植物单宁合成调控及其对环境的响应机制

doi:10.13560/j.cnki.biotech.bull.1985.2025-0022

摘要/Abstract

摘要：

单宁是植物中一类重要的次生代谢物，广泛存在于植物各组织和器官中，并在植物生长发育过程中发挥至关重要的作用。作为一类具有强大生物活性的化合物，单宁不仅可以帮助植物有效应对各种环境挑战，还在植物的免疫防御系统中发挥核心作用。单宁的合成途径复杂，通常分为前体合成和单宁聚合两个主要阶段。前体合成以芳香族氨基酸为起点，通过一系列酶促反应转化为单宁的基本单元；单宁的聚合则涉及这些单元的交联和聚合，形成具有生物活性的复杂大分子。目前，已有大量研究探讨了单宁的合成途径、调控机制及其对环境的响应，然而仍缺乏系统的总结和全面的概述。本文详细介绍了植物单宁的结构特征，分析了其在植物生长发育中的多重功能，并讨论了植物单宁的生物合成途径。此外，本文总结了环境因素对单宁合成的调控作用，进一步揭示了单宁在植物应对逆境中的潜力，特别是在增强植物抗逆性和生态适应性方面的应用前景。最后，对当前植物单宁研究中尚未解决的问题和未来研究方向进行了分析和展望，旨在为植物单宁的合理开发与应用提供理论依据，推动其在植物抗逆性提升、生态适应性增强以及植物生态功能优化等方面的应用。

关键词: 植物单宁, 生物合成, 转录调控, 环境响应

Abstract:

Tannins are an important class of secondary metabolites in plants, widely distributed in various tissues and organs of plants, and play a crucial role in plant growth and development. As compounds with strong biological activity, tannins not only help plants effectively cope with various environmental challenges but also play a core role in the plant immune defense system. The biosynthesis of tannins is complex and is typically divided into two main stages: Precursor synthesis and tannin polymerization. Precursor synthesis begins with aromatic amino acids, which are converted into the basic units of tannins through a series of enzymatic reactions. Tannin polymerization is involved in the cross-linking and polymerization of these units to form complex macromolecules with biological activity. Although extensive research has been conducted on the synthesis pathways, regulatory mechanisms, and environmental responses of tannins, there is still a lack of systematic summaries and comprehensive overviews. This paper provides a detailed introduction to the structural characteristics of plant tannins, analyzes their multiple functions in plant growth and development, and discusses the biosynthetic pathways of tannins. Additionally, this paper summarizes the regulatory effects of environmental factors on tannin synthesis, further revealing the potential of tannins in helping plants cope with stress, particularly in enhancing plant resistance to stress and ecological adaptability. Finally, this paper analyzes and provides an outlook on unresolved issues in current tannin research and future research directions, aiming to provide theoretical support for the rational development and application of plant tannins, and promoting their application in improving plant stress resistance, enhancing ecological adaptability, and optimizing plant ecological functions.

Key words: plant tannins, biosynthesis, transcriptional regulation, response to environment

高婧, 陈益存, 高暝, 赵耘霄, 汪阳东. 植物单宁合成调控及其对环境的响应机制[J]. 生物技术通报, 2025, 41(7): 49-59.

GAO Jing, CHENG Yi-cun, GAO Ming, ZHAO Yun-xiao, WANG Yang-dong. Regulation of Plant Tannin Synthesis and Mechanisms of Its Responses to Environment[J]. Biotechnology Bulletin, 2025, 41(7): 49-59.

图/表 4

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来源植物 Plants of source	单宁类型 Type of tannins	主要功能 Main functions	单宁含量 Tannin content （%）	参考文献 Reference
柿Diospyros kaki	缩合单宁	抗氧化、抗炎、抗辐射、抗菌、抗病毒、抗高血压、降血脂、降血糖、解毒	4	［22］
杨属Populus	缩合单宁	抗氧化、抗菌、抗紫外线、抗病原体	15-25	［23-24］
茶叶Camellia sinensis	水解单宁和缩合单宁	抗氧化、抗菌、抗过敏、抗紫外线、消炎、除臭、抗虫、细胞防御、促进免疫	20-35	［13， 25］
葡萄Vitis vinifera	水解单宁和缩合单宁	抗氧化、抗菌、抗病毒、抗衰老、降血糖、抑制络氨酸酶活性、抑制呼吸道炎症、保护心血管、保护神经元、保护软骨组织	40-50	［26-27］
栎属Quercus	水解单宁和缩合单宁	抗氧化、抗炎、抗菌、抗增殖、免疫调节、降血糖、降胆固醇、保胃保肝	25-35	［28］
石榴Punica granatum	水解单宁	抗氧化应激、抗炎、抗癌、抗真菌、抗病毒、促进伤口愈合	10-20	［29］
核桃Juglans regia	水解单宁和缩合单宁	抗氧化、抗菌、抗肿瘤、镇痛	20-30	［30］
五倍子Rhus chinensis	水解单宁	抗氧化、抗病原微生物、抗癌、抗龋齿、抗肥胖、抗炎、止血、降血糖	65-70	［31］
麻黄Ephedrae Herba	缩合单宁	抗炎、抗氧化、抗增殖、抗菌、促进伤口愈合、酶抑制活性	10-20	［32］
杨梅Morella rubra	水解单宁和缩合单宁	抗氧化、抗炎、抗过敏、抗癌、抗菌、止泻	10-20	［33］
桉属Eucalyptus	缩合单宁和水解单宁	抗氧化、抗菌、抗炎、抑制酶活性	40-50	［34-35］
油茶Camellia oleifera	水解单宁和缩合单宁	抗氧化、抗炎、抗菌、抗肿瘤、降血糖	10-20	［36-37］
黑荆Acacia mearnsii	缩合单宁	抗氧化、抗菌、抗糖尿病、抗虫、抗肥胖、抑制酶活性	30-45	［38］

来源植物 Plants of source	单宁类型 Type of tannins	主要功能 Main functions	单宁含量 Tannin content （%）	参考文献 Reference
柿Diospyros kaki	缩合单宁	抗氧化、抗炎、抗辐射、抗菌、抗病毒、抗高血压、降血脂、降血糖、解毒	4	［22］
杨属Populus	缩合单宁	抗氧化、抗菌、抗紫外线、抗病原体	15-25	［23-24］
茶叶Camellia sinensis	水解单宁和缩合单宁	抗氧化、抗菌、抗过敏、抗紫外线、消炎、除臭、抗虫、细胞防御、促进免疫	20-35	［13， 25］
葡萄Vitis vinifera	水解单宁和缩合单宁	抗氧化、抗菌、抗病毒、抗衰老、降血糖、抑制络氨酸酶活性、抑制呼吸道炎症、保护心血管、保护神经元、保护软骨组织	40-50	［26-27］
栎属Quercus	水解单宁和缩合单宁	抗氧化、抗炎、抗菌、抗增殖、免疫调节、降血糖、降胆固醇、保胃保肝	25-35	［28］
石榴Punica granatum	水解单宁	抗氧化应激、抗炎、抗癌、抗真菌、抗病毒、促进伤口愈合	10-20	［29］
核桃Juglans regia	水解单宁和缩合单宁	抗氧化、抗菌、抗肿瘤、镇痛	20-30	［30］
五倍子Rhus chinensis	水解单宁	抗氧化、抗病原微生物、抗癌、抗龋齿、抗肥胖、抗炎、止血、降血糖	65-70	［31］
麻黄Ephedrae Herba	缩合单宁	抗炎、抗氧化、抗增殖、抗菌、促进伤口愈合、酶抑制活性	10-20	［32］
杨梅Morella rubra	水解单宁和缩合单宁	抗氧化、抗炎、抗过敏、抗癌、抗菌、止泻	10-20	［33］
桉属Eucalyptus	缩合单宁和水解单宁	抗氧化、抗菌、抗炎、抑制酶活性	40-50	［34-35］
油茶Camellia oleifera	水解单宁和缩合单宁	抗氧化、抗炎、抗菌、抗肿瘤、降血糖	10-20	［36-37］
黑荆Acacia mearnsii	缩合单宁	抗氧化、抗菌、抗糖尿病、抗虫、抗肥胖、抑制酶活性	30-45	［38］

环境因子 Environmental factors	响应因子 Response factors	关键酶及转录因子 Key enzymes and transcription factors	对单宁合成的影响 Effects on tannin synthesis	参考文献 Reference
光照Light	苯丙氨酸解氨酶和过氧化物酶活性增加	PAL、C4H	促进	［64-65］
紫外线Ultraviolet radiation	抗氧化酶活性增加	MYB、DFR、ANS	促进	［23， 66］
高温High temperature	增强热应激信号的转导	HSPs、MBW	短时高温抑制；长时高温促进	［67-68］
寒冷Cold	水杨酸响应同时抗氧化酶活性提高	SOD、CAT、POX、bHLH	促进	［69］
干旱Drought	改变苯丙烷途径提高抗氧化酶活性	MBW、WRKY、UGT、	促进	［23， 70］
水分Water	抗氧化酶活性	MBW、C4H、CHI	随着胁迫程度的增加先促进后抑制	［71］
重金属Heavy metals	生物合成酶活性增加，基因表达增强	SDH、PAL、CHI、PPO	促进	［16， 18］
盐分Salt	基因表达增强	FNSII、bHLH、CHS	促进	［16， 72］
昆虫Insects	基因表达增强	DAPHS、UGT、CHS、LAR	促进	［10， 73］
病原体Pathogens	触发免疫反应	NBS-LRR、SDH	促进	［74-75］

环境因子 Environmental factors	响应因子 Response factors	关键酶及转录因子 Key enzymes and transcription factors	对单宁合成的影响 Effects on tannin synthesis	参考文献 Reference
光照Light	苯丙氨酸解氨酶和过氧化物酶活性增加	PAL、C4H	促进	［64-65］
紫外线Ultraviolet radiation	抗氧化酶活性增加	MYB、DFR、ANS	促进	［23， 66］
高温High temperature	增强热应激信号的转导	HSPs、MBW	短时高温抑制；长时高温促进	［67-68］
寒冷Cold	水杨酸响应同时抗氧化酶活性提高	SOD、CAT、POX、bHLH	促进	［69］
干旱Drought	改变苯丙烷途径提高抗氧化酶活性	MBW、WRKY、UGT、	促进	［23， 70］
水分Water	抗氧化酶活性	MBW、C4H、CHI	随着胁迫程度的增加先促进后抑制	［71］
重金属Heavy metals	生物合成酶活性增加，基因表达增强	SDH、PAL、CHI、PPO	促进	［16， 18］
盐分Salt	基因表达增强	FNSII、bHLH、CHS	促进	［16， 72］
昆虫Insects	基因表达增强	DAPHS、UGT、CHS、LAR	促进	［10， 73］
病原体Pathogens	触发免疫反应	NBS-LRR、SDH	促进	［74-75］