淫羊藿类黄酮生物合成相关基因研究进展

doi:10.11983/CBB23133

植物学报 ›› 2024, Vol. 59 ›› Issue (5): 834-846.DOI: 10.11983/CBB23133 cstr: 32102.14.CBB23133

淫羊藿类黄酮生物合成相关基因研究进展

范雪兰¹^,²^,^†, 落艳娇¹^,³^,^†, 徐超群¹^,^*(), 郭宝林¹^,^*()

¹中国医学科学院北京协和医学院药用植物研究所, 中草药物质基础与资源利用教育部重点实验室, 北京 100193
²江西中医药大学, 南昌 330000
³中国中医科学院中医药健康产业研究所, 南昌 330000

收稿日期:2023-09-22 接受日期:2024-03-18 出版日期:2024-09-10 发布日期:2024-08-19
通讯作者: 徐超群,郭宝林
作者简介:第一联系人：† 共同第一作者
基金资助:
中国医学科学院医学与健康科技创新工程(2021-I2M-1-031)

Research Progress on Genes Related to Flavonoids Biosynthesis in Herba Epimedii

Xuelan Fan¹^,²^,^†, Yanjiao Luo¹^,³^,^†, Chaoqun Xu¹^,^*(), Baolin Guo¹^,^*()

¹Key Laboratory of Bioactive Substances and Resources Utilization of Chinese Herbal Medicines, Ministry of Education, Institute of Medicinal Plant Development, Peking Union Medical College & Chinese Academy of Medical Sciences, Beijing 100193, China
²Jiangxi University of Chinese Medicine, Nanchang 330000, China
³Institute of Traditional Chinese Medicine Health Industry, China Academy of Chinese Medical Sciences, Nanchang 330000, China

Received:2023-09-22 Accepted:2024-03-18 Online:2024-09-10 Published:2024-08-19
Contact: Chaoqun Xu, Baolin Guo
About author:First author contact:† These authors contributed equally to this paper

摘要/Abstract

摘要： 淫羊藿(Herba Epimedii)是一种历史悠久的中草药(TCM), 药用价值高, 国内淫羊藿相关研究备受关注。黄酮醇苷类(PFGs)成分是淫羊藿的主要活性物质, 其含量决定了药用品质。了解淫羊藿类黄酮生物合成途径, 挖掘与淫羊藿黄酮醇苷类含量相关的基因, 阐明其生物合成途径的调控机制对于提高淫羊藿品质至关重要。该文系统综述了淫羊藿类黄酮生物合成相关结构基因和转录因子基因研究进展, 为揭示黄酮含量的分子调控机制, 进而为淫羊藿分子育种和黄酮醇苷的合成生物学研究奠定理论基础。

关键词: 淫羊藿, 黄酮醇苷, 生物合成, 结构基因, 调控基因

Abstract: Herba Epimedii is a traditional Chinese herb medicine (TCM) with a long history. Research on Herba Epimedii has attracted much attention in China due to its high medicinal value. C8-prenylated flavonol glycosides (PFGs) have been demonstrated to be the main bioactive components in Epimedium brevicornu, and their content determines the medicinal quality. Understanding the biosynthesis pathway of PFGs, exploring genes related to PFGs content, and elucidating the regulatory mechanisms of PFGs biosynthesis pathway is fundamental and essential for improving the quality of E. brevicornu. Here, we provide a comprehensive review of the research on structural and transcriptional factor genes related to the biosynthesis of PFGs, which not only contributes to unravel the regulatory mechanisms related to PFGs content, but also lay a foundation for research on molecular breeding and the synthetic biology in Epimedium plants.

Key words: Herba Epimedii, flavonol glycosides, biosynthesis, structural genes, regulatory genes

范雪兰, 落艳娇, 徐超群, 郭宝林. 淫羊藿类黄酮生物合成相关基因研究进展. 植物学报, 2024, 59(5): 834-846.

Xuelan Fan, Yanjiao Luo, Chaoqun Xu, Baolin Guo. Research Progress on Genes Related to Flavonoids Biosynthesis in Herba Epimedii. Chinese Bulletin of Botany, 2024, 59(5): 834-846.

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https://www.chinbullbotany.com/CN/Y2024/V59/I5/834

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参考文献 61

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基因名	物种	功能	验证方法	参考文献
EsPAL1	箭叶淫羊藿	随着叶片的成熟, 表达量增高, 推测其可能参与木质素生物合成	a	Zeng et al., 2013b
EsPAL2	箭叶淫羊藿	在各发育阶段及组织中均表达, 推测其可能参与木质素、类黄酮和花青素的生物合成	a	Zeng et al., 2013b
EsPAL3	箭叶淫羊藿	主要在叶、花和蒴果中表达, 推测其主要参与花青素和4种主要活性成分的生物合成	a	Zeng et al., 2013b
EwPAL	巫山淫羊藿	体外功能验证: 催化L-苯丙氨酸和L-酪氨酸分别转化为肉桂酸和4-香豆酸	b	Liu et al., 2021b
EpPAL2、EpC4H、Ep4CL3、Ep4CL2、EpCHS2、EpCHI1、EpCHI2、EpCHIL和EpF3H	柔毛淫羊藿	表达模式与4种主要活性成分的积累模式呈正相关	a	Zeng et al., 2013a; Huang et al., 2015; Xu et al., 2023
EsC4H和EsCHS1	箭叶淫羊藿	表达模式与4种主要活性成分的积累模式无显著相关性	a	Zeng et al., 2013a; Huang et al., 2015
Es4CL1、Es4CL2、EsCHS3和EsCHI2	箭叶淫羊藿	表达模式与4种主要活性成分的积累模式呈负相关	a	Zeng et al., 2013a; Huang et al., 2015
Ew4CL1	巫山淫羊藿	体外功能验证: 催化4-香豆酸和肉桂酸乙酰化, 分别转化为4-香豆素基-CoA和肉桂酰基-CoA	b	Liu et al., 2021b
Ew4CL2	巫山淫羊藿	与其它物种参与木质素生物合成的4CL基因聚成一簇, 推测其参与木质素的生物合成	c	Liu et al., 2021b
EwCHS1	巫山淫羊藿	体外功能验证: 催化4-香豆素酰-CoA转化为柚皮素查尔酮和对香豆酰三乙酸内酯; 催化肉桂酰-CoA转化为松属素查尔酮和肉桂酰基三乙酸内酯	b	Liu et al., 2021b
eppsCHS1	拟巫山淫羊藿	表达模式与4种主要活性成分的积累模式相关	a	Pan et al., 2017
EwCHI1	巫山淫羊藿	体外功能验证: 催化柚皮素查尔酮和松属素查尔酮环化形成(2S)-柚皮素和(2S)-松属素	b	Liu et al., 2021b
EwCHIL3	巫山淫羊藿	与EwCHS1相互作用, 抑制对香豆酰三乙酸内酯和肉桂酰基三乙酸内酯产物的生成, 增加柚皮素和松属素的产生	b	Liu et al., 2021b

基因名	物种	功能	验证方法	参考文献
EsPAL1	箭叶淫羊藿	随着叶片的成熟, 表达量增高, 推测其可能参与木质素生物合成	a	Zeng et al., 2013b
EsPAL2	箭叶淫羊藿	在各发育阶段及组织中均表达, 推测其可能参与木质素、类黄酮和花青素的生物合成	a	Zeng et al., 2013b
EsPAL3	箭叶淫羊藿	主要在叶、花和蒴果中表达, 推测其主要参与花青素和4种主要活性成分的生物合成	a	Zeng et al., 2013b
EwPAL	巫山淫羊藿	体外功能验证: 催化L-苯丙氨酸和L-酪氨酸分别转化为肉桂酸和4-香豆酸	b	Liu et al., 2021b
EpPAL2、EpC4H、Ep4CL3、Ep4CL2、EpCHS2、EpCHI1、EpCHI2、EpCHIL和EpF3H	柔毛淫羊藿	表达模式与4种主要活性成分的积累模式呈正相关	a	Zeng et al., 2013a; Huang et al., 2015; Xu et al., 2023
EsC4H和EsCHS1	箭叶淫羊藿	表达模式与4种主要活性成分的积累模式无显著相关性	a	Zeng et al., 2013a; Huang et al., 2015
Es4CL1、Es4CL2、EsCHS3和EsCHI2	箭叶淫羊藿	表达模式与4种主要活性成分的积累模式呈负相关	a	Zeng et al., 2013a; Huang et al., 2015
Ew4CL1	巫山淫羊藿	体外功能验证: 催化4-香豆酸和肉桂酸乙酰化, 分别转化为4-香豆素基-CoA和肉桂酰基-CoA	b	Liu et al., 2021b
Ew4CL2	巫山淫羊藿	与其它物种参与木质素生物合成的4CL基因聚成一簇, 推测其参与木质素的生物合成	c	Liu et al., 2021b
EwCHS1	巫山淫羊藿	体外功能验证: 催化4-香豆素酰-CoA转化为柚皮素查尔酮和对香豆酰三乙酸内酯; 催化肉桂酰-CoA转化为松属素查尔酮和肉桂酰基三乙酸内酯	b	Liu et al., 2021b
eppsCHS1	拟巫山淫羊藿	表达模式与4种主要活性成分的积累模式相关	a	Pan et al., 2017
EwCHI1	巫山淫羊藿	体外功能验证: 催化柚皮素查尔酮和松属素查尔酮环化形成(2S)-柚皮素和(2S)-松属素	b	Liu et al., 2021b
EwCHIL3	巫山淫羊藿	与EwCHS1相互作用, 抑制对香豆酰三乙酸内酯和肉桂酰基三乙酸内酯产物的生成, 增加柚皮素和松属素的产生	b	Liu et al., 2021b

基因名	物种	功能	验证方法	参考文献
EsFLS	箭叶淫羊藿	在烟草中过表达导致花中黄酮醇含量增加, 花青素含量降低	a	Zeng et al., 2013a; Huang et al., 2015
eppsFLS1	拟巫山淫羊藿	体外功能验证: 分别催化二氢山奈酚和二氢槲皮素生成山奈酚和槲皮素	b	Pan et al., 2017
EpFLS3和EsUF3GT	柔毛淫羊藿箭叶淫羊藿	表达模式与4种主要活性成分的积累模式呈正相关	a	Huang et al., 2015; Xu et al., 2023
EsPT2	箭叶淫羊藿	体外功能验证: 分别催化山奈酚、山奈素和柚皮素生成8-异戊烯基山奈酚、淫羊藿素(8-异戊烯基山奈素)和8-异戊烯基柚皮素	b	Wang et al., 2021
EpPT8	柔毛淫羊藿	体外功能验证: 分别催化山奈酚、槲皮素和芹菜素生成8-异戊烯基山奈酚、8-异戊烯基槲皮素和8-异戊烯基芹菜素	b	Shen et al., 2022a
EkF8DT3	朝鲜淫羊藿	体外功能验证: 催化山奈酚生成8-异戊烯基山奈酚	b	周景文等, 2023a
EkF4′OMT1	朝鲜淫羊藿	体外功能验证: 催化8-异戊烯基山奈酚C4′-OH甲基化, 生成淫羊藿素	b	周景文等, 2023b
EsUF7GT	箭叶淫羊藿	表达模式与4种主要活性成分的积累模式呈负相关	a	Huang et al., 2015
EkF3UGal	朝鲜淫羊藿	体外功能验证: 分别催化淫羊藿素和多种不带异戊烯基类黄酮的C3-OH半乳糖基化, 生成8-异戊烯基山奈素-3-O-半乳糖和类黄酮- 3-O-半乳糖苷	b	Lyu et al., 2020
EkF3UGluT	朝鲜淫羊藿	体外功能验证: 分别催化淫羊藿素和多种不带异戊烯基类黄酮的C3-OH葡萄糖基化, 生成8-异戊烯基山奈素-3-O-葡萄糖和类黄酮- 3-O-葡萄糖苷	b	Lyu et al., 2020
EkF3URhaT	朝鲜淫羊藿	体外功能验证: 分别催化淫羊藿素和多种不带异戊烯基黄酮醇的C3-OH鼠李糖基化, 生成宝藿苷I和不含异戊烯基的黄酮醇-3-O-鼠李糖苷	b	Lyu et al., 2020
EpGT60	柔毛淫羊藿	体外功能验证: 分别催化8-异戊烯基山奈酚和淫羊藿素的C3-OH鼠李糖基化, 生成宝藿苷II和宝藿苷I	b	Yao et al., 2022a
EpPF3RT	拟巫山淫羊藿	体外功能验证: 分别催化淫羊藿素、8-异戊烯基山奈酚、山奈酚和山奈素的C3-OH鼠李糖基化, 生成宝藿苷I、宝藿苷II、山奈酚- 3-O-鼠李糖苷和山奈素-3-O-鼠李糖苷	b	Feng et al., 2018
Ep7GT (EpGT8)	拟巫山淫羊藿	体外功能验证: 分别催化宝藿苷I、宝藿苷II和山奈酚的C7-OH葡萄糖基化, 生成淫羊藿苷、朝藿定A和山奈酚-7-O-葡萄糖苷	b	Feng et al., 2019
EsGT1	箭叶淫羊藿	体外功能验证: 分别催化宝藿苷I和山奈酚的C7-OH葡萄糖基化, 生成淫羊藿苷和山奈酚-7-O-葡萄糖苷	b	Yang et al., 2020
EwGGTa	巫山淫羊藿	体外功能验证: 催化鼠李糖2′′-OH位加1分子葡萄糖, 使淫羊藿苷生成朝藿定A	b	虞沂等, 2022
EwGGTb	巫山淫羊藿	体外功能验证: 催化鼠李糖2′′-OH位加1分子木糖, 使淫羊藿苷生成朝藿定B	b	虞沂等, 2022
EpF3R2"XylT	柔毛淫羊藿	体外功能验证: 催化鼠李糖2′′-OH位加1分子木糖, 分别使淫羊藿苷、宝藿苷I、宝藿苷II和朝藿定A生成朝藿定B、箭藿苷B、大花淫羊藿苷F和淫羊藿苷E	b	Yao et al., 2022b

基因名	物种	功能	验证方法	参考文献
EsFLS	箭叶淫羊藿	在烟草中过表达导致花中黄酮醇含量增加, 花青素含量降低	a	Zeng et al., 2013a; Huang et al., 2015
eppsFLS1	拟巫山淫羊藿	体外功能验证: 分别催化二氢山奈酚和二氢槲皮素生成山奈酚和槲皮素	b	Pan et al., 2017
EpFLS3和EsUF3GT	柔毛淫羊藿箭叶淫羊藿	表达模式与4种主要活性成分的积累模式呈正相关	a	Huang et al., 2015; Xu et al., 2023
EsPT2	箭叶淫羊藿	体外功能验证: 分别催化山奈酚、山奈素和柚皮素生成8-异戊烯基山奈酚、淫羊藿素(8-异戊烯基山奈素)和8-异戊烯基柚皮素	b	Wang et al., 2021
EpPT8	柔毛淫羊藿	体外功能验证: 分别催化山奈酚、槲皮素和芹菜素生成8-异戊烯基山奈酚、8-异戊烯基槲皮素和8-异戊烯基芹菜素	b	Shen et al., 2022a
EkF8DT3	朝鲜淫羊藿	体外功能验证: 催化山奈酚生成8-异戊烯基山奈酚	b	周景文等, 2023a
EkF4′OMT1	朝鲜淫羊藿	体外功能验证: 催化8-异戊烯基山奈酚C4′-OH甲基化, 生成淫羊藿素	b	周景文等, 2023b
EsUF7GT	箭叶淫羊藿	表达模式与4种主要活性成分的积累模式呈负相关	a	Huang et al., 2015
EkF3UGal	朝鲜淫羊藿	体外功能验证: 分别催化淫羊藿素和多种不带异戊烯基类黄酮的C3-OH半乳糖基化, 生成8-异戊烯基山奈素-3-O-半乳糖和类黄酮- 3-O-半乳糖苷	b	Lyu et al., 2020
EkF3UGluT	朝鲜淫羊藿	体外功能验证: 分别催化淫羊藿素和多种不带异戊烯基类黄酮的C3-OH葡萄糖基化, 生成8-异戊烯基山奈素-3-O-葡萄糖和类黄酮- 3-O-葡萄糖苷	b	Lyu et al., 2020
EkF3URhaT	朝鲜淫羊藿	体外功能验证: 分别催化淫羊藿素和多种不带异戊烯基黄酮醇的C3-OH鼠李糖基化, 生成宝藿苷I和不含异戊烯基的黄酮醇-3-O-鼠李糖苷	b	Lyu et al., 2020
EpGT60	柔毛淫羊藿	体外功能验证: 分别催化8-异戊烯基山奈酚和淫羊藿素的C3-OH鼠李糖基化, 生成宝藿苷II和宝藿苷I	b	Yao et al., 2022a
EpPF3RT	拟巫山淫羊藿	体外功能验证: 分别催化淫羊藿素、8-异戊烯基山奈酚、山奈酚和山奈素的C3-OH鼠李糖基化, 生成宝藿苷I、宝藿苷II、山奈酚- 3-O-鼠李糖苷和山奈素-3-O-鼠李糖苷	b	Feng et al., 2018
Ep7GT (EpGT8)	拟巫山淫羊藿	体外功能验证: 分别催化宝藿苷I、宝藿苷II和山奈酚的C7-OH葡萄糖基化, 生成淫羊藿苷、朝藿定A和山奈酚-7-O-葡萄糖苷	b	Feng et al., 2019
EsGT1	箭叶淫羊藿	体外功能验证: 分别催化宝藿苷I和山奈酚的C7-OH葡萄糖基化, 生成淫羊藿苷和山奈酚-7-O-葡萄糖苷	b	Yang et al., 2020
EwGGTa	巫山淫羊藿	体外功能验证: 催化鼠李糖2′′-OH位加1分子葡萄糖, 使淫羊藿苷生成朝藿定A	b	虞沂等, 2022
EwGGTb	巫山淫羊藿	体外功能验证: 催化鼠李糖2′′-OH位加1分子木糖, 使淫羊藿苷生成朝藿定B	b	虞沂等, 2022
EpF3R2"XylT	柔毛淫羊藿	体外功能验证: 催化鼠李糖2′′-OH位加1分子木糖, 分别使淫羊藿苷、宝藿苷I、宝藿苷II和朝藿定A生成朝藿定B、箭藿苷B、大花淫羊藿苷F和淫羊藿苷E	b	Yao et al., 2022b

基因名	物种	功能	验证方法	参考文献
EsF3′H､EsF3′5′H和EsANS	箭叶淫羊藿	表达模式与叶片花青素的积累模式呈正相关	a	Huang et al., 2012, 2015; Zeng et al., 2013a
EsDFR1	箭叶淫羊藿	表达模式与4种主要活性成分的积累模式无显著相关性	a	Zeng et al., 2013a; Huang et al., 2015
EsDFR2	箭叶淫羊藿	表达模式与4种主要活性成分的积累模式呈负相关	a	Huang et al., 2012, 2015; Zeng et al., 2013a
EpGT43	柔毛淫羊藿	体外功能验证: 以UDP-葡萄糖为供体, 催化矢车菊素3-O-鼠李糖苷。推测其酶活产物可能为矢车菊素3-O-(2- O-(-葡萄糖基))-鼠李糖苷	b, c	姚宇, 2023

淫羊藿类黄酮生物合成相关基因研究进展

Research Progress on Genes Related to Flavonoids Biosynthesis in Herba Epimedii

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基因名	物种	功能	验证方法	参考文献
EsMYB1	箭叶淫羊藿	与拟南芥中负调控肉桂酸-4-羟化酶的AtMYB4基因高度同源, 推测其为类黄酮生物合成转录抑制因子	b	Huang et al., 2013b
EsMYB5和EsMYB11	箭叶淫羊藿	与葡萄中调控原花青素生物合成的VvMYBPA2具有约50%的相似性, 推测其参与原花青素积累	b	Huang et al., 2013b
EsMYB7和EsMYB10	箭叶淫羊藿	与拟南芥中调控原花青素生物合成的AtTT2 (At- MYB123)高度同源, 推测其参与调节种皮中原花青素的积累	b	Huang et al., 2013b
EsMYB9	箭叶淫羊藿	单独作用时轻微激活EsCHS的启动子或与bHLH (EsTT8)相互作用强烈激活EsCHS、EsF3H、EsDFR1、EsDFR2和EsANS的启动子, 正调控花青素和黄酮醇的生物合成	a, c	Huang et al., 2013b, 2017
EsMYB12	箭叶淫羊藿	表达模式与4种主要活性成分的积累模式呈负相关	a, b	Huang et al., 2013b, 2015
EsMYBA1	箭叶淫羊藿	单独作用或者与bHLH (EsTT8)相互作用激活EsDFR和EsANS的启动子, 正调控花青素的生物合成	a, c	Huang et al., 2013a, 2015
EsMYBF1	箭叶淫羊藿	强烈激活EsF3H和EsFLS的启动子, 正调控叶片4种主要活性成分的生物合成	a, c	Huang et al., 2015, 2016a
EsAN2 (MYB)	箭叶淫羊藿	单独作用或与bHLH (EsTT8)相互作用激活早期和晚期花青素途径基因, 正调控花青素的生物合成	a, c	Huang et al., 2016b
Ebr05G057070、Ebr05G003750、Ebr05G010200、Ebr05G056880、Ebr01G039680、Ebr01G039880 (MYB)、EsGL3 (bHLH)、EsTTG1 (WD40)、Ebr05G038380 (bZIP) 和Ebr03G071730 (WRKY)	柔毛淫羊藿和箭叶淫羊藿	表达模式与4种主要活性成分的积累模式呈正相关	a	Huang et al., 2015; Xu et al., 2023
EsTT8 (bHLH)	箭叶淫羊藿	表达模式与4种主要活性成分的积累模式呈负相关	a	Huang et al., 2013a, 2015, 2016b, 2017

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