Synthetic Biology Journal ›› 2021, Vol. 2 ›› Issue (6): 920-941.DOI: 10.12211/2096-8280.2020-090
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Qingzhuo WANG, Ping SONG, He HUANG
Received:
2020-12-21
Revised:
2021-03-17
Online:
2022-01-21
Published:
2021-12-31
Contact:
He HUANG
汪庆卓, 宋萍, 黄和
通讯作者:
黄和
作者简介:
基金资助:
CLC Number:
Qingzhuo WANG, Ping SONG, He HUANG. Synthetic biotechnology drives the development of natural eukaryotic lipid cell factories[J]. Synthetic Biology Journal, 2021, 2(6): 920-941.
汪庆卓, 宋萍, 黄和. 合成生物技术驱动天然的真核油脂细胞工厂开发[J]. 合成生物学, 2021, 2(6): 920-941.
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URL: https://synbioj.cip.com.cn/EN/10.12211/2096-8280.2020-090
菌株 | 底物 | 油脂产量/(g/L) | 生产强度/[g/(L·h)] | 文献 |
---|---|---|---|---|
Chlorella vulgaris | CO2/葡萄糖 | 41.95 | 0.583 | [ |
Rhodococcus opacus | 乳制品废水 | 2.2 | 0.023 | [ |
Rhodobacter sphaeroides | 乳酸/光照 | — | 0.028 | [ |
Escherichia coli | 葡萄糖 | 3.6 | 0.075 | [ |
E. coli | 葡萄糖 | 7.0 | 0.12 | [ |
E. coli | 葡萄糖 | 21.5 | 0.5 | [ |
Saccharomyces cerevisiae | 葡萄糖 | 10.4 | 0.087 | [ |
S. cerevisiae | 葡萄糖 | 33.4 | 0.17 | [ |
Yarrowia lipolytica | 葡萄糖 | 15.25 | 0.105 | [ |
Y. lipolytica | 葡萄糖 | 98.9 | 1.2 | [ |
Rhodosporidium toruloides | 葡萄糖 | 78.7 | 0.57 | [ |
R. toruloides | 葡萄糖 | 89.4 | 0.62 | [ |
Schizochytrium sp. | 葡萄糖 | 55.3 | 0.46 | [ |
Schizochytrium sp. | 葡萄糖 | 80.14 | 0.53 | [ |
Lipomyces starkeyi | 甜高粱茎汁 | 6.4 | 0.033 | [ |
L. starkeyi | 玉米棒芯水解液 | 8.1 | 0.042 | [ |
Mortierella isabellina | 冷杉木水解液 | 18.55 | 0.086 | [ |
Cryptococcus curvatus | 废纸水解液 | 5.75 | 0.08 | [ |
Trichosporon fermentans | 甘薯藤水解液 | 6.98 | 0.024 | [ |
Trichosporon oleaginosus | 生物柴油副产物 | 21.87 | [ | |
Rhodotorula glutinis | 纯甘油 | 16.28 | 0.1 | [ |
Tab. 1 Research progress of microbial lipid cell factories in recent years
菌株 | 底物 | 油脂产量/(g/L) | 生产强度/[g/(L·h)] | 文献 |
---|---|---|---|---|
Chlorella vulgaris | CO2/葡萄糖 | 41.95 | 0.583 | [ |
Rhodococcus opacus | 乳制品废水 | 2.2 | 0.023 | [ |
Rhodobacter sphaeroides | 乳酸/光照 | — | 0.028 | [ |
Escherichia coli | 葡萄糖 | 3.6 | 0.075 | [ |
E. coli | 葡萄糖 | 7.0 | 0.12 | [ |
E. coli | 葡萄糖 | 21.5 | 0.5 | [ |
Saccharomyces cerevisiae | 葡萄糖 | 10.4 | 0.087 | [ |
S. cerevisiae | 葡萄糖 | 33.4 | 0.17 | [ |
Yarrowia lipolytica | 葡萄糖 | 15.25 | 0.105 | [ |
Y. lipolytica | 葡萄糖 | 98.9 | 1.2 | [ |
Rhodosporidium toruloides | 葡萄糖 | 78.7 | 0.57 | [ |
R. toruloides | 葡萄糖 | 89.4 | 0.62 | [ |
Schizochytrium sp. | 葡萄糖 | 55.3 | 0.46 | [ |
Schizochytrium sp. | 葡萄糖 | 80.14 | 0.53 | [ |
Lipomyces starkeyi | 甜高粱茎汁 | 6.4 | 0.033 | [ |
L. starkeyi | 玉米棒芯水解液 | 8.1 | 0.042 | [ |
Mortierella isabellina | 冷杉木水解液 | 18.55 | 0.086 | [ |
Cryptococcus curvatus | 废纸水解液 | 5.75 | 0.08 | [ |
Trichosporon fermentans | 甘薯藤水解液 | 6.98 | 0.024 | [ |
Trichosporon oleaginosus | 生物柴油副产物 | 21.87 | [ | |
Rhodotorula glutinis | 纯甘油 | 16.28 | 0.1 | [ |
菌株 | 工具 | 原理 | 载体 | 转化方法 | 文献 |
---|---|---|---|---|---|
Yarrowia lipolytica | Golden Gate或Biobrick体外多片段组装; | 体外分子砌块组装,体内同源重组(双交换) | 线性化质粒 | 醋酸锂转化 | [ |
Y. lipolytica | 体内多片段组装 | 同源重组(双交换) | 线性化DNA片段 | 醋酸锂转化 | [ |
Y. lipolytica | Cre-lox重组酶系统 | 位点特异性重组 | 复制型质粒 | 醋酸锂转化 | [ |
Y. lipolytica | 基于rDNA或zeta位点的多拷贝整合 | 同源重组(单交换或双交换) | 自杀质粒或者线性化片段 | 醋酸锂转化 | [ |
Y. lipolytica | 染色体随机整合 | 非同源末端连接 | 线性化片段 | 醋酸锂转化 | [ |
Y. lipolytica | CRISPR/Cas9介导的基因编辑,单基因或多基因敲除、整合 | Cas9切割基因组后,宿主启动同源重组及非同源末端连接修复 | 复制型质粒 | 醋酸锂转化或 电转化 | [ |
Y. lipolytica | CRISPR/dCas9介导的转录抑制 | 失去切割活性的Cas9结合在靶标序列可阻碍转录正常进行 | 复制型质粒 | 醋酸锂转化 | [ |
Y. lipolytica | CRISPR/dCas9介导的转录激活 | 失去切割活性的Cas9结合在启动子区域,招募激活因子后可上调转录 | 复制型质粒 | 醋酸锂转化 | [ |
Y. lipolytica | CRISPR/Cpf1介导的基因编辑 | Cas9切割基因组后,宿主进行非同源末端连接修复 | 复制型质粒 | 醋酸锂转化 | [ |
Y. lipolytica | CRISPR/Cas9辅助的碱基编辑系统 | 胞嘧啶脱氨酶作用后,宿主碱基对由C-G转变为T-A | 复制型质粒 | 醋酸锂转化 | [ |
Rhodosporidium toruloides | CRISPR/Cas9基因编辑,单基因或多基因敲除 | 非同源末端连接 | Ti质粒 | 农杆菌介导转化 | [ |
R. toruloides | CRISPR/Cas9基因编辑,单基因或多基因敲除 | 非同源末端连接 | 自杀质粒 | 醋酸锂化学转化 | [ |
R. toruloides | CRISPR/Cas9基因编辑,单基因或整合,多基因敲除 | 非同源末端连接或同源重组 | Ti质粒 | 农杆菌介导转化 | [ |
R. toruloides | 等位基因替换 | 同源重组(双交换) | Ti质粒 | 农杆菌介导转化 | [ |
R. toruloides | 重组酶系统(Cre-loxP, Flipase-FRT, I-SceI) | 位点特异性重组 | 自杀质粒或Ti质粒 | 农杆菌介导转化或PEG介导原生质体转化 | [ |
R. toruloides | 染色体随机整合 | 非同源末端连接 | Ti质粒 | 农杆菌介导转化 | [ |
R. toruloides | 染色体随机整合 | 非同源末端连接 | 自杀质粒 | PEG介导原生质体转化 | [ |
R. toruloides | 染色体随机整合 | 非同源末端连接 | 线性化DNA片段 | 电转化 | [ |
Schizochytrium sp. | 染色体定点整合 | 同源重组(双交换) | (线性化的)自杀质粒 | 电穿孔转化 | [ |
Schizochytrium sp. TIO1101 | 染色体定点整合 | 同源重组(单交换) | 线性化自杀质粒 | 电穿孔转化 | [ |
Schizochytrium sp. | 染色体随机整合 | 非同源末端连接 | 线性化自杀质粒 | 电穿孔转化 | [ |
Schizochytrium sp. | 染色体随机整合 | 非同源末端连接 | Ti质粒 | 农杆菌介导转化 | [ |
Lipomyces starkeyi | 染色体随机整合 | 非同源末端连接 | Ti质粒 | 农杆菌介导转化 | [ |
L. starkeyi | 染色体定点整合 | 同源重组(双交换) | 线性化的自杀质粒 | PEG介导原生质体转化 | [ |
Tab. 2 Research progress on genetic tools of typical lipid producing fungi
菌株 | 工具 | 原理 | 载体 | 转化方法 | 文献 |
---|---|---|---|---|---|
Yarrowia lipolytica | Golden Gate或Biobrick体外多片段组装; | 体外分子砌块组装,体内同源重组(双交换) | 线性化质粒 | 醋酸锂转化 | [ |
Y. lipolytica | 体内多片段组装 | 同源重组(双交换) | 线性化DNA片段 | 醋酸锂转化 | [ |
Y. lipolytica | Cre-lox重组酶系统 | 位点特异性重组 | 复制型质粒 | 醋酸锂转化 | [ |
Y. lipolytica | 基于rDNA或zeta位点的多拷贝整合 | 同源重组(单交换或双交换) | 自杀质粒或者线性化片段 | 醋酸锂转化 | [ |
Y. lipolytica | 染色体随机整合 | 非同源末端连接 | 线性化片段 | 醋酸锂转化 | [ |
Y. lipolytica | CRISPR/Cas9介导的基因编辑,单基因或多基因敲除、整合 | Cas9切割基因组后,宿主启动同源重组及非同源末端连接修复 | 复制型质粒 | 醋酸锂转化或 电转化 | [ |
Y. lipolytica | CRISPR/dCas9介导的转录抑制 | 失去切割活性的Cas9结合在靶标序列可阻碍转录正常进行 | 复制型质粒 | 醋酸锂转化 | [ |
Y. lipolytica | CRISPR/dCas9介导的转录激活 | 失去切割活性的Cas9结合在启动子区域,招募激活因子后可上调转录 | 复制型质粒 | 醋酸锂转化 | [ |
Y. lipolytica | CRISPR/Cpf1介导的基因编辑 | Cas9切割基因组后,宿主进行非同源末端连接修复 | 复制型质粒 | 醋酸锂转化 | [ |
Y. lipolytica | CRISPR/Cas9辅助的碱基编辑系统 | 胞嘧啶脱氨酶作用后,宿主碱基对由C-G转变为T-A | 复制型质粒 | 醋酸锂转化 | [ |
Rhodosporidium toruloides | CRISPR/Cas9基因编辑,单基因或多基因敲除 | 非同源末端连接 | Ti质粒 | 农杆菌介导转化 | [ |
R. toruloides | CRISPR/Cas9基因编辑,单基因或多基因敲除 | 非同源末端连接 | 自杀质粒 | 醋酸锂化学转化 | [ |
R. toruloides | CRISPR/Cas9基因编辑,单基因或整合,多基因敲除 | 非同源末端连接或同源重组 | Ti质粒 | 农杆菌介导转化 | [ |
R. toruloides | 等位基因替换 | 同源重组(双交换) | Ti质粒 | 农杆菌介导转化 | [ |
R. toruloides | 重组酶系统(Cre-loxP, Flipase-FRT, I-SceI) | 位点特异性重组 | 自杀质粒或Ti质粒 | 农杆菌介导转化或PEG介导原生质体转化 | [ |
R. toruloides | 染色体随机整合 | 非同源末端连接 | Ti质粒 | 农杆菌介导转化 | [ |
R. toruloides | 染色体随机整合 | 非同源末端连接 | 自杀质粒 | PEG介导原生质体转化 | [ |
R. toruloides | 染色体随机整合 | 非同源末端连接 | 线性化DNA片段 | 电转化 | [ |
Schizochytrium sp. | 染色体定点整合 | 同源重组(双交换) | (线性化的)自杀质粒 | 电穿孔转化 | [ |
Schizochytrium sp. TIO1101 | 染色体定点整合 | 同源重组(单交换) | 线性化自杀质粒 | 电穿孔转化 | [ |
Schizochytrium sp. | 染色体随机整合 | 非同源末端连接 | 线性化自杀质粒 | 电穿孔转化 | [ |
Schizochytrium sp. | 染色体随机整合 | 非同源末端连接 | Ti质粒 | 农杆菌介导转化 | [ |
Lipomyces starkeyi | 染色体随机整合 | 非同源末端连接 | Ti质粒 | 农杆菌介导转化 | [ |
L. starkeyi | 染色体定点整合 | 同源重组(双交换) | 线性化的自杀质粒 | PEG介导原生质体转化 | [ |
工程菌 | 宿主 | 遗传操作 | 产物 | 产量 | 底物 | 培养方式 |
---|---|---|---|---|---|---|
S14G19O-ACC1 Y-4311a[ | ATCC 20460 | 敲除甘油-3-磷酸脱氢酶(Gpd1和Gut2)和氧化物酶体基质蛋白Pex10;过表达乙酰辅酶A羧化酶 ACC1 | 游离脂肪酸 | 2.03 g/L | 甘油和 正十二烷 | 摇瓶 |
M-MHY1[ | ACA-DC 50109 | 敲除C2H2锌指蛋白Mhy1 | 三乙酰甘油酯 | 43.1% (DCW) | 葡萄糖 | 摇瓶 |
ylXYL+Obese-XA[ | PO1d | 敲除酰基辅酶A氧化酶(POX1-6)和脂肪酶Tgl4;过表达DAG酰基转移酶DGA1、磷酸转酮酶XPKA、乙酸激酶ACK、乙酰辅酶A合成酶ACS2、磷酸转乙酰酶XA、木糖还原酶XR、木糖醇脱氢酶XDH和木酮糖激酶XK | 三乙酰甘油酯 | 16.5 g/L | 蔗糖 | 发酵罐 |
JMY3580[ | ATCC 20460 | 敲除DAG酰基转移酶(Dga1、Dga2和Lro1)和酰基辅酶A甾醇酰基转移酶;过表达DAG酰基转移酶DGA2 | 脂肪酸 | 9.9 g/L | 甘油 | 摇瓶 |
ADgm-hia[ | Po1g | 过表达乙酰辅酶A羧化酶 ACC1、DAG酰基转移酶DGA1、NAD+激酶YEF和3-磷酸甘油醛脱氢酶GapC | 脂肪酸甲酯 | 98.9 g/L | 葡萄糖 | 发酵罐 |
W29 (Dpex10 ZWF1 ACBP) [ | ATCC 20460 | 敲除氧化物酶体基质蛋白Pex10;过表达葡萄糖-6-磷酸脱氢酶ZWF | 三乙酰甘油酯 | 30% (DCW) | 葡萄糖 | 摇瓶 |
Po1f-6-D[ | PO1f | 过表达来源Mortierella alpina的Δ6去饱和酶 | γ-亚油酸 | 71.6 mg/L | 葡萄糖 | 摇瓶 |
ALDH[ | PO1g | 过表达乙酰辅酶A羧化酶ACC1、DAG酰基转移酶DGA1、醛脱氢酶EcAldH(来源于E. coli)、葡萄糖-6-磷酸脱氢酶ZWF(来源于S. cerevisiae)、谷胱甘肽二硫还原酶GSR和硫氧还蛋白还原酶TRX | 三乙酰甘油酯 | 72.7 g/L | 葡萄糖 | 发酵罐 |
Tafar1-5copy-Δdga1 fao1[ | PO1f | 敲除DAG酰基转移酶Dga1和脂肪醇氧化酶Fao1;过表达脂肪酰辅酶A还原酶Tafar1 | 脂肪醇 | 690.21 mg/L | 甘油 | 发酵罐 |
AD pYLXP-ylFAS1-EcTesA[ | PO1g | 表达脂肪酸合酶FAS1和硫脂酶EcTesA(来源于E. coli) | 游离脂肪酸 | 9.67 g/L | 葡萄糖 | 发酵罐 |
AD pYLXP-AbAtfA-scCat2[ | PO1g | 表达蜡酯合酶AbAtfA(来源于A. baylyi ADP1)和肉碱乙酰转移 酶scCat2 | 脂肪酸乙酯 | 142.5 mg/L | 葡萄糖 | 摇瓶 |
AD pYLXP- Maqu2220-EcfadD[ | PO1g | 表达脂酰辅酶A合成酶Maqu2220和EcfadD(来源于Marinobacter aquaeolei和E. coli) | 脂肪酸醇 | 2.15 g/L | 葡萄糖 | 摇瓶 |
AD pYLXP-MmCAR-BsuSfp-PmADO[ | PO1g | 表达羧酸还原酶MmCAR(来源于M. marinum)、磷酸泛酰巯基乙胺基转移酶BsuSfp(来源于Bacillus subtilis)和醛脱甲氧化酶PmADO(Prochlorococcus marinus) | 脂肪烷烃 | 23.3 mg/L | 葡萄糖 | 摇瓶 |
AD pYLXP-ylACC1-ylDGA1-scCat2[ | PO1g | 表达乙酰辅酶A羧化酶ACC1、DAG酰基转移酶DGA1和肉碱乙酰转移酶Cat2 | 三乙酰甘油酯 | 66.4 g/L | 葡萄糖 | 发酵罐 |
PMOC[ | PO1h | 表达聚羟基脂肪酸合成酶PhaC1 | 聚羟基链烷酸酯 | 1.11 g/L | 正十二烷 | 摇瓶 |
YL-ad9[ | PO1f | 过表达乙酰辅酶A羧化酶 ACC1、DAG酰基转移酶DGA1和硬脂酰辅酶A去饱和酶SCD | 三乙酰甘油酯 | 55 g/L | 葡萄糖 | 发酵罐 |
Y4305[ | ATCC 20362 | 敲除氧化物酶体基质蛋白Pex10、脂肪酶1、固醇载体蛋白scp2和未知功能基因yali0c1871;过表达C16/18延伸酶和Δ12、Δ9、Δ8、Δ5和Δ17去饱和酶 | ω-3 二十碳五烯酸 | 15%(DCW) | 葡萄糖 | 摇瓶 |
Tab. 3 Research progress in metabolic engineering of Yarrowia lipolytica synthesizing lipids and derivatives
工程菌 | 宿主 | 遗传操作 | 产物 | 产量 | 底物 | 培养方式 |
---|---|---|---|---|---|---|
S14G19O-ACC1 Y-4311a[ | ATCC 20460 | 敲除甘油-3-磷酸脱氢酶(Gpd1和Gut2)和氧化物酶体基质蛋白Pex10;过表达乙酰辅酶A羧化酶 ACC1 | 游离脂肪酸 | 2.03 g/L | 甘油和 正十二烷 | 摇瓶 |
M-MHY1[ | ACA-DC 50109 | 敲除C2H2锌指蛋白Mhy1 | 三乙酰甘油酯 | 43.1% (DCW) | 葡萄糖 | 摇瓶 |
ylXYL+Obese-XA[ | PO1d | 敲除酰基辅酶A氧化酶(POX1-6)和脂肪酶Tgl4;过表达DAG酰基转移酶DGA1、磷酸转酮酶XPKA、乙酸激酶ACK、乙酰辅酶A合成酶ACS2、磷酸转乙酰酶XA、木糖还原酶XR、木糖醇脱氢酶XDH和木酮糖激酶XK | 三乙酰甘油酯 | 16.5 g/L | 蔗糖 | 发酵罐 |
JMY3580[ | ATCC 20460 | 敲除DAG酰基转移酶(Dga1、Dga2和Lro1)和酰基辅酶A甾醇酰基转移酶;过表达DAG酰基转移酶DGA2 | 脂肪酸 | 9.9 g/L | 甘油 | 摇瓶 |
ADgm-hia[ | Po1g | 过表达乙酰辅酶A羧化酶 ACC1、DAG酰基转移酶DGA1、NAD+激酶YEF和3-磷酸甘油醛脱氢酶GapC | 脂肪酸甲酯 | 98.9 g/L | 葡萄糖 | 发酵罐 |
W29 (Dpex10 ZWF1 ACBP) [ | ATCC 20460 | 敲除氧化物酶体基质蛋白Pex10;过表达葡萄糖-6-磷酸脱氢酶ZWF | 三乙酰甘油酯 | 30% (DCW) | 葡萄糖 | 摇瓶 |
Po1f-6-D[ | PO1f | 过表达来源Mortierella alpina的Δ6去饱和酶 | γ-亚油酸 | 71.6 mg/L | 葡萄糖 | 摇瓶 |
ALDH[ | PO1g | 过表达乙酰辅酶A羧化酶ACC1、DAG酰基转移酶DGA1、醛脱氢酶EcAldH(来源于E. coli)、葡萄糖-6-磷酸脱氢酶ZWF(来源于S. cerevisiae)、谷胱甘肽二硫还原酶GSR和硫氧还蛋白还原酶TRX | 三乙酰甘油酯 | 72.7 g/L | 葡萄糖 | 发酵罐 |
Tafar1-5copy-Δdga1 fao1[ | PO1f | 敲除DAG酰基转移酶Dga1和脂肪醇氧化酶Fao1;过表达脂肪酰辅酶A还原酶Tafar1 | 脂肪醇 | 690.21 mg/L | 甘油 | 发酵罐 |
AD pYLXP-ylFAS1-EcTesA[ | PO1g | 表达脂肪酸合酶FAS1和硫脂酶EcTesA(来源于E. coli) | 游离脂肪酸 | 9.67 g/L | 葡萄糖 | 发酵罐 |
AD pYLXP-AbAtfA-scCat2[ | PO1g | 表达蜡酯合酶AbAtfA(来源于A. baylyi ADP1)和肉碱乙酰转移 酶scCat2 | 脂肪酸乙酯 | 142.5 mg/L | 葡萄糖 | 摇瓶 |
AD pYLXP- Maqu2220-EcfadD[ | PO1g | 表达脂酰辅酶A合成酶Maqu2220和EcfadD(来源于Marinobacter aquaeolei和E. coli) | 脂肪酸醇 | 2.15 g/L | 葡萄糖 | 摇瓶 |
AD pYLXP-MmCAR-BsuSfp-PmADO[ | PO1g | 表达羧酸还原酶MmCAR(来源于M. marinum)、磷酸泛酰巯基乙胺基转移酶BsuSfp(来源于Bacillus subtilis)和醛脱甲氧化酶PmADO(Prochlorococcus marinus) | 脂肪烷烃 | 23.3 mg/L | 葡萄糖 | 摇瓶 |
AD pYLXP-ylACC1-ylDGA1-scCat2[ | PO1g | 表达乙酰辅酶A羧化酶ACC1、DAG酰基转移酶DGA1和肉碱乙酰转移酶Cat2 | 三乙酰甘油酯 | 66.4 g/L | 葡萄糖 | 发酵罐 |
PMOC[ | PO1h | 表达聚羟基脂肪酸合成酶PhaC1 | 聚羟基链烷酸酯 | 1.11 g/L | 正十二烷 | 摇瓶 |
YL-ad9[ | PO1f | 过表达乙酰辅酶A羧化酶 ACC1、DAG酰基转移酶DGA1和硬脂酰辅酶A去饱和酶SCD | 三乙酰甘油酯 | 55 g/L | 葡萄糖 | 发酵罐 |
Y4305[ | ATCC 20362 | 敲除氧化物酶体基质蛋白Pex10、脂肪酶1、固醇载体蛋白scp2和未知功能基因yali0c1871;过表达C16/18延伸酶和Δ12、Δ9、Δ8、Δ5和Δ17去饱和酶 | ω-3 二十碳五烯酸 | 15%(DCW) | 葡萄糖 | 摇瓶 |
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