Robert Hill Institute and Department of Animal and Plant Sciences, University of Sheffield, Sheffield S10 2TN, UK.
J Plant Physiol. 2013 Mar 1;170(4):378-88. doi: 10.1016/j.jplph.2012.10.011. Epub 2012 Dec 13.
C(3) photosynthesis is an inefficient process, because the enzyme that lies at the heart of the Benson-Calvin cycle, ribulose 1,5-bisphosphate carboxylase-oxygenase (Rubisco) is itself a very inefficient enzyme. The oxygenase activity of Rubisco is an unavoidable side reaction that is a consequence of its reaction mechanism. The product of oxygenation, glycollate 2-P, has to be retrieved by photorespiration, a process which results in the loss of a quarter of the carbon that was originally present in glycollate 2-P. Photorespiration therefore reduces carbon gain. Purely in terms of carbon economy, there is, therefore, a strong selection pressure on plants to reduce the rate of photorespiration so as to increase carbon gain, but it also improves water- and nitrogen-use efficiency. Possibilities for the manipulation of plants to decrease the amount of photorespiration include the introduction of improved Rubisco from other species, reconfiguring photorespiration, or introducing carbon-concentrating mechanisms, such as inorganic carbon transporters, carboxysomes or pyrenoids, or engineering a full C(4) Kranz pathway using the existing evolutionary progression in C(3)-C(4) intermediates as a blueprint. Possible routes and progress to suppressing photorespiration by introducing C(4) photosynthesis in C(3) crop plants will be discussed, including whether single cell C(4) photosynthesis is feasible, how the evolution of C(3)-C(4) intermediates can be used as a blueprint for engineering C(4) photosynthesis, which pathway for the C(4) cycle might be introduced and the extent to which processes and structures in C(3) plant might require optimisation.
C(3) 光合作用是一个低效的过程,因为位于 Benson-Calvin 循环核心的酶,核酮糖 1,5-二磷酸羧化酶-加氧酶(Rubisco)本身就是一种非常低效的酶。Rubisco 的加氧酶活性是其反应机制不可避免的副反应。加氧的产物,2-磷酸甘油酸,必须通过光呼吸来回收,这个过程导致原本存在于 2-磷酸甘油酸中的四分之一的碳损失。因此,光呼吸会降低碳的获取量。纯粹从碳经济的角度来看,植物有强烈的选择压力来降低光呼吸速率,以增加碳的获取量,但它也提高了水和氮的利用效率。操纵植物减少光呼吸的可能性包括引入来自其他物种的改良 Rubisco、重新配置光呼吸或引入碳浓缩机制,如无机碳转运蛋白、羧化体或淀粉核,或使用现有的 C(3)-C(4)中间产物的进化进展作为蓝图,设计完整的 C(4) Kranz 途径。将 C(4)光合作用引入 C(3)作物植物中以抑制光呼吸的可能途径和进展将进行讨论,包括单细胞 C(4)光合作用是否可行,C(3)-C(4)中间产物的进化如何可以作为工程 C(4)光合作用的蓝图,C(4)循环的哪个途径可能被引入,以及 C(3)植物的过程和结构需要优化的程度。