甜菜色素生物合成研究进展

doi:10.19586/j.2095-2341.2025.0171

生物技术进展 ›› 2026, Vol. 16 ›› Issue (3): 510-522.DOI: 10.19586/j.2095-2341.2025.0171

甜菜色素生物合成研究进展

韩田杰¹(), 魏金凤¹(), 燕永亮², 林敏²^,³, 周正富²()

^1.河南大学生命科学学院，河南开封 475004
^2.中国农业科学院生物技术研究所，农业微生物资源发掘与利用全国重点实验室，北京 100081
^3.河南大学农学院，河南开封 475004

收稿日期:2025-12-02 接受日期:2026-01-05 出版日期:2026-05-25 发布日期:2026-07-14
通讯作者: 周正富
作者简介:韩田杰E-mail: hantianjie2023@163.com；
魏金凤E-mail: wjf@henu.edu.cn第一联系人：（韩田杰、魏金凤并列第一作者）
基金资助:
国家重点研发计划项目(2023YFF1000300);国家重点研发计划项目(2023YFF1000304);中国农业科学院重大科技任务项目(CAAS-ZDRW202305)

Advances in the Biosynthesis of Betalains

Tianjie HAN¹(), Jinfeng WEI¹(), Yongliang YAN², Min LIN²^,³, Zhengfu ZHOU²()

^1.School of Life Sciences，Henan University，Henan Kaifeng 475004，China
^2.National Key Laboratory of Agricultural Microbiology，Biotechnology Research Institute，Chinese Academy of Agricultural Sciences，Beijing 100081，China
^3.College of Agriculture，Henan University，Henan Kaifeng 475004，China

Received:2025-12-02 Accepted:2026-01-05 Online:2026-05-25 Published:2026-07-14
Contact: Zhengfu ZHOU

摘要/Abstract

摘要：

甜菜色素是一类天然色素，主要分为红紫色甜菜红素和黄橙色甜菜黄素。因具有优良的着色性能及抗氧化、抗炎、抗肥胖等生物活性，甜菜色素在食品、营养保健品、医药等领域展现出广阔的应用前景。然而，天然提取存在成本高、效率低及产物稳定性差等瓶颈，制约了商业化发展，因而，采用合成生物技术构建微生物细胞工厂异源生产合成甜菜色素已成为研究热点。综述了甜菜色素的化学特性与天然合成途径，重点分析了原核、真核等不同底盘生物的合成策略，系统阐述了关键合成酶功能、代谢路径调控及优化方案，探讨了未来研究方向与应用前景，以期为甜菜色素工业化生产提供新思路。

关键词: 甜菜色素, 甜菜红素, 甜菜黄素, 生物合成, 微生物细胞工厂

Abstract:

Betalains are a class of natural pigments， mainly divided into red-purple betacyanins and yellow-orange betaxanthins. Due to their excellent coloring properties and bioactivities such as antioxidant， anti-inflammatory and anti-obesity effects， betalains show broad application prospects in food， nutraceuticals， medicine and other fields. However， natural extraction faces bottlenecks including high cost， low efficiency and poor product stability， which restrict their commercial development. In contrast， the construction of microbial cell factories via synthetic biotechnology for heterologous and high-yield synthesis of betalains has become a research focus. This article reviewed the chemical properties and natural synthesis pathways of betalains， focused on analyzing the synthesis strategies of different chassis organisms such as prokaryotes and eukaryotes， systematically expounded the functions of key synthases， metabolic pathway regulation and optimization schemes， discussed future research directions and application prospects， and was expected to provide new ideas for the industrial production of betalains.

Key words: betalains, betacyanins, betaxanthins, biosynthesis, microbial cell factories

中图分类号:

Q819

韩田杰, 魏金凤, 燕永亮, 林敏, 周正富. 甜菜色素生物合成研究进展[J]. 生物技术进展, 2026, 16(3): 510-522.

Tianjie HAN, Jinfeng WEI, Yongliang YAN, Min LIN, Zhengfu ZHOU. Advances in the Biosynthesis of Betalains[J]. Current Biotechnology, 2026, 16(3): 510-522.

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链接本文: https://swjsjz.magtechjournal.com/CN/10.19586/j.2095-2341.2025.0171

https://swjsjz.magtechjournal.com/CN/Y2026/V16/I3/510

图/表 6

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类别	底盘微生物	碳源	产量	发酵方式	路径构建与优化策略	参考文献
甜菜苷	酿酒酵母	葡萄糖	17 mg·L^-1	摇瓶	路径构建（外源引入BvCYP76AD1^W13LMjDODA和Mj-cDOPA5GT）	[30]
	酿酒酵母	20 g·L^-1葡萄糖	30.8 mg·L^-1	摇瓶	路径构建（运用组合工程构建酪氨酸羟化酶和多巴双加氧酶文库，筛选最佳组合，引入葡萄糖基转移酶变体）	[31]
	酿酒酵母	10 g·L^-1蔗糖	134.1 mg·L^-1	摇瓶	酶改造（通过设计CYP76AD1增加酶活性），路径重构（加强L-酪氨酸合成途径）和优化发酵条件	[32]
	解脂耶氏酵母	20 g·L^-1葡萄糖	1.27 g·L^-1	5 L发酵罐	路径构建（引入来自七索藤的酪氨酸羟化酶EvTYH、来自紫茉莉的多巴双加氧酶MjDODA和来自甜菜的蔗糖糖基转移酶BvSGT2）并优化发酵条件	[33]
	解脂耶氏酵母	40 g·L^-1葡萄糖	1.34 g·L^-1	250 mL发酵罐	路径构建（组合表达TyH、DODA、GT酶基因），路径重构（增加路径拷贝数；引入酪氨酸、UDP-葡萄糖合成基因与转运蛋白；阻断副产物黑色素合成，敲除4-羟基苯丙酮酸双加氧酶基因，敲除3个β葡萄糖苷酶）	[34]
	威尼斯镰刀菌	40 g·L^-1葡萄糖	1.91 g·L^-1	5 L发酵罐	路径构建（在基因组上整合了RUBY，包含CYP76AD1、DODA、葡萄糖基转移酶基因）	[35]
	马铃薯	/	0.5 mg·g^-1	/	路径构建（利用器官特异性启动子，采用1个基因1个启动子、1个终止子的策略，将BvCYP76AD1、MjDODA、Mj5GT在马铃薯中表达）	[36]
	番茄	/	1.6 mg·g^-1	/	路径构建（利用器官特异性启动子，采用1个基因1个启动子、1个终止子的策略，将BvCYP76AD1、MjDODA、Mj5GT在番茄中表达）	[36]
	家蚕	/	274 μg·mL^-1	/	路径构建（敲除CYP76AD1与DODA的终止密码子，在其开放阅读框末端无缝插入P2A自裂解肽编码序列，把3个酶基因串成“CYP76AD1-P2A-DODA-P2A-糖基转移酶”单转录单元，实现单启动子同步驱动三基因表达）	[37]
组氨酸-甜菜黄素	大肠杆菌	5 g·L^-1葡萄糖	445.84 mg·L^-1	摇瓶	路径构建（通过引入MjDODA、HpaBC实现了甜菜醛氨酸的生成），外源添加组氨酸获得了组氨酸-甜菜黄素	[38]
	大肠杆菌	15 g·L^-1葡萄糖	287.69 mg·L^-1	摇瓶	路径构建（在高产酪氨酸菌株中异源表达HpaBC、MjDODA生产甜菜醛氨酸，与高产组氨酸菌株共培养）	[39]
多巴黄质	大肠杆菌	30 g·L^-1葡萄糖	22.87 g·L^-1	5 L发酵罐	酶改造（通过定向进化左旋多巴4，5-双加氧酶），路径重构（连接阻断3莽草酸合成左旋多巴的途径）	[40]
丙二酰-甜菜苷	解脂耶氏酵母	20 g·L^-1葡萄糖	1.95 g·L^-1	250 mL发酵罐	路径构建（挖掘火龙果和鸡冠花酰基转移酶，筛选鉴定后在高产甜菜苷的解脂耶氏酵母表达）	[41]
苋菜红素	酿酒酵母	20 g·L^-1葡萄糖	10.2 mg·L^-1	摇瓶	路径构建（在高产甜菜苷的酵母中共表达UDP-葡萄糖脱氢酶AtUGD1和葡糖醛酸转移酶GlcATs）	[42]
	解脂耶氏酵母	20 g·L^-1葡萄糖	2.97 g·L^-1	250 mL发酵罐	路径构建（共表达UDP-葡萄糖脱氢酶AtUGD1和葡糖醛酸转移酶GlcATs）	[42]

类别	底盘微生物	碳源	产量	发酵方式	路径构建与优化策略	参考文献
甜菜苷	酿酒酵母	葡萄糖	17 mg·L^-1	摇瓶	路径构建（外源引入BvCYP76AD1^W13LMjDODA和Mj-cDOPA5GT）	[30]
	酿酒酵母	20 g·L^-1葡萄糖	30.8 mg·L^-1	摇瓶	路径构建（运用组合工程构建酪氨酸羟化酶和多巴双加氧酶文库，筛选最佳组合，引入葡萄糖基转移酶变体）	[31]
	酿酒酵母	10 g·L^-1蔗糖	134.1 mg·L^-1	摇瓶	酶改造（通过设计CYP76AD1增加酶活性），路径重构（加强L-酪氨酸合成途径）和优化发酵条件	[32]
	解脂耶氏酵母	20 g·L^-1葡萄糖	1.27 g·L^-1	5 L发酵罐	路径构建（引入来自七索藤的酪氨酸羟化酶EvTYH、来自紫茉莉的多巴双加氧酶MjDODA和来自甜菜的蔗糖糖基转移酶BvSGT2）并优化发酵条件	[33]
	解脂耶氏酵母	40 g·L^-1葡萄糖	1.34 g·L^-1	250 mL发酵罐	路径构建（组合表达TyH、DODA、GT酶基因），路径重构（增加路径拷贝数；引入酪氨酸、UDP-葡萄糖合成基因与转运蛋白；阻断副产物黑色素合成，敲除4-羟基苯丙酮酸双加氧酶基因，敲除3个β葡萄糖苷酶）	[34]
	威尼斯镰刀菌	40 g·L^-1葡萄糖	1.91 g·L^-1	5 L发酵罐	路径构建（在基因组上整合了RUBY，包含CYP76AD1、DODA、葡萄糖基转移酶基因）	[35]
	马铃薯	/	0.5 mg·g^-1	/	路径构建（利用器官特异性启动子，采用1个基因1个启动子、1个终止子的策略，将BvCYP76AD1、MjDODA、Mj5GT在马铃薯中表达）	[36]
	番茄	/	1.6 mg·g^-1	/	路径构建（利用器官特异性启动子，采用1个基因1个启动子、1个终止子的策略，将BvCYP76AD1、MjDODA、Mj5GT在番茄中表达）	[36]
	家蚕	/	274 μg·mL^-1	/	路径构建（敲除CYP76AD1与DODA的终止密码子，在其开放阅读框末端无缝插入P2A自裂解肽编码序列，把3个酶基因串成“CYP76AD1-P2A-DODA-P2A-糖基转移酶”单转录单元，实现单启动子同步驱动三基因表达）	[37]
组氨酸-甜菜黄素	大肠杆菌	5 g·L^-1葡萄糖	445.84 mg·L^-1	摇瓶	路径构建（通过引入MjDODA、HpaBC实现了甜菜醛氨酸的生成），外源添加组氨酸获得了组氨酸-甜菜黄素	[38]
	大肠杆菌	15 g·L^-1葡萄糖	287.69 mg·L^-1	摇瓶	路径构建（在高产酪氨酸菌株中异源表达HpaBC、MjDODA生产甜菜醛氨酸，与高产组氨酸菌株共培养）	[39]
多巴黄质	大肠杆菌	30 g·L^-1葡萄糖	22.87 g·L^-1	5 L发酵罐	酶改造（通过定向进化左旋多巴4，5-双加氧酶），路径重构（连接阻断3莽草酸合成左旋多巴的途径）	[40]
丙二酰-甜菜苷	解脂耶氏酵母	20 g·L^-1葡萄糖	1.95 g·L^-1	250 mL发酵罐	路径构建（挖掘火龙果和鸡冠花酰基转移酶，筛选鉴定后在高产甜菜苷的解脂耶氏酵母表达）	[41]
苋菜红素	酿酒酵母	20 g·L^-1葡萄糖	10.2 mg·L^-1	摇瓶	路径构建（在高产甜菜苷的酵母中共表达UDP-葡萄糖脱氢酶AtUGD1和葡糖醛酸转移酶GlcATs）	[42]
	解脂耶氏酵母	20 g·L^-1葡萄糖	2.97 g·L^-1	250 mL发酵罐	路径构建（共表达UDP-葡萄糖脱氢酶AtUGD1和葡糖醛酸转移酶GlcATs）	[42]

甜菜色素生物合成研究进展

Advances in the Biosynthesis of Betalains

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