面向碳达峰与碳中和的植物合成生物学

doi:10.12211/2096-8280.2022-034

合成生物学 ›› 2022, Vol. 3 ›› Issue (5): 847-869.DOI: 10.12211/2096-8280.2022-034

面向碳达峰与碳中和的植物合成生物学

杨健钊¹^,², 朱新广¹

^1.中国科学院分子植物科学卓越创新中心，上海植物生理生态研究所，植物分子遗传国家重点实验室，上海 200032
^2.中国科学院大学，北京 100049

收稿日期:2022-06-11 修回日期:2022-08-26 出版日期:2022-10-31 发布日期:2022-11-16
通讯作者: 朱新广
作者简介:杨健钊（1997—），男，博士研究生。主要研究方向为光合作用合成生物学与系统生物学。 E-mail：yangjianzhao@cemps.ac.cn
朱新广（1974—），男，研究员，博士生导师。长期开展多尺度光合系统模型创建、数字植物模型创建、高光效元件库创建、光合表型组学平台研发、碳汇植物创制、农林碳汇系统创制等领域的研究。 E-mail：zhuxg@cemps.ac.cn
基金资助:
国家重点研发计划(2018YFA0900600);国家自然科学基金(31870214);中科院战略性先导项目(XDB27020105)

Plant synthetic biology for carbon peak and carbon neutrality

Jianzhao YANG¹^,², Xinguang ZHU¹

^1.National Key Laboratory of Plant Molecular Genetics，CAS Center for Excellence in Molecular Plant Sciences，Shanghai Institute of Plant Physiology and Ecology，Chinese Academy of Sciences，Shanghai 200032，China
^2.University of Chinese Academy of Sciences，Beijing 100049，China

Received:2022-06-11 Revised:2022-08-26 Online:2022-10-31 Published:2022-11-16
Contact: Xinguang ZHU

摘要/Abstract

摘要：

合成生物学是一门包括生物学、数学、物理学、化学、工程及信息科学等多学科交叉的前沿学科。历经多年发展，目前在细菌、酵母等微生物和哺乳动物细胞底盘中已初步形成了完备的定量化研究体系，然而植物合成生物学依然处于起步阶段。植物合成生物学可在挖掘植物次生代谢天然产物、生产分子农业制品、提升光合作用光能利用效率、创制碳汇植物和建立植物工厂系统等方面发挥作用。特别是在当前面向碳达峰与碳中和过程中，植物合成生物学可以参与解决人类活动所面临的粮食短缺、能源危机及环境污染等问题。不同植物底盘中也在逐步建成和完善标准化的元件体系、基因线路设计以及定向进化等合成生物学技术。本文回顾了近年来植物合成生物学的主要研究进展，详述了植物合成生物学在“双碳”目标中的应用领域，包含植物天然产物合成与次生代谢、分子农业、光合作用、碳汇植物的创制、植物工厂等。提出了植物基因文件标准化、植物基因线路设计、植物基因编辑和定向进化等助力“双碳”目标的植物合成生物学技术，并展望和探讨了在实现“双碳”目标过程中植物合成生物学的重要作用和发展前景。

关键词: 植物合成生物学, 碳达峰, 碳中和, 天然产物, 光合作用, 分子农业, 碳汇植物, 植物工厂

Abstract:

Synthetic biology is an interdisciplinary research field, for which complete quantitative research systems have been established in bacteria, yeast, and mammalian cells. However, synthetic biology in plants is still at its infancy. Plant synthetic biology can play important roles in synthesizing plant natural products, developing molecular farming, improving photosynthesis to increase light energy utilization efficiency, designing carbon farming plants, and building plant factories. In the current efforts in creating a carbon neutral society, plant synthetic biology can help to address challenges of food shortage, energy crisis, and environmental pollution. Specifically, innovative methods can be developed to reduce the emission of CO₂ and pollutants through plant production of high value products, whose industrial production is mostly associated with high CO₂ emission. Moreover, plant synthetic biology can be used to optimize plant production through minimizing carbon emissions and reducing the use of chemical fertilizers and pesticides. Furthermore, plants specialized in carbon capturing, such as high photosynthetic efficiency, large root systems, and high resistance to degradation, should be developed as well. Various options for increased photosynthetic efficiency, such as optimizing the antenna size of photosystem, converting C₃ to C₄ photosynthesis, introducing CO₂ concentrating mechanisms, and establishing the photorespiration bypasses into C₃ crops, holds the potential to dramatically increase the carbon capturing capacity for improved productivity. In the future, in addition to crops, trees and algae can also be engineered to become efficient carbon sinks. Photosynthetic algae are expected to become a source of clean energy and industrial production system with zero or negative carbon emissions. In the long term, a complete plant factory system, which has optimal control of light, temperature, CO₂, water, and nutrient, will be developed to achieve optimal plant growth and production while maintaining maximal carbon capturing capacity. Finally, artificial photosynthesis also promises to be an ideal solution as an energy production system. These aspects will be facilitated by the rapid development of plant synthetic biology tools, including biological part standardization, genetic circuits design, and directed evolution. This paper summarizes the major progresses of plant synthetic biology and prospects the major roles of plant synthetic biology in the future efforts in carbon emission peak and carbon neutrality.

Key words: plant synthetic biology, carbon emission peak, carbon neutrality, natural product, photosynthesis, molecular farming, carbon farming, plant factory

中图分类号:

杨健钊, 朱新广. 面向碳达峰与碳中和的植物合成生物学[J]. 合成生物学, 2022, 3(5): 847-869.

Jianzhao YANG, Xinguang ZHU. Plant synthetic biology for carbon peak and carbon neutrality[J]. Synthetic Biology Journal, 2022, 3(5): 847-869.

图/表 5

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针对疾病	应用案例	植物底盘	参考文献
霍乱	叶绿体表达霍乱弧菌毒素B亚基（CTB）	烟草、生菜、玉米、水稻	[79-80]
乙肝	玉米胚芽表达乙肝抗原HBsAg	玉米	[81]
脊髓灰质炎	烟草表达人脊髓灰质炎病毒L1蛋白	烟草	[82]
疟疾	叶绿体表达疟原虫的AMA1和MSP1蛋白	衣藻	[83]
糖尿病	表达人源小胰岛素原	番茄、灵芝	[84]
口腔生物膜（龋齿）	生菜叶绿体中表达脂肪酶、葡聚糖酶和抗菌肽	生菜	[85]
新冠病毒肺炎	组装新冠病毒VLPs	烟草	[86]

针对疾病	应用案例	植物底盘	参考文献
霍乱	叶绿体表达霍乱弧菌毒素B亚基（CTB）	烟草、生菜、玉米、水稻	[79-80]
乙肝	玉米胚芽表达乙肝抗原HBsAg	玉米	[81]
脊髓灰质炎	烟草表达人脊髓灰质炎病毒L1蛋白	烟草	[82]
疟疾	叶绿体表达疟原虫的AMA1和MSP1蛋白	衣藻	[83]
糖尿病	表达人源小胰岛素原	番茄、灵芝	[84]
口腔生物膜（龋齿）	生菜叶绿体中表达脂肪酶、葡聚糖酶和抗菌肽	生菜	[85]
新冠病毒肺炎	组装新冠病毒VLPs	烟草	[86]

类型	改造方向	改造策略	改造效果	改造底盘	参考文献
光反应	改良NPQ	过表达拟南芥的NPQ相关元件VDE、ZEP、PsbS	生物量积累增加15%	烟草	[97]
	光合天线优化	减小或重分配天线尺寸	预测冠层光合可最多提升30%	水稻	[100]
	补充易损光合元件	核转基因过表达PS Ⅱ的D1转运进叶绿体	生物量较野生型显著增加（拟南芥43.7%～80.2%，烟草15.1%～22.3%，水稻20.6%～22.9%）	拟南芥、烟草、水稻	[101]
碳代谢	改造光呼吸支路	引入大肠杆菌的乙醛酸代谢途径	干重较野生型提高1倍以上	拟南芥	[117]
		构建乙醇酸-草酸（GOC）氧化途径	光合效率和生物量分别可提高15%～22%和14%～35%	水稻	[118]
		叶绿体内建立GCGT捷径	光合效率可提高6%～16%	水稻	[115]
		叶绿体内让乙醇酸转变为苹果酸	生物量可比野生型提高约40%	烟草	[119]
	改造RuBisCO	引入藻类Red-RuBisCO	加快植物生长，并且可以作为定向进化改造目标蛋白	烟草	[127]
光合转录调控	增强光合基因表达	过表达EmBP1	提高光合速率、生物量及产量可达25%	水稻	[106]
	敲除光合表达负调控因子	敲除NRP1	光合速率和生物量都有显著增加	水稻	[107]
	调控光合和氮素利用效率	过表达DREB1C	可提高产量30%以上	水稻	[109]
人工光合作用	人工光反应结合生物固碳体系	硅纳米线与固碳细菌共培养	最高能量转换效率可达3.6%	卵型孢子菌（Sporomusaovata）	[137-139]
		光伏电池与蓝细菌PS Ⅰ偶联驱动固碳体系	总能量转化效率可达到9%	蓝细菌	[140]
	人工模拟叶绿体	微流控包裹类囊体	在光照下连续固定CO₂转化为有机酸	菠菜	[141]
	大肠杆菌内构建固碳通路	重组RuBisCO	可以固定CO₂	大肠杆菌	[130]
		构建卡尔文循环	以CO₂为碳源合成糖和其他有机物	大肠杆菌	[142]
	无细胞体系构建固碳通路	CO₂为碳源从头合成淀粉	较玉米中淀粉合成速率高约8.5倍	无细胞体系	[143]

类型	改造方向	改造策略	改造效果	改造底盘	参考文献
光反应	改良NPQ	过表达拟南芥的NPQ相关元件VDE、ZEP、PsbS	生物量积累增加15%	烟草	[97]
	光合天线优化	减小或重分配天线尺寸	预测冠层光合可最多提升30%	水稻	[100]
	补充易损光合元件	核转基因过表达PS Ⅱ的D1转运进叶绿体	生物量较野生型显著增加（拟南芥43.7%～80.2%，烟草15.1%～22.3%，水稻20.6%～22.9%）	拟南芥、烟草、水稻	[101]
碳代谢	改造光呼吸支路	引入大肠杆菌的乙醛酸代谢途径	干重较野生型提高1倍以上	拟南芥	[117]
		构建乙醇酸-草酸（GOC）氧化途径	光合效率和生物量分别可提高15%～22%和14%～35%	水稻	[118]
		叶绿体内建立GCGT捷径	光合效率可提高6%～16%	水稻	[115]
		叶绿体内让乙醇酸转变为苹果酸	生物量可比野生型提高约40%	烟草	[119]
	改造RuBisCO	引入藻类Red-RuBisCO	加快植物生长，并且可以作为定向进化改造目标蛋白	烟草	[127]
光合转录调控	增强光合基因表达	过表达EmBP1	提高光合速率、生物量及产量可达25%	水稻	[106]
	敲除光合表达负调控因子	敲除NRP1	光合速率和生物量都有显著增加	水稻	[107]
	调控光合和氮素利用效率	过表达DREB1C	可提高产量30%以上	水稻	[109]
人工光合作用	人工光反应结合生物固碳体系	硅纳米线与固碳细菌共培养	最高能量转换效率可达3.6%	卵型孢子菌（Sporomusaovata）	[137-139]
		光伏电池与蓝细菌PS Ⅰ偶联驱动固碳体系	总能量转化效率可达到9%	蓝细菌	[140]
	人工模拟叶绿体	微流控包裹类囊体	在光照下连续固定CO₂转化为有机酸	菠菜	[141]
	大肠杆菌内构建固碳通路	重组RuBisCO	可以固定CO₂	大肠杆菌	[130]
		构建卡尔文循环	以CO₂为碳源合成糖和其他有机物	大肠杆菌	[142]
	无细胞体系构建固碳通路	CO₂为碳源从头合成淀粉	较玉米中淀粉合成速率高约8.5倍	无细胞体系	[143]

类型	应用领域	内容	应用对象	参考文献
标准化植物基因元件	克隆和元件标准化体系	植物标准化克隆体系（Plant MoClo Syntax）	蓝藻、衣藻等	[184-185]
		衣藻元件工具库（Chlamy MoClo Toolkit）	衣藻	[186]
	创制合成生物学底盘	衣藻突变体表型库	衣藻	[187]
		酵母内融合蓝细菌内共生	衣藻	[188]
植物基因线路设计	改造信号通路	人工脱落酸受体	拟南芥	[190]
	构建基因线路	TEV蛋白酶逻辑门	烟草	[191]
		叶绿体内乳糖操纵子	衣藻	[193]
		叶绿体核糖体开关（Riboswitch）	衣藻	[194]
植物基因编辑与定向进化技术	精准基因编辑	基因组先导编辑	水稻、番茄、玉米	[202-204]
植物基因编辑与定向进化技术	精准基因编辑	质体基因编辑	生菜、油菜	[205]
	定向进化	饱和靶向内源诱变编辑器（STEME）	水稻	[217]

面向碳达峰与碳中和的植物合成生物学

Plant synthetic biology for carbon peak and carbon neutrality

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