代谢工程改造微生物固定二氧化碳研究进展

doi:10.16085/j.issn.1000-6613.2022-1545

摘要/Abstract

摘要：

微生物固定CO₂是实现CO₂资源化利用的有效策略之一，为固碳减排、节能生产与绿色合成提供了借鉴。然而，微生物在固定CO₂过程中存在底物利用效率低、能量需求量大、路径难优化等问题。为了解决这些问题，本文总结了6种天然CO₂固定途径与5种人工CO₂固定途径，并从自养微生物、异养微生物和人工微生物三个方面系统分析了代谢工程改造微生物固定CO₂合成化学品的最新进展。在自养微生物固定CO₂方面，采用的策略主要包括提高CO₂固定途径效率、开发能量捕集系统与调节碳代谢流分布；在异养微生物固定CO₂方面，常用的方法主要有强化羧化途径、重构CO₂固定途径与优化能量供给；在人工微生物固定CO₂方面，主要的研究思路是设计人工CO₂固定途径与构建人工CO₂固定微生物。最后，从CO₂固定的关键酶、途径和微生物三方面展望了进一步提高CO₂固定效率的发展方向。

关键词: 微生物, 代谢工程, 二氧化碳固定, 固碳减排

Abstract:

Microbial CO₂ fixation is one of the effective strategies to realize CO₂ resource utilization, which provides a reference for carbon abatement, energy-saving production and green synthesis. However, there are problems in microbial CO₂ fixation process, such as insufficient CO₂ utilization rate, high energy demand, and difficult pathway optimization. In order to solve these problems, this paper summarizes six natural CO₂ fixation pathways and five artificial CO₂ fixation pathways, and systematically analyzes the latest progress of metabolic engineering of microorganisms for CO₂ fixation from three aspects: autotrophic microorganisms, heterotrophic microorganisms and artificial microorganisms. For CO₂ fixation by autotrophic microorganisms, the main strategies include improving the efficiency of CO₂ fixation pathways, developing energy capture systems, and regulating the distribution of carbon metabolism. For CO₂ fixation by heterotrophic microorganisms, the main methods include strengthening the carboxylation pathway, remodeling the CO₂ fixation pathway, and optimizing the energy supply. For CO₂ fixation by artificial microorganisms, the main research ideas include the design of artificial CO₂ fixation pathways and the construction of artificial CO₂ fixation microorganisms. Finally, we further prospect the development direction of improving CO₂ fixation efficiency by engineering key enzymes, pathways and microorganisms.

Key words: microorganisms, metabolic engineering, CO₂ fixation, carbon abatement

中图分类号:

Q815

陶雨萱, 郭亮, 高聪, 宋伟, 陈修来. 代谢工程改造微生物固定二氧化碳研究进展[J]. 化工进展, 2023, 42(1): 40-52.

TAO Yuxuan, GUO Liang, GAO Cong, SONG Wei, CHEN Xiulai. Progress in metabolic engineering of microorganisms for CO₂ fixation[J]. Chemical Industry and Engineering Progress, 2023, 42(1): 40-52.

图/表 6

表1 天然CO2固定途径

CO₂固定途径	固定CO₂/HCO $3 -$	消耗能量	重要产物	氧气需求	参考文献
CBB循环	3mol CO₂	9mol ATP + 6mol NAD(P)H	3-磷酸甘油醛	好氧	[13]
RTCA途径	2mol CO₂	2mol ATP + 4mol NAD(P)H	乙酰辅酶A	厌氧/微氧	[14]
3-HP双循环	3mol HCO $3 -$	5mol ATP + 5mol NAD(P)H	丙酮酸	好氧	[15]
WL途径	2mol CO₂	1mol ATP + 4mol NAD(P)H	乙酰辅酶A	厌氧	[16]
DC-4HB循环	1mol CO₂ + 1mol HCO $3 -$	3mol ATP + 4mol NAD(P)H	乙酰辅酶A	厌氧/微氧	[17]
3HP-4HB循环	2mol HCO $3 -$	4mol ATP + 4mol NAD(P)H	乙酰辅酶A	好氧	[18]

表1 天然CO2固定途径

CO₂固定途径	固定CO₂/HCO $3 -$	消耗能量	重要产物	氧气需求	参考文献
CBB循环	3mol CO₂	9mol ATP + 6mol NAD(P)H	3-磷酸甘油醛	好氧	[13]
RTCA途径	2mol CO₂	2mol ATP + 4mol NAD(P)H	乙酰辅酶A	厌氧/微氧	[14]
3-HP双循环	3mol HCO $3 -$	5mol ATP + 5mol NAD(P)H	丙酮酸	好氧	[15]
WL途径	2mol CO₂	1mol ATP + 4mol NAD(P)H	乙酰辅酶A	厌氧	[16]
DC-4HB循环	1mol CO₂ + 1mol HCO $3 -$	3mol ATP + 4mol NAD(P)H	乙酰辅酶A	厌氧/微氧	[17]
3HP-4HB循环	2mol HCO $3 -$	4mol ATP + 4mol NAD(P)H	乙酰辅酶A	好氧	[18]

图1 天然CO2固定途径RuBisCo—1,5-二磷酸核酮糖羧化酶；Pcs—丙酰辅酶A合成酶；Mcr—丙二酰辅酶A还原酶；Ogs—2-酮戊二酸合酶；Hcd—4-羟基丁酰辅酶A脱水酶；COdh—一氧化碳脱氢酶；Fdh—富马酸脱氢酶；Fmdc—甲酰基呋喃脱氢酶

图2 自养微生物CO2固定的强化策略PR—视紫红质；CdS—硫化镉纳米材料

图3 异养微生物CO2固定的强化策略PEP—磷酸烯醇式丙酮酸；PECP—磷酸烯醇式丙酮酸羧化酶；PRK—磷酸核糖激酶；PFP—聚（芴-共亚苯基）；PDI—苝二酰亚胺衍生物

表2 人工CO2固定途径

CO₂固定途径	固定CO₂/HCO₃^-	消耗能量	固碳效率	氧气需求	参考文献
CETCH循环	1mol CO₂	1mol ATP + 4mol NAD(P)H	5nmol CO₂/(mg 蛋白·min)	好氧	[50]
rGly途径	1mol CO₂	2mol ATP + 4mol NAD(P)H	—	好氧	[1]
POAP循环	2mol CO₂	2mol ATP和1mol NAD(P)H	8.0nmol CO₂/(mg CO₂固定酶·min)	厌氧	[72]
acetyl-CoA双循环	2mol CO₂	消耗2mol乙酰辅酶A并生产3mol	—	厌氧	[73]
rGPS-MCG系统	2mol HCO₃^–	5mol ATP + 5mol NAD(P)H	28.5nmol CO₂/(mg核心蛋白·min)	好氧/厌氧	[74]

图4 人工CO2固定途径NOG—非氧化型糖酵解；GNG—糖异生；POAP—丙酮酸羧化酶/草酰乙酸乙酰水解酶/乙酸辅酶A连接酶/铁氧化还原蛋白氧化还原酶；CETCH—巴豆酰辅酶A/乙基丙二酰辅酶A/羟基丁酰辅酶A；rGPS—还原乙醛酸-丙酮酸合成；MCG—苹果酰辅酶A-甘油酸

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