细胞色素 b6f 复合物的光合作用控制

Gustaf E Degen, Matthew P Johnson
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摘要

光合控制(PCON)是一种保护机制,通过确保通过线性电子传递(LET)产生的 NADPH 和 ATP 的速率与 CO2 固定反应中的消耗相平衡,从而防止光诱导对光子系统 I(PSI)造成损害。保护 PSI 是植物的当务之急,因为植物缺乏专门针对这一复合体的快速修复循环,这意味着任何损伤都会导致长时间的光抑制和生长衰退。LET 与 CO2 固定反应之间的不平衡可通过转紫函 ΔpH 水平来感知,当光照过量时,ΔpH 会升高。PCON 的典型机制包括 ΔpH 对细胞色素 b6f 中 LET 的质醌氧化步骤的反馈控制。因此,PCON 使 PSI 特殊配对叶绿素(P700)保持氧化状态,从而使二氧化碳固定反应中未使用的多余电子通过电荷重组被安全淬灭。在这篇综述中,我们将以被子植物为研究对象,探讨 PSI 光氧化损伤是如何产生的,探讨 PSI 光抑制对光合作用和生长的影响,讨论在了解 PCON 调节方面的最新进展,最后探讨未来在作物中利用 PCON 提高光合效率的前景。
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Photosynthetic control at the cytochrome b6f complex
Photosynthetic control (PCON) is a protective mechanism that prevents light-induced damage to photosystem I (PSI) by ensuring the rate of NADPH and ATP production via linear electron transfer (LET) is balanced by their consumption in the CO2 fixation reactions. Protection of PSI is a priority for plants since they lack a dedicated rapid-repair cycle for this complex, meaning that any damage leads to prolonged photoinhibition and decreased growth. The imbalance between LET and the CO2 fixation reactions is sensed at the level of the transthylakoid ΔpH, which increases when light is in excess. The canonical mechanism of PCON involves feedback control by ΔpH on the plastoquinol oxidation step of LET at cytochrome b6f. PCON thereby maintains the PSI special pair chlorophylls (P700) in an oxidized state, that allows excess electrons unused in the CO2 fixation reactions to be safely quenched via charge recombination. In this review we focus on angiosperms, considering how photo-oxidative damage to PSI comes about, explore the consequences of PSI photoinhibition on photosynthesis and growth, discuss recent progress in understanding PCON regulation, and finally consider the prospects for its future manipulation in crop plants to improve photosynthetic efficiency.
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