植物二氧化碳代谢途径改造研究进展

doi:10.12211/2096-8280.2022-002

合成生物学 ›› 2022, Vol. 3 ›› Issue (5): 985-1005.DOI: 10.12211/2096-8280.2022-002

植物二氧化碳代谢途径改造研究进展

史梦琳¹^,², 周琳¹^,³, 王庆¹^,³, 赵磊¹^,²^,³

^1.中国科学院天津工业生物技术研究所，天津 300308
^2.中国科学院系统微生物工程重点实验室，天津 300308
^3.国家合成生物技术创新中心，天津 300308

收稿日期:2022-01-12 修回日期:2022-02-17 出版日期:2022-10-31 发布日期:2022-11-16
通讯作者: 赵磊
作者简介:史梦琳（1991—），女，博士后。研究方向为植物固碳途径解析与改造等。 E-mail：shiml@tib.cas.cn
赵磊（1985—），男，博士，研究员。研究方向为植物合成生物学与代谢工程等。 E-mail：zhaol@tib.cas.cn
基金资助:
天津市合成生物技术创新能力提升行动项目(TSBICIP-CXRC-027);中国科技部科技合作计划项目(KY202001017)

Advances in the study on the modification of carbon dioxide metabolic pathways in plants

Menglin SHI¹^,², Lin ZHOU¹^,³, Qing WANG¹^,³, Lei ZHAO¹^,²^,³

^1.Tianjin Institute of Industrial Biotechnology，Chinese Academy of Sciences，Tianjin 300308，China
^2.Key Laboratory of Systems Microbial Biotechnology，Tianjin Institute of Industrial Biotechnology，Chinese Academy of Sciences，Tianjin 300308，China
^3.National Center of Technology Innovation for Synthetic Biology，Tianjin 300308，China

Received:2022-01-12 Revised:2022-02-17 Online:2022-10-31 Published:2022-11-16
Contact: Lei ZHAO

摘要/Abstract

摘要：

降碳减排是我国可持续发展过程中的一项重大战略决策。为如期实现“碳达峰、碳中和”宏伟目标，我国需在提高生态碳汇能力上取得突破。植物光合作用有利于增加地球碳汇，而光呼吸和呼吸作用过程则释放CO₂。自然状态下，上述CO₂代谢过程能量利用率低、人工改造并提高植物CO₂固定的难度较大。因此，在植物体内重构新的人工代谢途径，有望大幅提高植物CO₂固定能力，是解决人类社会发展瓶颈的有效途径之一。本文分别介绍了植物光合作用、光呼吸和呼吸作用中与CO₂固定、释放相关途径，并指出可用于改造的潜在靶点；重点综述了植物体内已构建的人工固碳途径及其代谢原理，系统分析评价了不同途径的CO₂固定能力和限制因素；最后，对人工设计及合成植物CO₂代谢通路、基于零碳排放的新型物质生产等关键问题进行了探讨，并对植物CO₂代谢途径的改造发展趋势进行了展望。

关键词: 植物, CO₂代谢, 改造, 合成生物学, 碳中和

Abstract:

Reducing carbon emissions is a major strategic decision in the process of sustainable development in China. In order to achieve the ambitious goal of “carbon peak， carbon neutrality” on schedule, China needs to make a breakthrough in improving the ecological carbon sink capacity. Plant photosynthesis is beneficial to increase the Earth's carbon sink by fixing atmospheric carbon dioxide or inorganic carbon to produce organic compounds, while CO₂ is released in the process of photorespiration and respiration which reduce carbon sinks by degrading the organic compounds into other substrates. Energy utilization rate of the above-mentioned CO₂ natural metabolic process is low, and those processes are difficult to be artificially modified and improved in plants due to the characters and limitations of the relevant enzymes. Therefore, reconstructing new artificial metabolic pathways with synthetic biology in plants is expected to greatly improve the plant CO₂ fixation capacity, which is one of the effective ways to solve the bottleneck of humanity development in the future. In the present review, we introduce the metabolic pathways associated with CO₂ fixation and release which are involved in plant photosynthesis, photorespiration and respiration, respectively, and then point out the potential targets that could be used for modification by synthetic biology. In each section, we mainly discuss the artificial carbon fixation pathways that have been implemented in plants and their underlying principles. Especially, the modification of photorespiration is particularly discussed and several pathways are mentioned in details which shed lights on the design of artificial pathway in the future. Then we compare the capacity of each pathway in carbon fixation and limitation. Finally, we propose the key questions of designing and synthesizing novel carbon fixation pathway in plants, and the zero-carbon releasing design is mainly discussed. The development trend of transformation of plant CO₂ metabolic pathway by synthetic biology is also forecasted.

Key words: plant, CO₂ metabolism, synthetic biology, modification, synthetic biology, carbon neutrality

中图分类号:

Q945

史梦琳, 周琳, 王庆, 赵磊. 植物二氧化碳代谢途径改造研究进展[J]. 合成生物学, 2022, 3(5): 985-1005.

Menglin SHI, Lin ZHOU, Qing WANG, Lei ZHAO. Advances in the study on the modification of carbon dioxide metabolic pathways in plants[J]. Synthetic Biology Journal, 2022, 3(5): 985-1005.

图/表 6

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涉及部分	改造对象或方式	改造效果
光合作用	光系统天线	降低光损耗、提高光能利用率和生物量
	光系统对波动光的响应能力	有望提高光系统对波动光的响应能力，减少光抑制
	天然光合系统吸光范围	有望将植物吸光范围扩展到远红区域，进而增强光合效率
	CO₂扩散能力	可改善叶片内部CO₂扩散特性，是提高光合效率的有效途径；其扩散能力与温度、细胞壁厚度和组成等均相关。由于叶片CO₂扩散速率测定较为困难，致该部分研究缓慢
	Rubisco催化特性	Rubisco催化特性改造尚未取得实质性进展；通过筛选不同物种特异性Rubisco或将Rubisco与碳浓缩机制改造相结合，有望改善植物固碳能力
	Rubisco附近的CO₂浓度	通过将CCM系统、蛋白核或将C₄光合系统引入C₃作物等，可增加CO₂供给。但需要考虑光合过程中生化反应的变化及叶片结构的改变，C₃向C₄植物转化研究仍待加强
光呼吸	天然光呼吸途径	改造较为复杂，需考虑较多因素；增加光呼吸通量可使植物更好地应对高光呼吸胁迫，提高生物量，但具体改造靶点和策略仍待探索
	新型光呼吸替代途径	导入新型光呼吸替代途径，可将毒性副产物化为其他生物质，同时将CO₂重新释放到叶绿体中，在提高CO₂浓度的同时减少植物固碳损失；基于零CO₂释放的新型替代通路，可避免光呼吸中的碳损失，在增加植物固碳方面有较大潜力
	光呼吸代谢通量模型计算	将计算技术与基因组工程、合成生物学技术等技术结合，进行代谢通量模型分析，可简化和评估光呼吸通路改造设计，优化实验设计
呼吸作用	交替氧化酶AOX	呼吸作用对植物基础代谢尤为重要，对其进行改造的操作空间较小；不同体系和生长条件下，AOX表达量变化对固碳和生长影响不同，对其进行有效改造的策略仍待探索中

涉及部分	改造对象或方式	改造效果
光合作用	光系统天线	降低光损耗、提高光能利用率和生物量
	光系统对波动光的响应能力	有望提高光系统对波动光的响应能力，减少光抑制
	天然光合系统吸光范围	有望将植物吸光范围扩展到远红区域，进而增强光合效率
	CO₂扩散能力	可改善叶片内部CO₂扩散特性，是提高光合效率的有效途径；其扩散能力与温度、细胞壁厚度和组成等均相关。由于叶片CO₂扩散速率测定较为困难，致该部分研究缓慢
	Rubisco催化特性	Rubisco催化特性改造尚未取得实质性进展；通过筛选不同物种特异性Rubisco或将Rubisco与碳浓缩机制改造相结合，有望改善植物固碳能力
	Rubisco附近的CO₂浓度	通过将CCM系统、蛋白核或将C₄光合系统引入C₃作物等，可增加CO₂供给。但需要考虑光合过程中生化反应的变化及叶片结构的改变，C₃向C₄植物转化研究仍待加强
光呼吸	天然光呼吸途径	改造较为复杂，需考虑较多因素；增加光呼吸通量可使植物更好地应对高光呼吸胁迫，提高生物量，但具体改造靶点和策略仍待探索
	新型光呼吸替代途径	导入新型光呼吸替代途径，可将毒性副产物化为其他生物质，同时将CO₂重新释放到叶绿体中，在提高CO₂浓度的同时减少植物固碳损失；基于零CO₂释放的新型替代通路，可避免光呼吸中的碳损失，在增加植物固碳方面有较大潜力
	光呼吸代谢通量模型计算	将计算技术与基因组工程、合成生物学技术等技术结合，进行代谢通量模型分析，可简化和评估光呼吸通路改造设计，优化实验设计
呼吸作用	交替氧化酶AOX	呼吸作用对植物基础代谢尤为重要，对其进行改造的操作空间较小；不同体系和生长条件下，AOX表达量变化对固碳和生长影响不同，对其进行有效改造的策略仍待探索中

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植物二氧化碳代谢途径改造研究进展

Advances in the study on the modification of carbon dioxide metabolic pathways in plants

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