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光系统 II 电子传递动力学:基于叶绿素荧光诱导的数学分析。

Kinetics of photosystem II electron transport: a mathematical analysis based on chlorophyll fluorescence induction.

机构信息

Institute of Technology, University of Tartu, Nooruse St. 1, Tartu, 50411, Estonia.

出版信息

Photosynth Res. 2018 Apr;136(1):63-82. doi: 10.1007/s11120-017-0439-y. Epub 2017 Sep 21.

Abstract

The OJDIP rise in chlorophyll fluorescence during induction at different light intensities was mathematically modeled using 24 master equations describing electron transport through photosystem II (PSII) plus ordinary differential equations for electron budgets in plastoquinone, cytochrome f, plastocyanin, photosystem I, and ferredoxin. A novel feature of the model is consideration of electron in- and outflow budgets resulting in changes in redox states of Tyrosine Z, P680, and Q as sole bases for changes in fluorescence yield during the transient. Ad hoc contributions by transmembrane electric fields, protein conformational changes, or other putative quenching species were unnecessary to account for primary features of the phenomenon, except a peculiar slowdown of intra-PSII electron transport during induction at low light intensities. The lower than F post-flash fluorescence yield F was related to oxidized tyrosine Z. The transient J peak was associated with equal rates of electron arrival to and departure from Q and requires that electron transfer from Q to Q be slower than that from Q to Q. Strong quenching by oxidized P680 caused the dip D. Reduced plastoquinone, a competitive product inhibitor of PSII, blocked electron transport proportionally with its concentration. Electron transport rate indicated by fluorescence quenching was faster than the rate indicated by O evolution, because oxidized donor side carriers quench fluorescence but do not transport electrons. The thermal phase of the fluorescence rise beyond the J phase was caused by a progressive increase in the fraction of PSII with reduced Q and reduced donor side.

摘要

在不同光强下诱导时,叶绿素荧光的 OJDIP 升高,使用描述通过光系统 II(PSII)的电子传递的 24 个主方程和用于质体醌、细胞色素 f、质体蓝素、光系统 I 和铁氧还蛋白中电子预算的常微分方程对其进行了数学建模。该模型的一个新特点是考虑了电子流入和流出预算,导致 Tyrosine Z、P680 和 Q 的氧化还原状态发生变化,从而导致瞬变期间荧光产率发生变化。除了在低光强下诱导时 PSII 内电子传递的奇特减速外,不需要跨膜电场、蛋白质构象变化或其他假定的猝灭物质的特定贡献来解释该现象的主要特征。低于 F 后闪光荧光产率 F 的值与氧化的酪氨酸 Z 有关。瞬态 J 峰与 Q 处电子到达和离开的相等速率相关,并且要求从 Q 到 Q 的电子转移比从 Q 到 Q 的电子转移慢。氧化的 P680 引起强烈猝灭,导致 D 点。还原的质体醌是 PSII 的竞争性产物抑制剂,其浓度与电子传递成比例地被阻断。荧光猝灭指示的电子传递速率比 O 释放指示的电子传递速率快,因为氧化的供体侧载体猝灭荧光但不传递电子。在 J 相之后的荧光上升的热相是由具有还原的 Q 和还原的供体侧的 PSII 分数逐渐增加引起的。

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