温度对真眼点藻生长、总脂及二十碳五烯酸（EPA）合成的影响

doi:10.13560/j.cnki.biotech.bull.1985.2021-1293

生物技术通报 ›› 2022, Vol. 38 ›› Issue (6): 261-271.doi: 10.13560/j.cnki.biotech.bull.1985.2021-1293

温度对真眼点藻生长、总脂及二十碳五烯酸（EPA）合成的影响

许瑾¹(), 李涛¹^,²^,³, 李楚琳², 朱顺妮¹, 王忠铭¹(), 向文洲²^,³()

1.中国科学院广州能源研究所广东省新能源和可再生能源研究开发与应用重点实验室中国科学院可再生能源重点实验室,广州 510640
2.中国科学院南海海洋研究所中国科学院热带海洋生物资源与生态重点实验室广东省海洋药物重点实验室,广州 510301
3.南方海洋科学与工程广东省实验室（广州）,广州 511458

收稿日期:2021-10-13 出版日期:2022-06-26 发布日期:2022-07-11
作者简介:许瑾,女,博士,助理研究员,藻类生物技术;E-mail： xujin@ms.giec.ac.cn
基金资助:
广东省新能源和可再生能源研究开发与应用重点实验室开放课题基金(E039kf0301);广东省新能源和可再生能源研究开发与应用重点实验室开放课题基金(E039010301);中国科学院热带海洋生物资源与生态重点实验室开放课题基金(LMB20211006);广东省自然科学基金研究团队项目(2016A030312007);南方海洋科学与工程广东省实验室（广州）人才团队引进重大专项(GML2019ZD0406)

Effects of Temperature on the Growth,Total Lipid and Eicosapentaenoic Acid Synthesis of Eustigmatos sp.

XU Jin¹(), LI Tao¹^,²^,³, LI Chu-lin², ZHU Shun-ni¹, WANG Zhong-ming¹(), XIANG Wen-zhou²^,³()

1. Guangzhou Institute of Energy Conversion,Chinese Academy of Sciences,Guangdong Provincial Key Laboratory of New and Renewable Energy Research and Development,CAS Key Laboratory of Renewable Energy,Guangzhou 510640
2. Key Laboratory of Tropical Marine Bio-resources and Ecology,Guangdong Key Laboratory of Marine Materia Medica,South China Sea Institute of Oceanology,Chinese Academy of Sciences,Guangzhou 510301
3. Southern Marine Science and Engineering Guangdong Laboratory（Guangzhou）,Guangzhou 511458

Received:2021-10-13 Published:2022-06-26 Online:2022-07-11

摘要/Abstract

摘要：

真眼点藻可以积累二十碳五烯酸（EPA）而受到广泛关注,温度是影响多不饱和脂肪酸合成的重要因素。本研究以真眼点藻（Eustigmatos sp. JHsu-01）为材料,设置高温组（30℃）和低温组（15℃）两种培养条件,通过测定生长、脂类积累、脂肪酸组成和甘油酯合成关键基因表达量的变化,探究温度对EPA合成规律的影响。结果表明,低温培养促进了真眼点藻JHsu-01膜脂和EPA的合成,EPA含量最高达到2.78% DW,糖脂是EPA的主要载体,但温度可以改变EPA在糖脂和中性脂之间的分配比例。转录组结果显示,低温条件下,脂肪酸从头合成、三酰甘油（GPAT、plsC、PLPP和DGAT）、糖脂（MGD和DGD）、硫脂（SQD1和SQD2）和ω-3合成途径（Δ5 Des、Δ6 Des和Δ15 Des）中多个关键酶基因表达上调。综上所述,低温可以促进真眼点藻EPA的合成,同时也是一种获得高含量糖脂型EPA的理想培养方式,研究结果为提高真眼点藻EPA产量提供理论和技术依据。

关键词: 多不饱和脂肪酸, 真眼点藻, EPA, 温度, 转录组

Abstract:

Eustigmatos sp. has received great attention due to its high content of eicosapentaenoic acid（EPA）. Temperature is an important factor affecting the synthesis of polyunsaturated fatty acids. In this study,Eustigmatos sp. JHsu-01 was used as the experimental strain and cultured in the 2 conditions,high temperature（30℃）and low temperature（15℃）. The growth,lipid accumulation,fatty acid composition and the expressions of key genes involved in lipid synthesis were measured and the effects of temperature on the EPA synthesis patterns were investigated. The results showed that the low-temperature enhanced the synthesis of membrane lipids and EPA in Eustigmatos sp.,and the highest EPA content was 2.78% DW. Glycolipids were the main carrier of EPA,but temperature changed the distribution of EPA between glycolipids and neutral lipids. The results of transcriptome demonstrated that under low temperature conditions,the expression of key genes in de novo fatty acid synthesis,triacylglycerols（GPAT,plsC,PLPP and DGAT）,glycolipids（MGD and DGD）,sulfolipids（SQD1 and SQD2）and ω-3 synthesis pathways（Δ5 Des,Δ6 Des and Δ15 Des）were up-regulated. In conclusion,low temperature is an important culture condition to promote the synthesis of EPA in Eustigmatos,and it is also an optimal cultivation approach for obtaining high-content glycolipid EPA. The research results may provide a theoretical and technical guidance to increase the production of EPA.

Key words: polyunsaturated fatty acids, Eustigmatos, EPA, temperature, transcriptome

许瑾, 李涛, 李楚琳, 朱顺妮, 王忠铭, 向文洲. 温度对真眼点藻生长、总脂及二十碳五烯酸（EPA）合成的影响[J]. 生物技术通报, 2022, 38(6): 261-271.

XU Jin, LI Tao, LI Chu-lin, ZHU Shun-ni, WANG Zhong-ming, XIANG Wen-zhou. Effects of Temperature on the Growth,Total Lipid and Eicosapentaenoic Acid Synthesis of Eustigmatos sp.[J]. Biotechnology Bulletin, 2022, 38(6): 261-271.

图/表 8

图1 不同温度培养下真眼点藻JHsu-01的细胞形态及生长特性 A：生物量;B：细胞密度;C：30℃条件下第12天细胞形态;D：15℃条件下第12天细胞形态

Fig.1 Cell morphology and growth characteristics of Eustigmatos sp. JHsu-01 cultured at different temperatures A：Biomass. B：Cell density. C：Cell morphology on day 12（30℃）. D：Cell morphology on day 12（15℃）

图2 温度对真眼点藻JHsu-01的PSII的实际光量子产量（A）和PSII的调节性能量耗散的量子产量（B）的影响

Fig.2 Effects of temperature on the effective photochemical quantum yield of PSII（A）and quantum yield of the light-induced non-photochemical fluorescence quenching of PSII（B）of Eustigmatos sp. JHsu-01

图3 温度对真眼点藻JHsu-01总脂含量（A）和脂类分级（B）的影响

Fig.3 Effects of temperature on the total lipid content（A）and lipid classification（B）of Eustigmatos sp. JHsu-01

图4 30℃（A）和15℃（B）培养温度对真眼点藻JHsu-01脂肪酸组成的影响

Fig.4 Effects of 30℃（A）and 15℃（B）on the fatty acid composition of Eustigmatos sp. JHsu-01

图5 EPA在中性脂、糖脂和磷脂中的分布 A：30℃条件下EPA含量;B：15℃条件下EPA含量;C：30℃条件下EPA在不同脂组分中的比例;D：15℃条件下EPA在不同脂组分中的比例

Fig.5 Distribution of EPA in neutral lipids,glycolipids and phospholipids A：EPA content at 30℃. B：EPA content at 15℃. C：The proportion of EPA in different lipid fractions at 30℃. D：The proportion of EPA in different lipid fractions at 15℃

图6 注释基因的统计分析 A：同源物种分布;B：E-value;C：序列覆盖度

Fig.6 Statistical analysis of annotated genes A：Species distribution. B：E-value distribution. C：Similarity distribution

图7 温度对真眼点藻JHsu-01脂肪酸代谢通路的变化 DHAP：磷酸二羟丙酮;G3P：三磷酸甘油;DG：二酰甘油;TG：三酰甘油;PG：磷脂酰甘油;PI：磷脂酰肌醇;PC：磷脂酰胆碱;PE：磷脂酰乙醇胺;PS：磷脂酰丝氨酸;SQDG：异鼠李糖甘油二酯;MGDG：单半乳糖甘油二酯;DGDG：双半乳糖甘油二酯;CDP-DG：胞嘧啶核苷二磷酸甘油二酯;PA：磷酸二酰甘油;LPA：溶血磷脂酸;DHAP：磷酸二羟丙酮（对照组为30℃,处理组为15℃;红色为上调,绿色为下调）

Fig.7 Changes of temperature on the fatty acid metabolic pathways of Eustigmatos sp. JHsu-01 DHAP：dihydroxyacetone phosphate;G3P：glycerol-3-phosphate;DG：diacylglycerol;TG：triacylglycerol;PG：phosphatidylglycerol;PI：phosphatidylinositol;PC：phosphatidylcholine; PE：phosphatidylethanolamine;PS：phosphatidylserine;SQDG：sulfoquinovosyldiacylglycerol; MGDG：monogalactosyldiacylglycerol;DGDG：digalactosyldiacylglycerol;CDP-DG：CDP-diglyceride;PA：diacylglycerol phosphate;LPA：lysophosphatidic acid. DHAP：Dihydroxyacetone phosphate（The control group is 30℃,the treatment group is 15℃;red is up-regulation,and green is down-regulation）

表1 脂肪酸合成相关酶基因的变化情况

Table 1 Expression changes of genes in lipid and fatty acid biosynthesis pathways

	基因功能Gene function	简写Abbreviation	变化情况Changes
脂肪酸合成 Fatty acid synthesis	乙酰辅酶A羧化酶羧基转移酶Acetyl-CoA carboxylase carboxyl transferase	accA	上调 Up
	3-酮酰基-ACP合成酶 3-oxoacyl-ACP synthase	fabF	上调 Up
	3-酮酰基-ACP还原酶 3-oxoacyl-ACP reductase	fabG	上调 Up
	3-酮酰基-ACP脱水酶 3-hydroxyacyl- ACP dehydratase	fabZ	上调 Up
	烯酯酰ACP还原酶I Enoyl-ACP reductase I	fabI	上调 Up
脂肪酸去饱和 Fatty acid desaturation	delta-5脂肪酸去饱和酶 Delta-5 fatty acid desaturase	Δ5 Des	上调 Up
	delta-6脂肪酸去饱和酶 Delta-6 fatty acid desaturase	Δ6 Des	上调 Up
	delta-15脂肪酸去饱和酶 Delta-15 fatty acid desaturase	Δ15 Des	上调 Up
三酰甘油合成 Triacylglycerol synthesis	3-磷酸甘油酰基转移酶 Glycerol-3-phosphate O-acyltransferase	GPAT	上调 Up
	甘油磷酸脱氢酶 Glycerophosphatedehydrogenase	GPDH	无变化 No
	1-酰基甘油-3-磷酸酰基转移酶 1-acyl-glycerol-3-phosphate acyltransferase	plsC	上调 Up
	磷脂酸磷酸酶 Phosphatidate phosphatase	PLPP	上调 Up
	二酰基甘油酰基转移酶 Diacylglycerol O-acyltransferase	DGAT	上调 Up
硫脂合成 Sulfolipid synthesis	硫代异鼠基转移酶 Sulfoquinovosyltransferase	SQD2	上调 Up
硫脂合成 Sulfolipid synthesis	UDP-硫代异鼠糖合酶 UDP-sulfoquinovose synthase	SQD1	上调 Up
糖脂合成 Glycolipid synthesis	二酰基甘油3-β-半乳糖基转移酶 1,2-diacylglycerol 3-beta-galactosyltransferase	MGD	上调 Up
糖脂合成 Glycolipid synthesis	二半乳糖二酰基甘油合酶 Digalactosyldiacylglycerol synthase	DGD	上调 Up
磷脂合成 Phospholipid synthesis	磷脂酰胞苷转移酶 Phosphatidate cytidylyltransferase	CDS1	无变化 No
	乙醇胺磷酸转移酶 Ethanolaminephosphotransferase	EPT1	下调 Down
	二酰基甘油胆碱磷酸转移酶 Diacylglycerol cholinephosphotransferase	CPT1	下调 Down
	磷脂酰丝氨酸合酶2 phosphatidylserine synthase 2	PTDSS2	下调 Down
	CDP-二酰基甘油-甘油-3-磷酸磷脂酰转移酶 CDP-diacylglycerol-glycerol-3-phosphate 3-phosphatidyltransferase	pgsA	无变化 No
	磷脂酰甘油磷酸酶 Phosphatidylglycerophosphatase	GEP4	无变化 No
	CDP-二酰基甘油-肌醇3-磷脂酰转移酶 CDP-diacylglycerol-inositol 3-phosphatidyltransferase	CDIPT	上调 Up

表1 脂肪酸合成相关酶基因的变化情况

Table 1 Expression changes of genes in lipid and fatty acid biosynthesis pathways

	基因功能Gene function	简写Abbreviation	变化情况Changes
脂肪酸合成 Fatty acid synthesis	乙酰辅酶A羧化酶羧基转移酶Acetyl-CoA carboxylase carboxyl transferase	accA	上调 Up
	3-酮酰基-ACP合成酶 3-oxoacyl-ACP synthase	fabF	上调 Up
	3-酮酰基-ACP还原酶 3-oxoacyl-ACP reductase	fabG	上调 Up
	3-酮酰基-ACP脱水酶 3-hydroxyacyl- ACP dehydratase	fabZ	上调 Up
	烯酯酰ACP还原酶I Enoyl-ACP reductase I	fabI	上调 Up
脂肪酸去饱和 Fatty acid desaturation	delta-5脂肪酸去饱和酶 Delta-5 fatty acid desaturase	Δ5 Des	上调 Up
	delta-6脂肪酸去饱和酶 Delta-6 fatty acid desaturase	Δ6 Des	上调 Up
	delta-15脂肪酸去饱和酶 Delta-15 fatty acid desaturase	Δ15 Des	上调 Up
三酰甘油合成 Triacylglycerol synthesis	3-磷酸甘油酰基转移酶 Glycerol-3-phosphate O-acyltransferase	GPAT	上调 Up
	甘油磷酸脱氢酶 Glycerophosphatedehydrogenase	GPDH	无变化 No
	1-酰基甘油-3-磷酸酰基转移酶 1-acyl-glycerol-3-phosphate acyltransferase	plsC	上调 Up
	磷脂酸磷酸酶 Phosphatidate phosphatase	PLPP	上调 Up
	二酰基甘油酰基转移酶 Diacylglycerol O-acyltransferase	DGAT	上调 Up
硫脂合成 Sulfolipid synthesis	硫代异鼠基转移酶 Sulfoquinovosyltransferase	SQD2	上调 Up
硫脂合成 Sulfolipid synthesis	UDP-硫代异鼠糖合酶 UDP-sulfoquinovose synthase	SQD1	上调 Up
糖脂合成 Glycolipid synthesis	二酰基甘油3-β-半乳糖基转移酶 1,2-diacylglycerol 3-beta-galactosyltransferase	MGD	上调 Up
糖脂合成 Glycolipid synthesis	二半乳糖二酰基甘油合酶 Digalactosyldiacylglycerol synthase	DGD	上调 Up
磷脂合成 Phospholipid synthesis	磷脂酰胞苷转移酶 Phosphatidate cytidylyltransferase	CDS1	无变化 No
	乙醇胺磷酸转移酶 Ethanolaminephosphotransferase	EPT1	下调 Down
	二酰基甘油胆碱磷酸转移酶 Diacylglycerol cholinephosphotransferase	CPT1	下调 Down
	磷脂酰丝氨酸合酶2 phosphatidylserine synthase 2	PTDSS2	下调 Down
	CDP-二酰基甘油-甘油-3-磷酸磷脂酰转移酶 CDP-diacylglycerol-glycerol-3-phosphate 3-phosphatidyltransferase	pgsA	无变化 No
	磷脂酰甘油磷酸酶 Phosphatidylglycerophosphatase	GEP4	无变化 No
	CDP-二酰基甘油-肌醇3-磷脂酰转移酶 CDP-diacylglycerol-inositol 3-phosphatidyltransferase	CDIPT	上调 Up

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温度对真眼点藻生长、总脂及二十碳五烯酸（EPA）合成的影响

Effects of Temperature on the Growth,Total Lipid and Eicosapentaenoic Acid Synthesis of Eustigmatos sp.

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