光合作用碳同化的合成生物学研究进展

doi:10.12211/2096-8280.2022-019

摘要/Abstract

摘要：

随着人口增多及耕地面积的减少，人类对粮食的需求日益增加，因此保障足够的粮食供给尤为重要。光合作用通过光反应和碳同化把无机物转换成有机物，是地球上最重要的化学反应。90%以上的植物干物质来源于光合作用固碳反应，光合作用同化的有机物是作物产量形成的物质基础，因此提高作物光能利用效率是提高作物产量的重要途径。近年来，合成生物学在能源、材料、健康和环境等多领域的快速发展，为提高植物光合效率提供了新的机遇。本文着重讨论了合成生物学在提高光合作用碳同化效率方面的研究进展，主要集中在提高Rubisco酶的羧化活性、引进CO₂浓缩机制、降低光呼吸等方面；最后，对新型光合固碳回路进行探讨，通过合成生物学对光合作用碳同化模块进行设计、改造、优化和重组，必将有效提高碳同化效率，最终提高作物产量。

关键词: 光合作用, 光合作用效率, 合成生物学, 碳同化

Abstract:

With the increase of population and the decrease of cultivated land, human demand for food is increasing. Therefore, it is particularly important to ensure adequate food supply. Photosynthesis is the most important chemical reaction on the earth, which converts inorganic matter into organic matter through light reaction and carbon assimilation. More than 90% of plant dry matter comes from the carbon fixation reaction of photosynthesis. The assimilated organic matter of photosynthesis is the material basis for the formation of crop yield. Therefore, improving the efficiency of crop light energy utilization is an important way to improve crop yield. In recent years, the rapid development of synthetic biology in the fields of energy, materials, health and environment has provided new opportunities for improving plant photosynthetic efficiency. This paper highlights the research progress of synthetic biology in improving the carbon assimilation efficiency of photosynthesis, mainly focusing on: (1) Improving the carboxylation activity of Rubisco enzymes, including identifying Rubisco enzymes with high carboxylation activity, optimizing gene expression regulatory sequences on Rubisco, and co-expressing Rubisco chaperone proteins; (2) Introducing CO₂ concentrating mechanisms, including C₄ photosynthetic enzymes, cyanobacterial transporter proteins, and cyanobacterial carboxysomes; (3) Reducing photorespiration through the introduction of four photorespiratory branches: chloroplast glycerate bypass, peroxisomal glycerate bypass, chloroplast glycolate oxidation bypass, and 3-hydroxypropionate bypass, and the exploration of new branches of photorespiration; Finally, the new photosynthetic carbon fixation circuit is discussed. The design, transformation, optimization and reorganization of photosynthetic carbon assimilation module through synthetic biology will effectively improve the efficiency of carbon assimilation and ultimately improve crop yield.

Key words: photosynthesis, photosynthetic efficiency, synthetic biology, carbon assimilation

中图分类号:

盛阳阳, 徐秀美, 张巧红, 张立新. 光合作用碳同化的合成生物学研究进展[J]. 合成生物学, 2022, 3(5): 870-883.

Yangyang SHENG, Xiumei XU, Qiaohong ZHANG, Lixin ZHANG. Advances in synthetic biology for photosynthetic carbon assimilation[J]. Synthetic Biology Journal, 2022, 3(5): 870-883.

图/表 5

图1 提高光合作用碳同化效率总思路Ru-5-P—5-磷酸核酮糖；RUBP—1，5-二磷酸核酮糖；G2P—2-磷酸甘油酸；3-PGA—3-磷酸甘油酸；GAP—丙糖磷酸Ru-5-P—Ribulose 5-phosphate; RUBP—Ribulose 1,5-bisphosphate; G2P—2-phosphoglyceric acid; 3-PGA—3-磷酸甘油酸; GAP—Triose phosphate

Fig. 1 Overview of improving carbon assimilation efficiency of photosynthesis

表1 Rubisco酶活性的合成生物学研究汇总

Tab. 1 Summary of Rubisco enzyme activity by synthetic biological research

提高Rubisco酶的羧化活性	研究策略	参考文献
寻找其他物种中高活性的Rubisco酶	高羧化酶活性	[29-31]
寻找其他物种中高活性的Rubisco酶	高亲和力	[32]
筛选Rubisco酶高活性品种	高羧化酶活性	[33]
人工合成肽	无明显作用	[34]
引进Rubisco生物合成依赖的辅助因子	折叠伴侣蛋白、组装伴侣蛋白及活化酶	[35-37]

表2 CO2浓缩机制的合成生物学研究汇总

Tab. 2 Summary of CO2 enrichment mechanisms by synthetic biological research

引进CO₂浓缩机制	研究策略	参考文献
GLK基因的引入	促进产量增加	[65-66]
转运蛋白的引入	条件促进，可以提高光合速率和碳同化率	[68-69]
蓝藻羧酶体的引入	有待进一步探索研究	[84-85]

表3 光呼吸支路的合成生物学研究汇总

Tab. 3 Summary of photorespiration pathways by synthetic biological research

降低光呼吸	研究策略	参考文献
叶绿体甘油酸支路	叶绿体乙醇酸转化为甘油酸，生物量增加，无NH₃的释放	[96-97]
过氧化物酶体甘油酸支路	绕过了线粒体中的甘氨酸到丝氨酸的转化，同时将CO₂释放的位置从线粒体转移到过氧化物酶体，无NH₃的释放	[98]
叶绿体乙醇酸氧化支路	光呼吸途径的碳全部丢失，无NH₃的释放	[99-100]
3-羟基丙酸盐支路	实现光呼吸期间CO₂的净同化，无NH₃的释放	[101]

图2 天然和合成的光呼吸支路黑色箭头，经典的光呼吸旁路；蓝色箭头，叶绿体甘油酸支路；橙色箭头，过氧化物酶体甘油酸支路；绿色箭头，叶绿体乙醇酸氧化支路；紫色箭头，3-羟基丙酸盐支路；红色箭头，水稻中新的光呼吸支路RUBP—1，5-二磷酸核酮糖；G2P—2-磷酸甘油酸；3-PGA—3-磷酸甘油酸

Fig. 2 Natural and synthetic photorespiratory bypassesBlack arrow, classic photorespiratory bypass; Blue arrow, chloroplastic glycerate bypass; Orange arrow, peroxisomal glycerate bypass; Green arrow, chloroplastic glycolate oxidation bypass; Purple arrow, 3-hydroxypropionate bypass; Red arrow, A new photorespiratory bypasses in riceRUBP—Ribulose 1,5-bisphosphate; G2P—2-phosphoglyceric acid; 3-PGA—3-phosphoglyceric acid

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