized (neutral) after dark adaptation, while the redox states with closed and/or semiquinone QQ forms were supposedly generated after preillumination, and did not relax within the 10 s dark interval. In qE simulations, a pH-dependent Hill relationship was used. The rate constant of heat losses in PSII antenna k(t) was found to increase from the basic value k, at the onset of illumination, to its maximal level k due to lumenal acidification. In dark-adapted leaves, a low value of k of ∼ 2 × 10 s was found. Simulations on the microsecond to 30 s time scale revealed that the slow P-S-M-T phases of the fluorescence induction were sensitive to light-induced FNR activation and high-energy qE quenching. Thus, the corresponding time-dependent rate constants k(t) and k(t) change substantially upon the release of electron transport on the acceptor side of PSI and during the NPQ development. The transitions between the cyclic and linear electron transport modes have also been quantified in this paper.
从黑暗到光明的转变使类囊体膜(TM)被激活,这反映在荧光诱导(FI)的快速和慢速(OJIPSMT 或 OABCDE)阶段以及 P700 氧化还原变化(ΔA)中。一个特别强调铁氧还蛋白-NADP-氧化还原酶(FNR)激活和能量依赖的 qE 猝灭的类囊体膜模型(T-M 模型)被应用于量化 FI 和 ΔA 的动力学。豌豆叶片在黑暗中放置 15 分钟,然后在光照射下测量 FI 和 ΔA 信号,直接施加光或在 10 秒光脉冲后施加 10 秒暗间隔。在 40 μs 到 30 s 的时间尺度上,获得了对 FI 和 ΔA 信号的并行 T-M 模型拟合。发现暗适应和预照光叶片的 FNR 激活和 qE 猝灭的建立参数不同。在光照射开始时,PSII 受体在暗适应后被氧化(中性),而在预照光后,应该产生了具有闭合和/或半醌 QQ 形式的氧化还原状态,并且在 10 秒的暗间隔内不会松弛。在 qE 模拟中,使用了 pH 依赖的希尔关系。在光照射开始时,PSII 天线的热损失速率常数 k(t) 从基本值 k 增加到由于腔室酸化而达到最大水平 k。在暗适应叶片中,发现 k 的低值约为 2×10 s。在微秒到 30 s 的时间尺度上的模拟表明,荧光诱导的慢 P-S-M-T 相对光诱导的 FNR 激活和高能 qE 猝灭敏感。因此,在 PSI 受体侧电子传递释放和 NPQ 发展过程中,相应的时变速率常数 k(t) 和 k(t) 会发生很大变化。本文还量化了循环和线性电子传递模式之间的转变。