从CO2到有机物——碳中和的微藻绿色生物制造

doi:10.12211/2096-8280.2022-023

合成生物学 ›› 2022, Vol. 3 ›› Issue (5): 953-965.DOI: 10.12211/2096-8280.2022-023

从CO₂到有机物——碳中和的微藻绿色生物制造

孙中亮, 陈辉, 王强

河南大学省部共建作物逆境适应与改良国家重点实验室，河南　开封　475004

收稿日期:2022-04-15 修回日期:2022-08-17 出版日期:2022-10-31 发布日期:2022-11-16
通讯作者: 王强
作者简介:孙中亮（1989—），男，博士，副教授。主要从事微藻培养工程和藻基生物产品开发相关研究，包括利用新型光生物反应器高密度培养微藻以及虾青素、多不饱和脂肪酸高附加值产品的工程化制备。E-mail：zlsun@henu.edu.cn
王强（1973—），男，教授，博士生导师，河南大学“杰出人才特区支持计划”特聘教授。主要从事光合成生物学研究，以光合作用研究模式材料蓝细菌和小球藻为研究对象，立足光合作用与合成生物学，聚焦环境问题，围绕微藻环境适应、生物能源与生物减排等科学问题，开展系统性研究。 E-mail：wangqiang@henu.edu.cn
基金资助:
国家重点研发计划(2021YFA0909600);国家自然科学基金(32170138);河南省高校科技创新团队(22IRTSTHN024);河南省自然科学基金(212300410024);河南省科技攻关项目(222102110131);高等学校学科创新引智计划（#D16014）

From CO₂ to value-added products—carbon neutral microalgal green biomanufacturing

Zhongliang SUN, Hui CHEN, Qiang WANG

State Key Laboratory of Crop Stress Adaptation and Improvement，School of Life Sciences，Henan University，Kaifeng 475004，China

Received:2022-04-15 Revised:2022-08-17 Online:2022-10-31 Published:2022-11-16
Contact: Qiang WANG

摘要/Abstract

摘要：

微藻可以利用太阳能固定CO₂并转化为有机物，其作为合成生物物质的细胞工厂具有众多生物学和工程学优点。当前，全球正面临着碳减排和资源短缺的双重压力，通过微藻固碳合成化合物技术的攻关和突破，实现直接利用微藻固定CO₂，有望建立以CO₂为原料、以太阳能为能源，规模化生产大宗食物、能源、化学品和医药保健品的未来新兴绿色生物制造产业，对于解决当前面临的粮食安全、环境污染和能源紧缺等问题具有战略意义。本文从光驱自养的角度，首先总结了微藻作为细胞工厂生产平台化合物、生物能源和高附加值化合物的途径、底盘改造策略等最新进展，进而对该技术的未来发展方向进行展望。最后，提出了微藻作为合成生物学高效底盘细胞，其广泛应用还应该从建立标准化的藻类基因与基因组编辑技术体系、深刻理解合成物质在藻细胞中的代谢流和控制机制以及提高生物量产率和光合作用效率等几个环节进行攻关，以加强微藻绿色生物制造产业的可控性和可复制性。

关键词: 微藻, CO₂, 平台化合物, 生物能源, 高附加值化合物, 合成生物学

Abstract:

Currently, our world is facing the dual pressure of carbon emission reduction and resource shortage. China has also put forward a goal of reaching CO₂ emission peak by 2030 and achieving carbon neutrality by 2060. At present, the production and manufacture of fuels and bulk chemical products mainly rely on petrochemical refining, which is facing the challenges of high risk of production safety, great pressure of environmental protection, and contradiction between supply and demand of oil and gas resources. In this context, the use of microalgae for direct CO₂ fixation is expected to establish large-scale biomanufacturing with CO₂ as raw material and sunlight as energy source, this is a new manufacturing mode that breaks away from the route of petrochemical industry, and has the typical characteristics of low carbon, recyclable, green, and clean. This emerging green industry is of strategic significance for solving the current issues of food security and energy shortages, through sustainable production of food, energy, chemicals, and pharmaceuticals. In addition, microalgae possess great potential in environment protection, thanks to their strong stress resistance, effective remediation of eutrophic elements such as nitrogen and phosphorus from wastewater, and simultaneous removal of SO _x and NO _x during CO₂ utilization in flue gas. Therefore, compared with heterotrophic chassis cells, microalgae-based synthetic biology and bio-manufacturing also play a role in carbon sequestration and emission reduction, and microalgae have then attracted much attention in recent years as “green cell factories”. From the perspective of light-driven autotrophy, we summarize the latest progress of microalgae as a cell factory, introduce chassis transformation strategies, and then look into the future development of this technology. In particular, improved genetic manipulation and larger cultivation scales are critical for microalgae to serve as high efficiency chassis for synthetic biology, whereas promising directions include establishment of standardized systems for algal genome editing, deep understanding of metabolic flux and control for robust biosynthesis, as well as improvement of biomass productivity and photosynthesis efficiency. All in all, this review provides a useful reference to establish controllable and replicable processes for microalgae green biomanufacturing.

Key words: microalgae, carbon dioxide, carbon platform compound, bio-energy, high-value compounds, synthetic biology

中图分类号:

Q816

孙中亮, 陈辉, 王强. 从CO₂到有机物——碳中和的微藻绿色生物制造[J]. 合成生物学, 2022, 3(5): 953-965.

Zhongliang SUN, Hui CHEN, Qiang WANG. From CO₂ to value-added products—carbon neutral microalgal green biomanufacturing[J]. Synthetic Biology Journal, 2022, 3(5): 953-965.

图/表 6

参考文献 103

1	IEA. Global Energy Review 2021 [R/OL]. IEA, 2021. .
2	ESPOSITO R A, MONROE L S, FRIEDMAN J S. Deployment models for commercialized carbon capture and storage[J]. Environmental Science & Technology, 2011, 45(1): 139-146.
3	DETHERIDGE A, HOSKING L J, THOMAS H R, et al. Deep seam and minesoil carbon sequestration potential of the South Wales Coalfield, UK[J]. Journal of Environmental Management, 2019, 248: 109325.
4	SANNA A, STEEL L, MAROTO-VALER M M. Carbon dioxide sequestration using NaHSO₄ and NaOH: a dissolution and carbonation optimisation study[J]. Journal of Environmental Management, 2017, 189: 84-97.
5	SONG Z L, LIU H Y, STRÖMBERG C A E, et al. Phytolith carbon sequestration in global terrestrial biomes[J]. Science of the Total Environment, 2017, 603/604: 502-509.
6	SMITH P. Soil carbon sequestration and biochar as negative emission technologies[J]. Global Change Biology, 2016, 22(3): 1315-1324.
7	WANG Q. Microbial photosynthesis[M]. Singapore: Springer Singapore, 2020.
8	CHEN H, WANG Q. Microalgae-based green bio-manufacturing-how far from us[J]. Frontiers in Microbiology, 2022, 13: 832097.
9	CHEN H, WANG Q. Regulatory mechanisms of lipid biosynthesis in microalgae[J]. Biological Reviews of the Cambridge Philosophical Society, 2021, 96(5): 2373-2391.
10	CHEN H, WANG X Y, WANG Q. Microalgal biofuels in China: The past, progress and prospects[J]. GCB Bioenergy, 2020, 12(12): 1044-1065.
11	NG I S, KESKIN B B, TAN S I. A critical review of genome editing and synthetic biology applications in metabolic engineering of microalgae and cyanobacteria[J]. Biotechnology Journal, 2020, 15(8): e1900228.
12	NADUTHODI M I S, CLAASSENS N J, D'ADAMO S, et al. Synthetic biology approaches to enhance microalgal productivity[J]. Trends in Biotechnology, 2021, 39(10): 1019-1036.
13	POLINER E, FARRÉ E M, BENNING C. Advanced genetic tools enable synthetic biology in the oleaginous microalgae Nannochloropsis sp[J]. Plant Cell Reports, 2018, 37(10): 1383-1399.
14	ZHANG X, CHEN H, CHEN W X, et al. Evaluation of an oil-producing green alga Chlorella sp. C2 for biological DeNO _x of industrial flue gases[J]. Environmental Science & Technology, 2014, 48(17): 10497-10504.
15	ZHENG S Y, CHEN S Y, ZOU S Y, et al. Bioremediation of Pyropia-processing wastewater coupled with lipid production using Chlorella sp[J]. Bioresource Technology, 2021, 321: 124428.
16	PACHAPUR V L, SARMA S J, BRAR S K, et al. Platform chemicals[M]//Platform Chemical Biorefinery. Amsterdam: Elsevier, 2016: 1-20.
17	WERPY T, PETERSEN G. Top Value added chemicals from biomass: volume i—results of screening for potential candidates from sugars and synthesis gas[R]. US Department of Energy, Office of Scientific and Technical Information (OSTI), 2004.
18	FACT.MR. Bio-based Platform Chemicals Market[R/OL]. 2021. .
19	MORALES-DELANUEZ A, HERNÁNDEZ-CASTELLANO L E, MORENO-INDIAS I, et al. Use of glycerol and propylene glycol as additives in heat-treated goat colostrum[J]. Journal of Dairy Science, 2020, 103(3): 2756-2761.
20	TIAN C L, LIU L, XIA M Q, et al. The evaluations of menthol and propylene glycol on the transdermal delivery system of dual drug-loaded lyotropic liquid crystalline gels[J]. AAPS PharmSciTech, 2020, 21(6): 224.
21	FIUME M M, BERGFELD W F, BELSITO D V, et al. Safety assessment of propylene glycol, tripropylene glycol, and PPGs as used in cosmetics[J]. International Journal of Toxicology, 2012, 31(5 ): 245S-260S.
22	LEE C S, AROUA M K, DAUD W M A W, et al. A review: conversion of bioglycerol into 1,3-propanediol via biological and chemical method[J]. Renewable and Sustainable Energy Reviews, 2015, 42: 963-972.
23	HIROKAWA Y, DEMPO Y, FUKUSAKI E, et al. Metabolic engineering for isopropanol production by an engineered cyanobacterium, Synechococcus elongatus PCC 7942, under photosynthetic conditions[J]. Journal of Bioscience and Bioengineering, 2017, 123(1): 39-45.
24	HIROKAWA Y, MATSUO S, HAMADA H, et al. Metabolic engineering of Synechococcus elongatus PCC 7942 for improvement of 1,3-propanediol and glycerol production based on in silico simulation of metabolic flux distribution[J]. Microbial Cell Factories, 2017, 16(1): 212.
25	HIROKAWA Y, MAKI Y, HANAI T. Improvement of 1, 3-propanediol production using an engineered cyanobacterium, Synechococcus elongatus by optimization of the gene expression level of a synthetic metabolic pathway and production conditions[J]. Metabolic Engineering, 2017, 39: 192-199.
26	LIU H Y, NI J, XU P, et al. Enhancing light-driven 1, 3-propanediol production by using natural compartmentalization of differentiated cells[J]. ACS Synthetic Biology, 2018, 7(10): 2436-2446.
27	LI H, LIAO J C. Engineering a cyanobacterium as the catalyst for the photosynthetic conversion of CO₂ to 1,2-propanediol[J]. Microbial Cell Factories, 2013, 12(1): 4.
28	DAVID C, SCHMID A, ADRIAN L, et al. Production of 1, 2-propanediol in photoautotrophic Synechocystis is linked to glycogen turn-over[J]. Biotechnology and Bioengineering, 2018, 115(2): 300-311.
29	MATOS C T, GOUVEIA L, MORAIS A R C, et al. Green metrics evaluation of isoprene production by microalgae and bacteria[J]. Green Chem, 2013, 15(10): 2854-2864.
30	GAO X, GAO F, LIU D, et al. Engineering the methylerythritol phosphate pathway in cyanobacteria for photosynthetic isoprene production from CO₂ [J]. Energy & Environmental Science, 2016, 9(4): 1400-1411.
31	CHAVES J E, MELIS A. Biotechnology of cyanobacterial isoprene production[J]. Applied Microbiology and Biotechnology, 2018, 102(15): 6451-6458.
32	PADE N, ERDMANN S, ENKE H K, et al. Insights into isoprene production using the cyanobacterium Synechocystis sp. PCC 6803[J]. Biotechnology for Biofuels, 2016, 9: 89.
33	WANG Y, TAO F, NI J, et al. Production of C₃ platform chemicals from CO₂ by genetically engineered cyanobacteria[J]. Green Chemistry, 2015, 17(5): 3100-3110.
34	HIROKAWA Y, MAKI Y, TATSUKE T, et al. Cyanobacterial production of 1,3-propanediol directly from carbon dioxide using a synthetic metabolic pathway[J]. Metabolic Engineering, 2016, 34: 97-103.
35	CHAVES J E, RUEDA-ROMERO P, KIRST H, et al. Engineering isoprene synthase expression and activity in cyanobacteria[J]. ACS Synthetic Biology, 2017, 6(12): 2281-2292.
36	VEETIL V P, ANGERMAYR S A, HELLINGWERF K J. Ethylene production with engineered Synechocystis sp PCC 6803 strains[J]. Microbial Cell Factories, 2017, 16(1): 34.
37	DURALL C, LINDBERG P, YU J P, et al. Increased ethylene production by overexpressing phosphoenolpyruvate carboxylase in the cyanobacterium Synechocystis PCC 6803[J]. Biotechnology for Biofuels, 2020, 13: 16.
38	WANG Y P, CHEN L, ZHANG W W. Proteomic and metabolomic analyses reveal metabolic responses to 3-hydroxypropionic acid synthesized internally in cyanobacterium Synechocystis sp. PCC 6803[J]. Biotechnology for Biofuels, 2016, 9: 209.
39	LAN E I, CHUANG D S, SHEN C R, et al. Metabolic engineering of cyanobacteria for photosynthetic 3-hydroxypropionic acid production from CO₂ using Synechococcus elongatus PCC 7942[J]. Metabolic Engineering, 2015, 31: 163-170.
40	KANNO M, ATSUMI S. Engineering an obligate photoautotrophic cyanobacterium to utilize glycerol for growth and chemical production[J]. ACS Synthetic Biology, 2017, 6(1): 69-75.
41	MCEWEN J T, KANNO M, ATSUMI S. 2, 3 Butanediol production in an obligate photoautotrophic cyanobacterium in dark conditions via diverse sugar consumption[J]. Metabolic Engineering, 2016, 36: 28-36.
42	NOZZI N E, CASE A E, CARROLL A L, et al. Systematic approaches to efficiently produce 2,3-butanediol in a marine cyanobacterium[J]. ACS Synthetic Biology, 2017, 6(11): 2136-2144.
43	CHOWDHURY H, LOGANATHAN B. Third-generation biofuels from microalgae: a review[J]. Current Opinion in Green and Sustainable Chemistry, 2019, 20: 39-44.
44	ARUN J, VARSHINI P, PRITHVINATH P K, et al. Enrichment of bio-oil after hydrothermal liquefaction (HTL) of microalgae C. vulgaris grown in wastewater: bio-char and post HTL wastewater utilization studies[J]. Bioresource Technology, 2018, 261: 182-187.
45	XU S N, ELSAYED M, ISMAIL G A, et al. Evaluation of bioethanol and biodiesel production from Scenedesmus obliquus grown in biodiesel waste glycerol: a sequential integrated route for enhanced energy recovery[J]. Energy Conversion and Management, 2019, 197: 111907.
46	CHISTI Y. Biodiesel from microalgae[J]. Biotechnology Advances, 2007, 25(3): 294-306.
47	RAN W Y, WANG H T, LIU Y H, et al. Storage of starch and lipids in microalgae: biosynthesis and manipulation by nutrients[J]. Bioresource Technology, 2019, 291: 121894.
48	SINGH P, KUMARI S, GULDHE A, et al. Trends and novel strategies for enhancing lipid accumulation and quality in microalgae[J]. Renewable and Sustainable Energy Reviews, 2016, 55: 1-16.
49	GUARNIERI M T, PIENKOS P T. Algal omics: unlocking bioproduct diversity in algae cell factories[J]. Photosynthesis Research, 2015, 123(3): 255-263.
50	XUE J, NIU Y F, HUANG T, et al. Genetic improvement of the microalga Phaeodactylum tricornutum for boosting neutral lipid accumulation[J]. Metabolic Engineering, 2015, 27: 1-9.
51	ISKANDAROV U, SITNIK S, SHTAIDA N, et al. Cloning and characterization of a GPAT-like gene from the microalga Lobosphaera incisa (Trebouxiophyceae): overexpression in Chlamydomonas reinhardtii enhances TAG production[J]. Journal of Applied Phycology, 2016, 28(2): 907-919.
52	TRENTACOSTE E M, SHRESTHA R P, SMITH S R, et al. Metabolic engineering of lipid catabolism increases microalgal lipid accumulation without compromising growth[J]. Proceedings of the National Academy of Sciences of the United States of America, 2013, 110(49): 19748-19753.
53	NIU Y F, ZHANG M H, LI D W, et al. Improvement of neutral lipid and polyunsaturated fatty acid biosynthesis by overexpressing a type 2 diacylglycerol acyltransferase in marine diatom Phaeodactylum tricornutum [J]. Marine Drugs, 2013, 11(11): 4558-4569.
54	FAN J H, NING K, ZENG X W, et al. Genomic foundation of starch-to-lipid switch in oleaginous Chlorella spp[J]. Plant Physiology, 2015, 169(4): 2444-2461.
55	LI D W, BALAMURUGAN S, YANG Y F, et al. Transcriptional regulation of microalgae for concurrent lipid overproduction and secretion[J]. Science Advances, 2019, 5(1): eaau3795.
56	SHIN Y S, JEONG J, NGUYEN T H T, et al. Targeted knockout of phospholipase A2 to increase lipid productivity in Chlamydomonas reinhardtii for biodiesel production[J]. Bioresource Technology, 2019, 271: 368-374.
57	RYU A J, KANG N K, JEON S, et al. Development and characterization of a Nannochloropsis mutant with simultaneously enhanced growth and lipid production[J]. Biotechnology for Biofuels, 2020, 13: 38.
58	SZPYRKA E, BRODA D, OKLEJEWICZ B, et al. A non-vector approach to increase lipid levels in the microalga Planktochlorella nurekis [J]. Molecules, 2020, 25(2): 270.
59	GOSWAMI R K, MEHARIYA S, OBULISAMY P K, et al. Advanced microalgae-based renewable biohydrogen production systems: a review[J]. Bioresource Technology, 2021, 320: 124301.
60	CHEN C Y, CHANG H Y, CHANG J S. Producing carbohydrate-rich microalgal biomass grown under mixotrophic conditions as feedstock for biohydrogen production[J]. International Journal of Hydrogen Energy, 2016, 41(7): 4413-4420.
61	LAKATOS G, BALOGH D, FARKAS A, et al. Factors influencing algal photobiohydrogen production in algal-bacterial co-cultures[J]. Algal Research, 2017, 28: 161-171.
62	WANG Y T, JIANG X Q, HU C X, et al. Optogenetic regulation of artificial microRNA improves H₂ production in green alga Chlamydomonas reinhardtii [J]. Biotechnology for Biofuels, 2017, 10: 257.
63	XU F Q, MA W M, ZHU X G. Introducing pyruvate oxidase into the chloroplast of Chlamydomonas reinhardtii increases oxygen consumption and promotes hydrogen production[J]. International Journal of Hydrogen Energy, 2011, 36(17): 10648-10654.
64	LIN H D, LIU B H, KUO T T, et al. Knockdown of PsbO leads to induction of HydA and production of photobiological H₂ in the green alga Chlorella sp. DT[J]. Bioresource Technology, 2013, 143: 154-162.
65	KOSOUROV S N, GHIRARDI M L, SEIBERT M. A truncated antenna mutant of Chlamydomonas reinhardtii can produce more hydrogen than the parental strain[J]. International Journal of Hydrogen Energy, 2011, 36(3): 2044-2048.
66	BURROWS E H, CHAPLEN F W R, ELY R L. Effects of selected electron transport chain inhibitors on 24-h hydrogen production by Synechocystis sp. PCC 6803[J]. Bioresource Technology, 2011, 102(3): 3062-3070.
67	LI H, LIU Y M, WANG Y T, et al. Improved photobio-H₂ production regulated by artificial miRNA targeting psbA in green microalga Chlamydomonas reinhardtii [J]. Biotechnology for Biofuels, 2018, 11: 36.
68	YANG D W, SYN J W, HSIEH C H, et al. Genetically engineered hydrogenases promote biophotocatalysis-mediated H₂ production in the green alga Chlorella sp. DT[J]. International Journal of Hydrogen Energy, 2019, 44(5): 2533-2545.
69	YACOBY I, POCHEKAILOV S, TOPORIK H, et al. Photosynthetic electron partitioning between [FeFe]-hydrogenase and ferredoxin: NADP⁺-oxidoreductase (FNR) enzymes in vitro [J]. Proceedings of the National Academy of Sciences of the United States of America, 2011, 108(23): 9396-9401.
70	LOSH J L, YOUNG J N, MOREL F M M. Rubisco is a small fraction of total protein in marine phytoplankton[J]. The New Phytologist, 2013, 198(1): 52-58.
71	HU G R, FAN Y, ZHENG Y L, et al. Photoprotection capacity of microalgae improved by regulating the antenna size of light-harvesting complexes[J]. Journal of Applied Phycology, 2020, 32(2): 1027-1039.
72	LI T P, JIANG Q Y, HUANG J F, et al. Reprogramming bacterial protein organelles as a nanoreactor for hydrogen production[J]. Nature Communications, 2020, 11: 5448.
73	JOHN R P, ANISHA G S, NAMPOOTHIRI K M, et al. Micro and macroalgal biomass: a renewable source for bioethanol[J]. Bioresource Technology, 2011, 102(1): 186-193.
74	LUAN G D, QI Y J, WANG M, et al. Combinatory strategy for characterizing and understanding the ethanol synthesis pathway in cyanobacteria cell factories[J]. Biotechnology for Biofuels, 2015, 8: 184.
75	CHOCHOIS V, DAUVILLÉE D, BEYLY A, et al. Hydrogen production in Chlamydomonas: photosystem II-dependent and-independent pathways differ in their requirement for starch metabolism[J]. Plant Physiology, 2009, 151(2): 631-640.
76	HE M, WU B, QIN H, et al. Zymomonas mobilis: a novel platform for future biorefineries[J]. Biotechnology for Biofuels, 2014, 7(1): 101.
77	GAO Z X, ZHAO H, LI Z M, et al. Photosynthetic production of ethanol from carbon dioxide in genetically engineered cyanobacteria[J]. Energy Environmental Science, 2012, 5(12): 9857-9865.
78	REPPAS N. Metabolic switch: US20120164705A1[P]. 2012.
79	DEXTER J, ARMSHAW P, SHEAHAN C, et al. The state of autotrophic ethanol production in Cyanobacteria[J]. Journal of Applied Microbiology, 2015, 119(1): 11-24.
80	BILAL M, RASHEED T, AHMED I, et al. High-value compounds from microalgae with industrial exploitability-a review[J]. Frontiers in Bioscience (Scholar Edition), 2017, 9(3): 319-342.
81	GAO E B, KYERE-YEBOAH K, WU J H, et al. Photoautotrophic production of p-coumaric acid using genetically engineered Synechocystis sp. Pasteur Culture Collection 6803[J]. Algal Research, 2021, 54: 102180.
82	LEHMANN M, VAMVAKA E, TORRADO A, et al. Introduction of the carotenoid biosynthesis α-branch into Synechocystis sp. PCC 6803 for lutein production[J]. Frontiers in Plant Science, 2021, 12: 699424.
83	LIN P C, ZHANG F Z, PAKRASI H B. Enhanced limonene production in a fast-growing cyanobacterium through combinatorial metabolic engineering[J]. Metabolic Engineering Communications, 2021, 12: e00164.
84	VARMAN A M, YU Y, YOU L, et al. Photoautotrophic production of D-lactic acid in an engineered cyanobacterium[J]. Microbial Cell Factories, 2013, 12: 117.
85	HASUNUMA T, TAKAKI A, MATSUDA M, et al. Single-stage astaxanthin production enhances the nonmevalonate pathway and photosynthetic central metabolism in Synechococcus sp. PCC 7002[J]. ACS Synthetic Biology, 2019, 8(12): 2701-2709.
86	HALFMANN C, GU L P, GIBBONS W, et al. Genetically engineering cyanobacteria to convert CO₂, water, and light into the long-chain hydrocarbon farnesene[J]. Applied Microbiology and Biotechnology, 2014, 98(23): 9869-9877.
87	WANG X, LIU W, XIN C P, et al. Enhanced limonene production in cyanobacteria reveals photosynthesis limitations[J]. Proceedings of the National Academy of Sciences of the United States of America, 2016, 113(50): 14225-14230.
88	LAUERSEN K J. Eukaryotic microalgae as hosts for light-driven heterologous isoprenoid production[J]. Planta, 2019, 249(1): 155-180.
89	POURMIR A, NOOR-MOHAMMADI S, JOHANNES T W. Production of xylitol by recombinant microalgae[J]. Journal of Biotechnology, 2013, 165(3/4): 178-183.
90	LAUERSEN K J, BAIER T, WICHMANN J, et al. Efficient phototrophic production of a high-value sesquiterpenoid from the eukaryotic microalga Chlamydomonas reinhardtii [J]. Metabolic Engineering, 2016, 38: 331-343.
91	HAMILTON M L, HASLAM R P, NAPIER J A, et al. Metabolic engineering of Phaeodactylum tricornutum for the enhanced accumulation of omega-3 long chain polyunsaturated fatty acids[J]. Metabolic Engineering, 2014, 22: 3-9.
92	TANAKA T, MAEDA Y, SUHAIMI N, et al. Intron-mediated enhancement of transgene expression in the oleaginous diatom Fistulifera solaris towards bisabolene production[J]. Algal Research, 2021, 57: 102345.
93	WEI H H, SHI Y, MA X N, et al. A type-I diacylglycerol acyltransferase modulates triacylglycerol biosynthesis and fatty acid composition in the oleaginous microalga, Nannochloropsis oceanica [J]. Biotechnology for Biofuels, 2017, 10: 174.
94	EILERS U, BIKOULIS A, BREITENBACH J, et al. Limitations in the biosynthesis of fucoxanthin as targets for genetic engineering in Phaeodactylum tricornutum [J]. Journal of Applied Phycology, 2016, 28(1): 123-129.
95	LANH P T, NGUYEN H M, DUONG B T T, et al. Generation of microalga Chlamydomonas reinhardtii expressing VP28 protein as oral vaccine candidate for shrimps against White Spot Syndrome Virus (WSSV) infection[J]. Aquaculture, 2021, 540: 736737.
96	GREGORY J A, TOPOL A B, DOERNER D Z, et al. Alga-produced cholera toxin-Pfs25 fusion proteins as oral vaccines[J]. Applied and Environmental Microbiology, 2013, 79(13): 3917-3925.
97	TRAN M, HENRY R E, SIEFKER D, et al. Production of anti-cancer immunotoxins in algae: ribosome inactivating proteins as fusion partners[J]. Biotechnology and Bioengineering, 2013, 110(11): 2826-2835.
98	TRAN M, VAN C, BARRERA D J, et al. Production of unique immunotoxin cancer therapeutics in algal chloroplasts[J]. Proceedings of the National Academy of Sciences of the United States of America, 2013, 110(1): E15-E22.
99	HU L N, FENG S Y, LIANG G F, et al. CRISPR/Cas9-induced β-carotene hydroxylase mutation in Dunaliella salina CCAP19/18[J]. AMB Express, 2021, 11(1): 83.
100	BECKER I, PRASAD B, NTEFIDOU M, et al. Agrobacterium tumefaciens-mediated nuclear transformation of a biotechnologically important microalga-euglena gracilis[J]. International Journal of Molecular Sciences, 2021, 22(12): 6299.
101	KIM J, CHANG K S, LEE S, et al. Establishment of a genome editing tool using CRISPR-Cas9 in Chlorella vulgaris UTEX395[J]. International Journal of Molecular Sciences, 2021, 22(2): 480.
102	LIN W R, TAN S I, HSIANG C C, et al. Challenges and opportunity of recent genome editing and multi-omics in cyanobacteria and microalgae for biorefinery[J]. Bioresource Technology, 2019, 291: 121932.
103	PERIN G, BELLAN A, BERNARDI A, et al. The potential of quantitative models to improve microalgae photosynthetic efficiency[J]. Physiologia Plantarum, 2019, 166(1): 380-391.

表达产物	合成聚合物	表达株系	最终产率/[mg/(L·d)]	参考文献
1，3-丙二醇	聚酯、聚醚、聚氨酯、PTT	Synechococcus sp. PCC 7942	61	[25]
		Synechococcus sp. PCC 7942	8	[33]
		Synechococcus sp. PCC 7942	20.6	[34]
		Synechococcus sp. PCC 7120	2.3	[26]
1，2-丙二醇	聚丙二醇	Synechococcus sp. PCC 7942	15	[27]
		Synechocystis sp. PCC 6803	100	[28]
异戊二烯	橡胶	Synechocystis sp. PCC 6803	60	[30]
		Synechocystis sp. PCC 6803	1.7	[35]
乙烯	聚乙烯、聚苯乙烯、PVC、聚酯	Synechocystis sp. PCC 6803	2104	[36]
		Synechocystis sp. PCC 6803	19.2	[37]
3-羟基丙酸	聚3-羟基丙酸	Synechocystis sp. PCC 6803	139.5	[38]
		Synechococcus sp. PCC 7942	329.5	[39]
2，3-丁二醇	树脂	Synechococcus sp. PCC 7942	54.36	[40]
		Synechococcus sp. PCC 7942	300	[41]
		Synechococcus sp. PCC 7942	110	[40]
		Synechococcus sp. PCC 7942	100	[42]

表达产物	合成聚合物	表达株系	最终产率/[mg/(L·d)]	参考文献
1，3-丙二醇	聚酯、聚醚、聚氨酯、PTT	Synechococcus sp. PCC 7942	61	[25]
		Synechococcus sp. PCC 7942	8	[33]
		Synechococcus sp. PCC 7942	20.6	[34]
		Synechococcus sp. PCC 7120	2.3	[26]
1，2-丙二醇	聚丙二醇	Synechococcus sp. PCC 7942	15	[27]
		Synechocystis sp. PCC 6803	100	[28]
异戊二烯	橡胶	Synechocystis sp. PCC 6803	60	[30]
		Synechocystis sp. PCC 6803	1.7	[35]
乙烯	聚乙烯、聚苯乙烯、PVC、聚酯	Synechocystis sp. PCC 6803	2104	[36]
		Synechocystis sp. PCC 6803	19.2	[37]
3-羟基丙酸	聚3-羟基丙酸	Synechocystis sp. PCC 6803	139.5	[38]
		Synechococcus sp. PCC 7942	329.5	[39]
2，3-丁二醇	树脂	Synechococcus sp. PCC 7942	54.36	[40]
		Synechococcus sp. PCC 7942	300	[41]
		Synechococcus sp. PCC 7942	110	[40]
		Synechococcus sp. PCC 7942	100	[42]

藻株	操作方法	有益结果	油脂含量或产率	参考文献
Phaeodactylum tricornutum	过表达内源性苹果酸脱氢酶	油脂含量增加250%	57.8%	[50]
Chlamydomonas reinhardtii	Lobosphaera incise中的GPAT基因（LiGPAT）在莱茵衣藻中过表达	TAG含量比野生型增加50%	50.0%	[51]
Thalassiosira pseudonana	转化反义结构体，阻止脂质分解代谢	油脂含量比野生型增加230%	18.8%	[52]
Phaeodactylum tricornutum	过表达Ⅱ型甘油二酯酰基转移酶（DGAT2D）基因	TAG含量增加35%	37.2%	[53]
Chlorella spp	拟南芥激酶的上调和过表达	油脂产量增加110.4%	27.5%	[54]
Nannochloropsis	过表达新型bZIP1转录因子NobZIP1N	油脂含量增加而不影响微藻生长	40%	[55]
Chlamydomonas reinhardtii	使用CRISPR-Cas9敲除磷脂酶A2基因	油脂含量可达64.25%	80.92 mg/(L·d)	[56]
Chlamydomonas reinhardtii	使用CRISPR-Cas9 RNP方法产生葡萄糖焦磷酸化酶基因（AGP）突变的突变体	油脂含量比野生型增加274%	57.67%	[56]
Nannochloropsis	通过插入突变产生突变体Mut68	脂肪酸甲酯含量和产量分别比野生型增加34%和75%	78.3 mg/(L·d)	[57]
Planktochlorella nurekis	利用细胞松弛素B和秋水仙碱调控DNA水平	油脂含量比野生型增加10%～60%	12%～26%	[58]

藻株	操作方法	有益结果	油脂含量或产率	参考文献
Phaeodactylum tricornutum	过表达内源性苹果酸脱氢酶	油脂含量增加250%	57.8%	[50]
Chlamydomonas reinhardtii	Lobosphaera incise中的GPAT基因（LiGPAT）在莱茵衣藻中过表达	TAG含量比野生型增加50%	50.0%	[51]
Thalassiosira pseudonana	转化反义结构体，阻止脂质分解代谢	油脂含量比野生型增加230%	18.8%	[52]
Phaeodactylum tricornutum	过表达Ⅱ型甘油二酯酰基转移酶（DGAT2D）基因	TAG含量增加35%	37.2%	[53]
Chlorella spp	拟南芥激酶的上调和过表达	油脂产量增加110.4%	27.5%	[54]
Nannochloropsis	过表达新型bZIP1转录因子NobZIP1N	油脂含量增加而不影响微藻生长	40%	[55]
Chlamydomonas reinhardtii	使用CRISPR-Cas9敲除磷脂酶A2基因	油脂含量可达64.25%	80.92 mg/(L·d)	[56]
Chlamydomonas reinhardtii	使用CRISPR-Cas9 RNP方法产生葡萄糖焦磷酸化酶基因（AGP）突变的突变体	油脂含量比野生型增加274%	57.67%	[56]
Nannochloropsis	通过插入突变产生突变体Mut68	脂肪酸甲酯含量和产量分别比野生型增加34%和75%	78.3 mg/(L·d)	[57]
Planktochlorella nurekis	利用细胞松弛素B和秋水仙碱调控DNA水平	油脂含量比野生型增加10%～60%	12%～26%	[58]

藻株	操作方法	有益结果	氢气产量或产率	参考文献
Chlamydomonas reinhardtii	设计光诱导系统，利用该系统设计蓝光诱导产氢转基因藻类	成功激活了人工miRNA，提高微藻的产氢能力	20 μL/(L·h)	[62]
Chlamydomonas reinhardtii	将大肠杆菌的丙酮酸氧化酶和过氧化氢酶基因整合到衣藻叶绿体基因组中	生物氢产量增加了2倍	1.04 μmol/(L·h)	[63]
Chlorella sp. DT.	敲除psbO基因	生物氢产量比野生型提高了9倍	350 mL/L	[64]
Chlamydomonas reinhardtii	截短捕光天线	氢气产量比野生型增加6倍	30 mL/L	[65]
Synechocystis sp. PCC 6803	添加抑制光合和呼吸作用电子传递链的抑制剂	氢气产量增加了30倍	1.25 μmol/(L·h)	[66]
Chlamydomonas reinhardtii	热诱导人工miRNA表达系统	氢气合成增加60%	90 μL/mg Chl	[67]

从CO₂到有机物——碳中和的微藻绿色生物制造

From CO₂ to value-added products—carbon neutral microalgal green biomanufacturing

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分类	藻株	合成产物	参考文献
原核藻	Synechocystis sp. PCC 6803	对香豆酸	[81]
	Synechocystis sp. PCC 6803	叶黄素	[82]
	Synechococcus elongatus UTEX 2973	柠檬烯	[83]
	Synechocystis sp. PCC 6803	乳酸	[84]
	Synechococcus sp. PCC 7002	虾青素	[85]
	Anabaena sp. PCC7120	法尼烯	[86]
	Synechococcus sp. PCC 7942	柠檬烯	[87]
真核藻	Chlamydomonas reinhardtii	类异戊二烯	[88]
	Chlamydomonas reinhardtii	木糖醇	[89]
	Chlamydomonas reinhardtii	倍半萜	[90]
	Phaeodactylum tricornutum	DHA	[91]
	Bacillariophyta	没药烯	[92]
	Nannochloropsis	EPA	[93]
	Phaeodactylum tricornutum	岩藻黄质	[94]

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