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Temperature-dependent regulation of electron transport and ATP synthesis in chloroplasts in vitro and in silico.
Photosynthesis Research ( IF 2.9 ) Pub Date : 2020-08-11 , DOI: 10.1007/s11120-020-00777-0
Alexander N Tikhonov 1, 2 , Alexey V Vershubskii 1
Affiliation  

The significance of temperature-dependent regulation of photosynthetic apparatus (PSA) is determined by the fact that plant temperature changes with environmental temperature. In this work, we present a brief overview of temperature-dependent regulation of photosynthetic processes in class B chloroplasts (thylakoids) and analyze these processes using a computer model that takes into account the key stages of electron and proton transport coupled to ATP synthesis. The rate constants of partial reactions were parametrized on the basis of experimental temperature dependences of partial photosynthetic processes: (1) photosystem II (PSII) turnover and plastoquinone (PQ) reduction, (2) the plastoquinol (PQH2) oxidation by the cytochrome (Cyt) b6f complex, (3) the ATP synthase activity, and (4) the proton leak from the thylakoid lumen. We consider that PQH2 oxidation is the rate-limiting step in the intersystem electron transport. The parametrization of the rate constants of these processes is based on earlier experimental data demonstrating strong correlations between the functional and structural properties of thylakoid membranes that were probed with the lipid-soluble spin labels embedded into the membranes. Within the framework of our model, we could adequately describe a number of experimental temperature dependences of photosynthetic reactions in thylakoids. Computer modeling of electron and proton transport coupled to ATP synthesis supports the notion that PQH2 oxidation by the Cyt b6f complex and proton pumping into the lumen are the basic temperature-dependent processes that determine the overall electron flux from PSII to molecular oxygen and the net ATP synthesis upon variations of temperature. The model describes two branches of the temperature dependence of the post-illumination reduction of \( {\text{P}}_{700}^{ + } \) characterized by different activation energies (about 60 and ≤ 3.5 kJ mol−1). The model predicts the bell-like temperature dependence of ATP formation, which arises from the balance of several factors: (1) the thermo-induced acceleration of electron transport through the Cyt b6f complex, (2) deactivation of PSII photochemistry at sufficiently high temperatures, and (3) acceleration of the passive proton outflow from the thylakoid lumen bypassing the ATP synthase complex. The model describes the temperature dependence of experimentally measured parameter P/2e, determined as the ratio between the rates of ATP synthesis and pseudocyclic electron transport (H2O → PSII → PSI → O2).



中文翻译:

体外和计算机中叶绿体中电子传递和 ATP 合成的温度依赖性调节。

光合装置(PSA)的温度依赖性调节的重要性取决于植物温度随环境温度变化的事实。在这项工作中,我们简要概述了 B 类叶绿体(类囊体)光合过程的温度依赖性调节,并使用计算机模型分析了这些过程,该模型考虑了与 ATP 合成耦合的电子和质子传输的关键阶段。部分反应的速率常数基于部分光合作用过程的实验温度依赖性进行参数化:(1)光系统 II(PSII)转换和质体醌(PQ)还原,(2)细胞色素对质体醌(PQH 2)氧化(细胞色素) b 6 f复合体,(3) ATP 合酶活性,和 (4) 类囊体腔的质子泄漏。我们认为 PQH 2氧化是系统间电子传输的限速步骤。这些过程的速率常数的参数化基于早期的实验数据,表明类囊体膜的功能和结构特性之间存在很强的相关性,这些膜通过嵌入膜中的脂溶性自旋标记进行探测。在我们的模型框架内,我们可以充分描述类囊体中光合作用反应的许多实验温度依赖性。与 ATP 合成耦合的电子和质子传输的计算机建模支持Cyt b 6氧化PQH 2的观点f复合物和质子泵入管腔是基本的温度依赖性过程,它决定了从 PSII 到分子氧的总电子通量以及温度变化时的净 ATP 合成。该模型描述了\( {\text{P}}_{700}^{ + } \)的光照后还原温度依赖性的两个分支,其特征在于不同的活化能(约 60 和 ≤ 3.5 kJ mol -1)。该模型预测了 ATP 形成的钟状温度依赖性,这源于以下几个因素的平衡:(1) 通过 Cyt b 6 f的电子传输的热诱导加速复合物,(2)在足够高的温度下使 PSII 光化学失活,以及(3)加速被动质子从类囊体腔绕过 ATP 合酶复合物流出。该模型描述了实验测量参数 P/2e 的温度依赖性,确定为 ATP 合成速率与伪环电子传输速率之间的比率(H 2 O → PSII → PSI → O 2)。

更新日期:2020-08-11
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