银杏萜内酯合成代谢调控研究进展

doi:10.11707/j.1001-7488.LYKX20250497

摘要/Abstract

摘要：

银杏为银杏科银杏属国家一级保护孑遗植物，具有较高的观赏、经济和药用价值。萜内酯作为银杏叶中最重要的药用活性成分之一，具有抗登革热、抗炎、抗癌和抗氧化特性，对血小板活化因子受体有强大的特异性抑制作用，是治疗阿尔兹海默病的有效制剂，故而在银杏优良树种的选育过程中，萜内酯含量是极为重要的考量指标。本文深入探讨了萜内酯生物合成调控网络，从甲基赤藓糖醇4-磷酸（MEP）途径和甲羟戊酸（MVA）途径出发，全面综述了已鉴定的20个结构基因、5类关键转录因子（MYB、WRKY、bZIP、bHLH和AP2/ERF）以及2类非编码RNA（miRNA和lncRNA）对萜内酯合成的正向或负向调控作用。同时，系统分析了环境因素（光照、温度、水分、养分等）和植物激素（ABA、SA、MeJA、IAA、ETH、GA₃、CCC等）对萜内酯合成的间接影响，明确了紫外照射、适度干旱等外部环境刺激，以及外源施加MeJA、SA、CCC等激素可显著提升银杏叶片中萜内酯含量，而土壤肥力、种植密度以及病虫害等因素也对其积累存在一定影响。最后，本文提出构建“内部基因调控”与“外部环境优化”协同策略，以促进银杏萜内酯含量，旨在为后续的生产实践和科学研究提供具有重要参考价值的理论指导，进而实现银杏药用价值和商业价值的提升。

关键词: 银杏, 萜内酯, 生物合成, 调控网络, 环境因素, 植物激素

Abstract:

: Ginkgo biloba is a national first-class protected relict plant belonging to the genus Ginkgo in the family Ginkgoaceae, boasting high ornamental, economic, and medicinal values. Terpene trilactones, as one of the most significant bioactive components in G. biloba leaves, exhibit anti-dengue, anti-inflammatory, anti-cancer, and antioxidant properties. Specifically, they demonstrate a potent and specific inhibitory effect on platelet-activating factor receptors and serve as effective agents for the treatment of Alzheimer's disease. Consequently, the content of terpene trilactone is considered a crucial selection index in the breeding of superior G. biloba varieties. This paper provides a comprehensive review of the regulatory network underlying the biosynthesis of terpene trilactones. Based on the Methylerythritol 4-phosphate (MEP) and Mevalonate (MVA) pathways, the positive and negative regulatory roles of 20 identified structural genes, five categories of key transcription factors (including MYB, WRKY, bZIP, bHLH, and AP2/ERF), and two types of non-coding RNAs (miRNA and lncRNA) are thoroughly discussed. Furthermore, the indirect effects of environmental factors (such as light, temperature, water, and nutrients) and plant hormones (including ABA, SA, MeJA, IAA, ETH, GA₃, and CCC) on the biosynthesis of terpene trilactones are systematically analyzed. It is clarified that external environmental stimuli, such as ultraviolet radiation and moderate drought, as well as the exogenous application of hormones, such as MeJA, SA, and CCC, can significantly enhance the accumulation of terpene trilactones in G. biloba leaves. Additionally, the factors such as soil fertility, planting density, and the occurrence of pests and diseases also influence their accumulation. Finally, a synergistic strategy integrating “endogenous gene regulation” and “exogenous environmental optimization” is proposed to promote the content of terpene trilactones. This review aims to provide theoretical guidance with significant reference value for future production practices and scientific research, thereby maximizing the medicinal and commercial potential of G. biloba.

Key words: Ginkgo biloba, terpene trilactones, biosynthesis, regulation network, environmental factors, plant hormones

中图分类号:

S722.3

王采妮,李颖,苏东雪,耿微微,叶家保,许锋,张威威,廖咏玲. 银杏萜内酯合成代谢调控研究进展[J]. 林业科学, 2026, 62(4): 217-230.

Caini Wang,Ying Li,Dongxue Su,Weiwei Geng,Jiabao Ye,Feng Xu,Weiwei Zhang,Yongling Liao. Research Progress in Metabolic Regulation of Terpene Trilactones in Ginkgo biloba[J]. Scientia Silvae Sinicae, 2026, 62(4): 217-230.

图/表 7

图S1

图1

图2

图3

表S1

不同环境条件对银杏萜内酯积累的影响"

序号 No.	环境条件 Environmental conditions	处理方法 Treatment methods	总萜内酯含量 TTLs content （%）	调控基因 Regulatory genes	参考文献 References
1	光照 Light	UV，24 h	18.92	—	冯如等，2017 Feng et al.，2017
2	光照 Light	紫膜（涤纶滤光膜，厚0.2 mm），3个月 Purple membrane （polyester filter membrane， 0.2 mm thick）， 3 months	85.23	—	冷平生等，2002 Leng et al.，2002
3	光照 Light	1500 J·m^–2 UV-B，12 h	—	GbERF13	Li et al.，2019
4	光照 Light	100 μmol·m^?2 s^?1，白天16 h/夜晚8 h，24 ℃，4天 100 μmol·m^?2s^?1， 16 h during the day/8 h at night， 24 ℃， 4 days	15.10	GbMVD	Liao et al.，2016
5	光照 Light	1500 μJ·m^–2 UV，48 h	21.89	GbDXS、GbGGPPS、GbLPS、GbMVD	Zheng et al.，2020
6	温度 Temperature	4 ℃，8天 4 ℃， 8 days	14.50	GbDXS、GbGGPPS、GbLPS、GbMVD	Zheng et al.，2020
7	水分 Water	干旱（土壤含水量55%~60%），7个月 Drought （soil moisture 55%-60%）， 7 months	23.00	GbHMGR、GbMVD、GbDXS、GbDXR、MECT、MECPs	朱灿灿等，2010；2011 Zhu et al.，2010；2011
8	水分 Water	干旱（土壤相对含水量约50%），30天 Drought （soil relative water content about 50%）， 30 days	15.68	—	冯如等，2017 Feng et al.，2017
9	水分 Water	干旱（连续不浇水15天），12 h Drought （no watering for 15 days）， 12 h	—	GbEFR1、GbERF25	Li et al.，2019
10	水分 Water	PEG-6000（20%，W/V）溶液，6 h PEG-6000（20%，W/V）solution， 6 h	—	GbHMGS1、GbHMGR1、GbHMGR2	Yu et al.，2021
11	养分 Nutrition	叶面喷施，15 mg·L^?1 Na₂SeO₃，5周 Foliar application， 15 mg·L^?1 Na₂SeO₃， 5 weeks	58.00	GbWRKY、GbAP2/ERF、GbDXS、GbDXR、GbHMGR、GbMECPs、GbCYP450	Li et al.，2022
12	养分 Nutrition	叶面喷施，0.5%尿素，6%过磷酸钙溶液，接种G.mosseae Foliar application， 0.5% urea， 6% superphosphate solution， inoculation with G.mosseae	17.90	—	冷平生等，2001 Leng et al.，2001

表S1

表S2

不同外源激素处理对银杏萜内酯积累的影响"

序号 No.	激素 Hormones	处理方法 Treatment methods	总萜内酯含量 TTLs content （%）	调控基因 Regulatory genes	参考文献 References
1	赤霉素 GA₃	叶面喷施，0.04 mg·mL^?1，7天 Foliar application， 0.04 mg·mL^?1， 7 days	16.76	—	冯如等，2017 Feng et al.，2017
2	矮壮素 CCC	叶面喷施，0.2 mg·mL^?1，7天 Foliar application， 0.2 mg·mL^?1， 7 days	29.19	—	冯如等，2017 Feng et al.，2017
3	矮壮素 CCC	叶面喷施，2.0 g·L^?1，8周（每隔2周处理1次，共4次） Foliar application， 2.0 g·L^?1， 8 weeks （4 treatments every 2 weeks）	TTLs：14.80； GA：18.60； GB：12.50； BB：17.50	DXS、DXR、GGPPS、LPS	许锋等，2011 Xu et al.，2011
4	矮壮素 CCC	叶面喷施，2000 μL·L^?1，19天 Foliar application， 2000 μL·L^?1， 19 days	叶Leaves：107.12；根Roots：62.03；茎Stems：59.80	—	冷平生等，2004 Leng et al.，2004
5	萘乙酸 NAA	叶片喷施，5 mg·L^?1，15天 Foliar application， 5 mg·L^?1， 15 days	13.50	—	王鼎豪，2021 Wang et al.，2021
6	多效唑 PBZ	叶片喷施，50 mg·L^?1，10天 Foliar application， 50 mg·L^?1， 10 days	39.30	—	王鼎豪，2021 Wang et al.，2021
7	ET	叶面喷施，0.02%，30天 Foliar application， 0.02%， 30 days	110.71	—	谢宝东等，2002 Xie et al.，2002
8	脱落酸ABA	叶面喷施，100 μmol·L^?1，2天 Foliar application， 100 μmol·L^?1， 2 days	1.77	DXS、GGPPS、LPS、MVD	Zheng et al.，2020
9	茉莉酸甲酯MeJA	叶面喷施，100 μmol·L^?1，8天 Foliar application， 100 μmol·L^?1， 8 days	14.50	DXS、GGPPS、LPS、MVD	Zheng et al.，2020
10	乙烯 ETH	叶面喷施，10 mmol·L^?1，8天 Foliar application， 10 mmol·L^?1， 8 days	10.20	DXS、GGPPS、LPS、MVD	Zheng et al.，2020
11	水杨酸 SA	叶面喷施，10 mmol·L^?1，8天 Foliar application， 10 mmol·L^?1， 8 days	9.20	DXS、GGPPS、LPS、MVD	Zheng et al.，2020
12	茉莉酸甲酯MeJA	细胞悬浮液培养，0.1 mmol·L^?1，12 h， 3%（W/V）蔗糖，18.8 mmol·L^?1 NAA，25 ℃ Cell suspension culture， 0.1 mmol·L^?1， 12 h， 3% （W/V） sucrose， 18.8 mmol·L^?1 NAA， 25 ℃	GA：430；GB：820	—	Kang et al.，2006
13	茉莉酸甲酯MeJA	细胞悬浮液培养，1.0 mmol·L^?1，12 h 3%（W/V）蔗糖，18.8 mmol·L^?1 NAA，25 ℃ Cell suspension culture， 0.1 mmol·L^?1， 12 h， 3% （W/V） sucrose， 18.8 mmol·L^?1 NAA， 25 ℃	BB：625	—	Kang et al.，2006
14	水杨酸 SA	细胞悬浮液培养，1.0 mmol·L^?1，12 h 3% （W/V）蔗糖，18.8 mmol·L^?1 NAA，25 ℃ Cell suspension culture， 0.1 mmol·L^?1， 12 h， 3% （W/V） sucrose， 18.8 mmol·L^?1 NAA， 25 ℃	GA：310；GB：610	—	Kang et al.，2006
15	水杨酸 SA	细胞悬浮液培养，0.01 mmol·L^?1，12 h， 3%（W/V）蔗糖，0.3%（W/V）结冷胶， 18.8 mmol·L^?1 NAA，pH 5.6，25 ℃避光 Cell suspension culture， 0.01 mmol·L^?1， 12 h， 3% sucrose， 0.3% gellan gum， 18.8 mmol·L^?1 NAA， pH 5.6， 25 ℃ dark	BB：540	—	Kang et al.，2006
16	茉莉酸甲酯+水杨酸MeJA+SA	细胞悬浮液培养，0.1 mmol·L^?1，24 h，30 g·L^?1蔗糖，2 g结冷胶，2 mg·L^?1 NAA，0.1 mg·L^?1 KT，pH 5.8，25 ℃避光 Cell suspension culture， 0.1 mmol·L^?1， 24 h， 30 g·L^?1 sucrose， 2 g gellan gum， 2 mg·L^?1 NAA 0.1 mg·L^?1 KT， pH 5.8， 25 ℃ dark	BB：178；GA：195；GB：205；GC：295	—	Sukito et al.，2016
17	茉莉酸甲酯MeJA	叶面喷施，400 μmol·L^?1，60天（每隔10天处理1次，共6次） Foliar application， 400 μmol·L^?1， 60 days （6 treatments every 10 days）	30.76	CYP450	Tan et al.，2022
18	水杨酸 SA	叶面喷施，2.0 mmol·L^?1，50天（每隔10天处理1次，共5次） Foliar application， 2.0 mmol·L^?1， 50 days （5 treatments every 10 days）	23.63	HMGR、TPS、CYPs、CPR、MYB、WRKY	Ye et al.，2020a
19	矮壮素 CCC	叶面喷施，10 mmol·L^?1，6 h Foliar application， 10 mmol·L^?1， 6 h	—	ERF25、ERF27	Li et al.，2019
20	茉莉酸甲酯MeJA	叶面喷施，0.5%（V/V），3周 Foliar application， 0.5% （V/V）， 3 weeks	GA：16-25； GB：10	—	Horbowicz et al.，2021

表S2

图4

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