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能量耗散是维持植物生存能力的一个重要机制:光合作用改良的生理限制。

Energy dissipation is an essential mechanism to sustain the viability of plants: The physiological limits of improved photosynthesis.

机构信息

Institut für Biologie I, Universität Leipzig, Johannisallee 21-23, Leipzig, Germany.

出版信息

J Plant Physiol. 2011 Jan 15;168(2):79-87. doi: 10.1016/j.jplph.2010.07.012. Epub 2010 Oct 8.

Abstract

In bright sunlight photosynthetic activity is limited by the enzymatic machinery of carbon dioxide assimilation. This supererogation of energy can be easily visualized by the significant increases of photosynthetic activity under high CO(2) conditions or other metabolic strategies which can increase the carbon flux from CO(2) to metabolic pools. However, even under optimal CO(2) conditions plants will provide much more NADPH+H(+) and ATP that are required for the actual demand, yielding in a metabolic situation, in which no reducible NADP(+) would be available. As a consequence, excited chlorophylls can activate oxygen to its singlet state or the photosynthetic electrons can be transferred to oxygen, producing highly active oxygen species such as the superoxide anion, hydroxyl radicals and hydrogen peroxide. All of them can initiate radical chain reactions which degrade proteins, pigments, lipids and nucleotides. Therefore, the plants have developed protection and repair mechanism to prevent photodamage and to maintain the physiological integrity of metabolic apparatus. The first protection wall is regulatory energy dissipation on the level of the photosynthetic primary reactions by the so-called non-photochemical quenching. This dissipative pathway is under the control of the proton gradient generated by the electron flow and the xanthophyll cycle. A second protection mechanism is the effective re-oxidation of the reduction equivalents by so-called "alternative electron cycling" which includes the water-water cycle, the photorespiration, the malate valve and the action of antioxidants. The third system of defence is the repair of damaged components. Therefore, plants do not suffer from energy shortage, but instead they have to invest in proteins and cellular components which protect the plants from potential damage by the supererogation of energy. Under this premise, our understanding and evaluation for certain energy dissipating processes such as non-photochemical quenching or photorespiration appear in a quite new perspective, especially when discussing strategies to improve the solar energy conversion into plant biomass.

摘要

在明亮的阳光下,光合作用受到二氧化碳同化酶机制的限制。通过在高 CO2 条件下或其他代谢策略下显著增加光合作用活性,可以很容易地看到这种能量的过剩,这些策略可以增加从 CO2 到代谢物池的碳通量。然而,即使在最佳 CO2 条件下,植物提供的 NADPH+H(+) 和 ATP 也远远超过实际需求,导致代谢状态下,没有可还原的 NADP(+)可用。因此,激发态的叶绿素可以将氧气激活到其单线态,或者光合作用电子可以转移到氧气上,产生高度活跃的氧物种,如超氧阴离子、羟基自由基和过氧化氢。所有这些都可以引发自由基链式反应,导致蛋白质、色素、脂质和核苷酸降解。因此,植物已经开发出保护和修复机制,以防止光损伤并维持代谢装置的生理完整性。第一道保护墙是通过所谓的非光化学猝灭来调节光合作用初级反应水平上的能量耗散。这种耗散途径受电子流和叶黄素循环产生的质子梯度的控制。第二种保护机制是通过所谓的“替代电子循环”有效地再氧化还原当量,其中包括水-水循环、光呼吸、苹果酸阀和抗氧化剂的作用。第三道防御系统是受损成分的修复。因此,植物不会遭受能量短缺,而是必须投资于蛋白质和细胞成分,以保护植物免受多余能量的潜在损害。在此前提下,我们对某些能量耗散过程的理解和评估,如非光化学猝灭或光呼吸,会呈现出全新的视角,尤其是在讨论提高太阳能转化为植物生物质的策略时。

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