微生物学报

Category	Compounds	Functions	References
Root exudates	Phenolic compounds	Antioxidant, antibacterial, and antiviral properties improve plant disease resistance	[10]
Organic acids	Promote mineral absorption and regulate microbial growth by either inhibiting or promoting it	[11]
Amino acids	Provide carbon and nitrogen sources that affect the composition of microbial communities	[12]
Carbohydrates	Enhance the diversity and complexity of microbial communities to support the proliferation of beneficial microorganisms	[13]
Volatile organic compounds (VOCs)	Aboveground VOCs	Attract natural enemies of pests, induce defense responses in neighboring plants, and indirectly influence soil microbial perception as signaling factors	[14]
Root VOCs	Antibacterial or bacteriostatic agents attract beneficial microorganisms and serve as carbon sources for some of them	[15]
Plant hormones and their analogs	Salicylic acid (SA)	Inhibit pathogen growth and recruit disease-resistant microorganisms	[16]
Jasmonic acid (JA)	Involved in plant defense and affecting microbial colonization	[17]
Ethylene (ET)	Involved in plant development, senescence, and stress responses	[18]
Strigolactones (SLs)	Specifically attract arbuscular mycorrhizal fungi (AMF), induce their spore germination and hyphal branching, and promote the establishment of mycorrhizal symbiosis	[19]
Cytokinins (CK)	Regulate growth and development; perceive microorganisms	[20]
Abscisic acid (ABA)	Limit pathogen invasion while enhancing osmotic protection and antioxidant capacity	[21]
Gibberellins (GA)	Promote growth and development, enhance JA-related defense mechanisms, and restrict pathogen expansion	[22]
Defense-related secondary metabolites	Alkaloids	Directly inhibit or kill pathogens	[23]
Pathogenesis-related (PR) proteins and antimicrobial peptides (AMPs)	PR proteins, serine protease inhibitors	Defensive proteins that inhibit protein digestion in attacking pests	[24]
AMPs	The plant innate immune system is widely used as a source of diagnostic molecular markers for defense signaling pathways	[25]

Category	Compounds	Functions	References
Root exudates	Phenolic compounds	Antioxidant, antibacterial, and antiviral properties improve plant disease resistance	[10]
Organic acids	Promote mineral absorption and regulate microbial growth by either inhibiting or promoting it	[11]
Amino acids	Provide carbon and nitrogen sources that affect the composition of microbial communities	[12]
Carbohydrates	Enhance the diversity and complexity of microbial communities to support the proliferation of beneficial microorganisms	[13]
Volatile organic compounds (VOCs)	Aboveground VOCs	Attract natural enemies of pests, induce defense responses in neighboring plants, and indirectly influence soil microbial perception as signaling factors	[14]
Root VOCs	Antibacterial or bacteriostatic agents attract beneficial microorganisms and serve as carbon sources for some of them	[15]
Plant hormones and their analogs	Salicylic acid (SA)	Inhibit pathogen growth and recruit disease-resistant microorganisms	[16]
Jasmonic acid (JA)	Involved in plant defense and affecting microbial colonization	[17]
Ethylene (ET)	Involved in plant development, senescence, and stress responses	[18]
Strigolactones (SLs)	Specifically attract arbuscular mycorrhizal fungi (AMF), induce their spore germination and hyphal branching, and promote the establishment of mycorrhizal symbiosis	[19]
Cytokinins (CK)	Regulate growth and development; perceive microorganisms	[20]
Abscisic acid (ABA)	Limit pathogen invasion while enhancing osmotic protection and antioxidant capacity	[21]
Gibberellins (GA)	Promote growth and development, enhance JA-related defense mechanisms, and restrict pathogen expansion	[22]
Defense-related secondary metabolites	Alkaloids	Directly inhibit or kill pathogens	[23]
Pathogenesis-related (PR) proteins and antimicrobial peptides (AMPs)	PR proteins, serine protease inhibitors	Defensive proteins that inhibit protein digestion in attacking pests	[24]
AMPs	The plant innate immune system is widely used as a source of diagnostic molecular markers for defense signaling pathways	[25]

Category	Compounds	Functions	References
Hormone analogs	1-aminocyclopropane-1-carboxylic acid (ACC) deaminase	Alleviate the adverse effects of ethylene on plant growth, promote development, and delay the senescence of medicinal plants	[62-63]
Indole-3-acetic acid (IAA)	Affect the growth and development of medicinal plants and play a key role in plant-microbe communication	[64]
Quorum sensing molecules	N-acyl homoserine lactones (AHLs)	Regulate the immunity of medicinal plants to enhance their resistance to various pathogens	[65]
Pseudomonas secretions	Siderophores/antibiotics	Antagonize pathogens	[66]
Mycorrhizal factors	Lipochitooligosaccharides (LCOs)	Recognized by plant root cell receptors, these molecules activate symbiotic signaling pathways and promote the formation of arbuscular mycorrhizal symbiosis	[67]
Short-chain chitin oligomers (COs)	Act synergistically with LCOs to trigger calcium oscillation signals in plant cells and regulate the expression of genes related to mycorrhizal symbiosis	[68]
Effector proteins	Interact with host plant transcription factors, inhibit plant defense responses, and promote the establishment of ectomycorrhizal symbiosis	[69]

Category	Compounds	Functions	References
Hormone analogs	1-aminocyclopropane-1-carboxylic acid (ACC) deaminase	Alleviate the adverse effects of ethylene on plant growth, promote development, and delay the senescence of medicinal plants	[62-63]
Indole-3-acetic acid (IAA)	Affect the growth and development of medicinal plants and play a key role in plant-microbe communication	[64]
Quorum sensing molecules	N-acyl homoserine lactones (AHLs)	Regulate the immunity of medicinal plants to enhance their resistance to various pathogens	[65]
Pseudomonas secretions	Siderophores/antibiotics	Antagonize pathogens	[66]
Mycorrhizal factors	Lipochitooligosaccharides (LCOs)	Recognized by plant root cell receptors, these molecules activate symbiotic signaling pathways and promote the formation of arbuscular mycorrhizal symbiosis	[67]
Short-chain chitin oligomers (COs)	Act synergistically with LCOs to trigger calcium oscillation signals in plant cells and regulate the expression of genes related to mycorrhizal symbiosis	[68]
Effector proteins	Interact with host plant transcription factors, inhibit plant defense responses, and promote the establishment of ectomycorrhizal symbiosis	[69]

药用植物-微生物互作的化学信号及其作用

PDF下载

王爱媛 , 冷春燕 , 黄信洁 , 邢咏梅 , 陈娟

微生物学报 | 综述 2026,66(8): 3718-3731

收起

微生物学报 |综述 2026 , 66 (8) : 3718 -3731

药用植物-微生物互作的化学信号及其作用

全屏

[Author(id=1297571208107684826, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, orderNo=0, firstName=null, middleName=null, lastName=null, nameCn=null, orcid=null, stid=null, country=null, authorPic=null, dead=0, email=null, emailSecond=null, emailThird=null, correspondingAuthor=0, authorType=1, ext={EN=AuthorExt(id=1297571208187376604, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, authorId=1297571208107684826, language=EN, stringName=Aiyuan WANG, firstName=Aiyuan, middleName=null, lastName=WANG, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing, China, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null), CN=AuthorExt(id=1297571208283845597, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, authorId=1297571208107684826, language=CN, stringName=王爱媛, firstName=null, middleName=null, lastName=null, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=中国医学科学院北京协和医学院，药用植物研究所，道地药材品质保障与资源持续利用全国重点实验室，北京, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null)}, companyList=[AuthorCompany(id=1297571207998632918, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, xref=null, ext=[AuthorCompanyExt(id=1297571208011215831, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, companyId=1297571207998632918, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing, China), AuthorCompanyExt(id=1297571208019604440, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, companyId=1297571207998632918, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=中国医学科学院北京协和医学院，药用植物研究所，道地药材品质保障与资源持续利用全国重点实验室，北京)])]), Author(id=1297571208359343071, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, orderNo=1, firstName=null, middleName=null, lastName=null, nameCn=null, orcid=null, stid=null, country=null, authorPic=null, dead=0, email=null, emailSecond=null, emailThird=null, correspondingAuthor=0, authorType=1, ext={EN=AuthorExt(id=1297571208451617761, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, authorId=1297571208359343071, language=EN, stringName=Chunyan LENG, firstName=Chunyan, middleName=null, lastName=LENG, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing, China, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null), CN=AuthorExt(id=1297571208543892450, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, authorId=1297571208359343071, language=CN, stringName=冷春燕, firstName=null, middleName=null, lastName=null, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=中国医学科学院北京协和医学院，药用植物研究所，道地药材品质保障与资源持续利用全国重点实验室，北京, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null)}, companyList=[AuthorCompany(id=1297571207998632918, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, xref=null, ext=[AuthorCompanyExt(id=1297571208011215831, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, companyId=1297571207998632918, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing, China), AuthorCompanyExt(id=1297571208019604440, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, companyId=1297571207998632918, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=中国医学科学院北京协和医学院，药用植物研究所，道地药材品质保障与资源持续利用全国重点实验室，北京)])]), Author(id=1297571208615195620, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, orderNo=2, firstName=null, middleName=null, lastName=null, nameCn=null, orcid=null, stid=null, country=null, authorPic=null, dead=0, email=null, emailSecond=null, emailThird=null, correspondingAuthor=0, authorType=1, ext={EN=AuthorExt(id=1297571208707470310, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, authorId=1297571208615195620, language=EN, stringName=Xinjie HUANG, firstName=Xinjie, middleName=null, lastName=HUANG, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing, China, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null), CN=AuthorExt(id=1297571208791356391, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, authorId=1297571208615195620, language=CN, stringName=黄信洁, firstName=null, middleName=null, lastName=null, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=中国医学科学院北京协和医学院，药用植物研究所，道地药材品质保障与资源持续利用全国重点实验室，北京, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null)}, companyList=[AuthorCompany(id=1297571207998632918, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, xref=null, ext=[AuthorCompanyExt(id=1297571208011215831, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, companyId=1297571207998632918, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing, China), AuthorCompanyExt(id=1297571208019604440, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, companyId=1297571207998632918, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=中国医学科学院北京协和医学院，药用植物研究所，道地药材品质保障与资源持续利用全国重点实验室，北京)])]), Author(id=1297571208866853865, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, orderNo=3, firstName=null, middleName=null, lastName=null, nameCn=null, orcid=null, stid=null, country=null, authorPic=null, dead=0, email=null, emailSecond=null, emailThird=null, correspondingAuthor=0, authorType=1, ext={EN=AuthorExt(id=1297571208959128555, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, authorId=1297571208866853865, language=EN, stringName=Yongmei XING, firstName=Yongmei, middleName=null, lastName=XING, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing, China, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null), CN=AuthorExt(id=1297571209022043116, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, authorId=1297571208866853865, language=CN, stringName=邢咏梅, firstName=null, middleName=null, lastName=null, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=中国医学科学院北京协和医学院，药用植物研究所，道地药材品质保障与资源持续利用全国重点实验室，北京, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null)}, companyList=[AuthorCompany(id=1297571207998632918, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, xref=null, ext=[AuthorCompanyExt(id=1297571208011215831, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, companyId=1297571207998632918, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing, China), AuthorCompanyExt(id=1297571208019604440, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, companyId=1297571207998632918, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=中国医学科学院北京协和医学院，药用植物研究所，道地药材品质保障与资源持续利用全国重点实验室，北京)])]), Author(id=1297571209080763374, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, orderNo=4, firstName=null, middleName=null, lastName=null, nameCn=null, orcid=null, stid=null, country=null, authorPic=null, dead=0, email=chenjuan@implad.ac.cn, emailSecond=null, emailThird=null, correspondingAuthor=1, authorType=1, ext={EN=AuthorExt(id=1297571209168843760, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, authorId=1297571209080763374, language=EN, stringName=Juan CHEN, firstName=Juan, middleName=null, lastName=CHEN, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing, China, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null), CN=AuthorExt(id=1297571209235952625, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, authorId=1297571209080763374, language=CN, stringName=陈娟, firstName=null, middleName=null, lastName=null, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=null, address=中国医学科学院北京协和医学院，药用植物研究所，道地药材品质保障与资源持续利用全国重点实验室，北京, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null)}, companyList=[AuthorCompany(id=1297571207998632918, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, xref=null, ext=[AuthorCompanyExt(id=1297571208011215831, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, companyId=1297571207998632918, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing, China), AuthorCompanyExt(id=1297571208019604440, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571206853587921, companyId=1297571207998632918, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=中国医学科学院北京协和医学院，药用植物研究所，道地药材品质保障与资源持续利用全国重点实验室，北京)])])]

王爱媛, 冷春燕, 黄信洁, 邢咏梅, 陈娟

作者信息

中国医学科学院北京协和医学院，药用植物研究所，道地药材品质保障与资源持续利用全国重点实验室，北京

通讯作者:

陈娟

作者简介:

作者贡献声明

王爱媛：文献收集与论文初稿撰写；冷春燕：论文修改；黄信洁：表格整理与校对；邢咏梅：文献核查与修改；陈娟：构思与最终定稿。

Chemical signals and their roles in medicinal plant-microorganism interactions

Aiyuan WANG, Chunyan LENG, Xinjie HUANG, Yongmei XING, Juan CHEN

Affiliations

State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing, China

Corresponding Author:

E-mail: chenjuan@implad.ac.cn

出版时间: 2026-08-04 doi: 10.13343/j.cnki.wsxb.20260109

文章导航

摘要

收起

药用植物与微生物间存在复杂的相互作用，互作影响着药用植物的生长发育、次生代谢产物的合成、对病原菌的防御及对环境的适应等多个方面。因此，理解作为介导二者建立联系关键纽带的化学信号物质至关重要。本文系统总结了药用植物与微生物互作过程中化学信号的类型及作用机制，分别分析了植物与微生物来源的信号物质对根际微生物群落结构、植物生长及次生代谢产物积累的影响，并探讨了双向信号在共生与竞争互作中的功能差异。尽管目前仍有许多化学信号相关科学问题有待深入研究，但可通过解析已知化学信号及其分子机制，基于“信号-微生物-功能”模式调控植物化学信号或筛选功能性微生物，重塑健康微生物区系，改善根际生态环境，促进药用植物生长，提高药用成分含量，增强对病原菌的抵抗能力。这对提升中药材质量、实现生态种植具有重要意义。

关键词

药用植物 / 根际微生物 / 化学信号 / 次生代谢产物 / 共生互作

Abstract

收起

Medicinal plants and microorganisms engage in complex, dynamic interactions that profoundly influence plant growth and development, the biosynthesis of secondary metabolites, resistance to pathogens, and adaptation to environmental stressors. Therefore, elucidating the chemical signaling molecules that mediate these interactions is of fundamental importance. The review systematically summarizes the types and functional mechanisms of chemical signals involved in medicinal plant-microorganism interactions. It highlights the roles of both plant- and microorganism-derived signaling molecules in shaping the rhizosphere microbial community assembly, regulating plant growth, and modulating the accumulation of bioactive secondary metabolites. In addition, we discuss the functional divergence of bidirectional signaling in both symbiotic and competitive interactions. Although many mechanism aspects of these chemical signals remain to be fully elucidated, current knowledge provides a solid framework for understanding their regulatory roles. With a “signal-microbiome-function” framework, it is possible to manipulate plant chemical signaling or screen functional microbial strains to reconstruct beneficial rhizosphere microbiomes. Such strategies hold potential for improving soil-plant-microorganism interactions, promoting plant growth, enhance the accumulation of medicinal compounds, and strengthen resistance to pathogens. Overall, these insights are of considerable significance for improving the quality of Chinese medicinal materials and promoting the sustainable and ecological cultivation of medicinal plants.

Key words

medicinal plant / rhizosphere microorganism / chemical signal / secondary metabolites / symbiotic interaction

引用本文

王爱媛, 冷春燕, 黄信洁, 邢咏梅, 陈娟. 药用植物-微生物互作的化学信号及其作用. 微生物学报, 2026 , 66 (8) : 3718 -3731 . DOI: 10.13343/j.cnki.wsxb.20260109

Aiyuan WANG, Chunyan LENG, Xinjie HUANG, Yongmei XING, Juan CHEN. Chemical signals and their roles in medicinal plant-microorganism interactions[J]. Acta Microbiologica Sinica, 2026 , 66 (8) : 3718 -3731 . DOI: 10.13343/j.cnki.wsxb.20260109

正文

收起

药用植物作为天然药物的宝库，其药效的物质基础源于植物体内结构多样且活性丰富的化学物质，包括初生代谢产物和次生代谢产物。初生代谢产物一般指碳水化合物、蛋白质、脂类等物质，而次生代谢产物则是以初生代谢产物为前体，在植物不同器官或组织部位产生的、非植物正常生长发育所必需的小分子有机化合物，包括生物碱、黄酮类、酚类等物质^[1]。这些物质不仅通过影响植物自身的正常生理功能以增强其对环境的适应性、抵御外界胁迫(如抑制周围杂草生长^[2]、防止昆虫啃食^[3]等)，还在植物与其他生物(特别是微生物)的互作中扮演重要角色^[4]。现代药理学研究表明，药用植物体内合成的次生代谢产物是其发挥药效的物质基础^[5]，因其活性多样而被广泛应用于医药学领域。例如，家喻户晓的抗疟药——青蒿素(artemisinin)以及临床上用于治疗癌症的紫杉醇(taxol)，均分别提取自黄花蒿和红豆杉中，属于高效、低毒、广谱的次生代谢产物^[6]。为提高药用植物的质量与产量，关注其生长发育过程至关重要，该过程与多种生态因素、生物因素等密切相关。近年来，药用植物与微生物的相互作用受到普遍关注，光、温、土、气等因素直接或间接影响土壤微生物，进而影响该区域所生长药用植物的品质。如本课题组Zhou等^[7]前期研究表明，不同产区铁皮石斛根际微生物组特征与石斛活性成分累积存在关联，证实了根际微生物群落结构对药用植物次生代谢产物积累的调控作用。因此，深入认识微生物对药用植物生长及次生代谢产物合成与积累的作用具有重要意义。

在药用植物与微生物相互作用的过程中，化学信号在植物-微生物“对话”中发挥着不可忽视的桥梁作用，是二者相互识别的第一步(图1)。化学物质的产生、释放、检测和反应均属于化学信号的组成环节^[8]。药用植物根际的各种过程，如根际效应的构建、根系分泌物的释放、根际微生物群落的组成、菌根的形成以及土壤健康与根际微生态平衡几乎都建立在植物-微生物-化学信号传导的基础之上^[9]。

系统概述药用植物-微生物之间的化学信号传导及相关化合物在二者互作中发挥的不同作用，有助于进一步阐明药用植物生长过程中的各类互作关系，为调控药用植物生长发育及次生代谢产物的合成与积累奠定基础。深入理解这些化学信号及其分子机制对未来更好地开发应用微生物肥料、生物农药，通过调控微生物群落组成提高中药材质量、实现生态种植具有重要意义。

1 植物来源的化学信号影响微生物的组成、多样性及功能

收起

植物来源的化学信号是连接宿主植物与微生物群落的关键因子，通过介导微生物招募、生态位筛选、维持定殖及功能调控等在植物相关微生物组的组装与稳态维持中发挥核心作用。这些化学信号来源广泛、类型多样，主要包括根系分泌物、挥发性有机化合物(volatile organic compounds, VOCs)以及植物激素等，不同类型信号在作用机制与功能上各具特征(表1)。

1.1 根系分泌物

根际是指植物根系与土壤及微生物相互作用的区域，包括根系表面和受根系影响的周围土壤^[26]，是药用植物与土壤间物质交换的重要场所^[27]。根系分泌物(植物生长过程中通过根系主动或被动向生长介质释放的所有物质)含有丰富的有机化合物，如酚类物质(酚酸、类黄酮等)、有机酸(柠檬酸、苹果酸、琥珀酸等)、氨基酸(谷氨酸、天冬氨酸等)及糖类(葡萄糖、果糖、蔗糖等)^[28]，不仅可作为根际微生物生长代谢的重要碳源和能源底物^[29]，选择性招募或排斥特定微生物类群可塑造根际微生态位^[30]，甚至通过化感作用抑制竞争植物的生长^[31]；还可通过调控植物免疫与抗逆相关过程提高药用植物对生物胁迫的适应能力，从而在一定程度上降低对化肥和农药的依赖。例如，有机酸作为根系分泌物的重要成分之一，可降低土壤有机质的分解速率，调控土壤碳氮矿化过程；还能通过降低根际土壤酸碱度、改变土壤氧化还原条件、影响土壤微生物群落等方式增加土壤中铁、磷及其他微量营养素的有效性^[32]。药用植物还可通过释放酚类化合物、非挥发性萜类化合物和含硫化合物，诱导产生对土壤中病原菌和害虫具有显著抑制作用的抗菌微生物，直接抑制其生长^[33]。成熟抑制剂(ripening-inhibitor, RIN)是一种调控番茄果实成熟的MADS-box转录因子，可通过改变植物根系分泌物(如3-羟基黄酮等)抑制土壤中植物病原菌茄科罗尔斯通氏菌(Ralstonia solanacearum)的积累，并促进富含抗病基因的细菌群落[如链霉菌属(Streptomyces)]在根际的聚集，增强植物抗病能力^[34]。

此外，根系分泌物在非生物胁迫响应中同样发挥关键作用，可通过内部螯合或清除活性氧(reactive oxygen species, ROS)维持细胞质或液泡中游离金属的无毒水平^[35]，建立金属螯合与隔离的防御机制^[36]以应对重金属胁迫。根系分泌物中的低分子量有机酸(low molecular weight organic acids, LMWOA)是至少含有1个羧基的小分子有机化合物，可活化土壤中无效态磷，提高磷的有效性，是药用植物应对低磷胁迫的重要手段^[37]。

1.2 挥发性有机化合物

挥发性有机化合物(volatile organic compounds, VOCs)是具有低分子量(小于300 Da)和高蒸汽压的有机化合物，种类繁多，包括醇类、醛类、酯类、萜类、硫醇等。药用植物中的VOCs是植物通过次生代谢途径产生并向周围释放的一类挥发性非甲烷碳氢化合物，地上部VOCs通过花、叶和果实排放至大气中，根系部VOCs则通过根系排放至根际^[15]。在药用植物中，VOCs参与多种生态互作，如作为化感物质吸引传粉者、防御草食动物，以及介导植物间、植物-微生物间的通讯^[38-39]。植物产生的VOCs可被微生物种群用作营养素或信号，而微生物释放的VOCs也能直接或间接地激活药用植物的免疫系统，或调节其生长和形态发生^[40]。由于其高蒸气压和低沸点，VOCs易在空气和水中扩散，使这类小分子次生代谢物能够在空气或水膜介质中实现远距离传递^[41]，即使不直接接触病原体，也可通过扰乱病原菌膜结构、干扰呼吸代谢、诱导氧化胁迫，或抑制群体感应(quorum sensing, QS)、生物膜形成及毒力基因表达等发挥抗菌作用^[42]。单萜、倍半萜等VOCs，既可通过疏水性作用破坏病原菌细胞膜完整性，又可通过调控微生物基因表达发挥抗菌作用^[43]。

1.3 植物激素及其类似物

植物激素是指由植物产生并能被自身感知的化合物，在植物与病原微生物和有益微生物的相互作用以及植物生长发育中起着重要作用^[44-45]。其中，水杨酸(salicylic acid, SA)和茉莉酸(jasmonic acid, JA)被认为是植物免疫中的主要激素，而乙烯(ethylene, ET)、独脚金内酯(strigolactones, SLs)、植物细胞分裂素(cytokinins, CK)、脱落酸(abscisic acid, ABA)和赤霉素(gibberellins, GA)等也通过激素间的串扰发挥重要作用^[46-47]。当植物遭受病原菌侵染后，SA、JA和ET等信号迅速响应，激活下游转录因子调控萜类合成相关基因的表达^[48]，进而调节植物免疫等生理过程。SLs作为关键分枝因子，能在低磷等养分限制条件下显著增加分泌量，特异性诱导丛枝菌根真菌(arbuscular mycorrhizal fungi, AMF)孢子萌发和菌丝分枝，促进AMF向根际趋化以增大与根系的接触概率，既加速了菌根共生的建立，也增强了药用植物对矿质养分的吸收能力。然而，SLs也可被某些寄生植物[如列当科(Orobanchaceae)独脚金属(Striga)植物]识别，诱导其种子萌发^[49]。GA在药用植物与微生物的共生互作中发挥关键调控作用，本课题组Chen等^[50]提出GA是石斛种子共生萌发的关键信号分子之一，并克隆鉴定了铁皮石斛赤霉素3-氧化酶(gibberellin 3-oxidases, GA3ox)基因^[51]，进一步丰富了激素类似物在药用植物-微生物共生互作中的作用机制。此外，本课题组阎波等^[52]还系统梳理了植物菌根共生中的激素调控作用，证明了GA、生长素等多种植物激素在菌根共生建立与维持中的重要地位。

1.4 防御相关次生代谢产物

尽管药用植物次生代谢产物并非植物生长发育所必需，但其在药用植物抵御外界环境胁迫、维持正常生长^[43]以及塑造微生物群落结构和调控其代谢活动方面具有重要作用^[53]。部分次生代谢产物可作为碳源和能源促进有益微生物的生长^[54]，另一些则具有抗菌活性，可抑制特定微生物的生长。人参皂苷可与根际微生物群相互作用^[55]并促进益生菌的生长^[56]。已有研究表明，人参皂苷可以刺激土壤中伯克霍尔德氏菌属(Burkholderia)的生长和生物膜形成，有助于降解自毒物质并拮抗根腐病病原体，从而缓解负向植物-土壤反馈(negative plantsoil feedback, NPSF)^[57]。此外，多数生物碱类化合物对真菌也表现出一定的抑制作用^[23]。例如，石蒜碱(lycorine, LYC)和水芹碱(narciclasine, NAR)已被证实可抑制水稻稻瘟病菌(Magnaporthe oryzae)的生长，并通过影响脂质代谢和黑色素合成等途径发挥抗真菌作用^[58]。

1.5 病程相关蛋白和抗菌肽

药用植物为应对各种不利环境而合成了多种病程相关(pathogenesis-related, PR)蛋白和抗菌肽(antimicrobial peptides, AMPs)，如凝集素、核糖体失活蛋白、蛋白酶抑制剂、尿素酶、青蒿素和致孔毒素等，在抵御捕食者和有害微生物方面发挥着关键作用^[59]。PR蛋白由多种蛋白质组成，植物受到病原菌攻击后防御信号通路SA和JA被激活，诱导PR蛋白大量积累，从而最大限度地减少病原菌负载，避免未受感染的植物器官发病^[25]。AMPs是富含半胱氨酸的分子，具有广泛的潜在抗菌活性，包括PR6蛋白家族(酶抑制剂)、PR12蛋白家族(植物防御素)、PR13蛋白家族(植物硫蛋白)和PR14蛋白家族(脂转移蛋白)^[60]，其主要靶标为微生物的细胞膜，如植物多肽可破坏其完整性^[61]。

2 微生物来源的化学信号对植物的多重影响

收起

微生物来源的化学信号是连接微生物群落与宿主植物互作的重要媒介，通过调控植物生长发育、营养吸收和免疫应答等多种生理过程，在植物适应环境变化及维持健康稳态中发挥重要作用。这些化学信号来源复杂、功能多样，主要包括激素类似物、群体感应信号分子、假单胞菌分泌物和菌根因子等，不同类型信号对植物具有多重影响(表2)。

2.1 激素类似物

微生物产生的激素类似物，如1-氨基环丙烷-1-羧酸(1-aminocyclopropane-1-carboxylic acid, ACC)脱氨酶和吲哚-3-乙酸(indole-3-acetic acid, IAA)等是重要的信号分子，能够调节药用植物的生理过程^[70-71]。ACC是植物体内ET的直接前体，由S-腺苷甲硫氨酸(S-adenosyl-L-methionine, SAM)在ACC合成酶催化下生成，随后在ACC氧化酶作用下转化为ET。逆境条件下，药用植物体内ET的合成显著增加，ET胁迫抑制植物生长^[72]。然而，部分微生物，特别是植物根际促生菌(plant growth-promoting bacteria, PGPB)能够产生ACC脱氨酶分解ACC，降低植物组织中ET水平^[62-63]，进而促进植物根和茎的生长，并延缓药用植物的衰老。研究表明，接种表达ACC脱氨酶的PGPB可提高药用植物在盐胁迫和干旱胁迫下的生物量^[63]。

IAA是一种重要的植物生长素，与植物细胞伸长、根系形成和顶端优势等生长发育过程密切相关。多种根际细菌和内生细菌均具有合成IAA的能力，主要通过吲哚-3-乙酰胺(indole-3-acetamide, IAM)途径和吲哚-3-丙酮酸(indole-3-pyruvate, IPyA)途径合成，此外还有色胺(tryptamine, TAM)途径、吲哚-3-乙腈(indole-3-acetonitrile, IAN)途径以及色氨酸侧链氧化酶(tryptophan side-chain oxidase, TSO)途径等^[73]。合成的IAA进入植物根际后可促进TIR1/AFB受体复合体对Aux/IAA抑制蛋白的识别与降解，解除其对生长素响应因子(auxin response factor, ARF)转录因子的抑制，进而激活一系列生长素响应基因的表达^[74]。IAA不仅影响药用植物的生长发育，还在植物-微生物通讯中发挥关键作用^[71]。IAA促进根系结构形成，不仅提高了水分和矿质养分的吸收效率，也显著增加了根系与AMF、外生菌根真菌(ectomycorrhizal fungi, ECMF)及其他有益微生物的接触面积，从而促进植物-微生物共生的建立及根际功能微生物的持续富集。同时，产IAA细菌可通过时空调节植物生长素库影响植物生长发育，而寄主植物也可积极参与调节细菌的IAA生物合成^[64]。

2.2 群体感应分子

药用植物生长过程中，微生物群体感应系统持续进行细胞间通讯。群体感应分子N-酰基高丝氨酸内酯(N-acyl homoserine lactones, AHLs)是革兰氏阴性菌中最常见的自诱导物^[75]，有益微生物可通过AHLs触发植物的诱导系统抗性(induced systemic resistance, ISR)，使植物对病原体入侵产生快速、强烈的免疫反应^[65]，提高植物对多种病原体的抵抗力。部分AHLs (如oxo-C14-HSL和oxo-C8-HSL)可被拟南芥识别为激活免疫反应、增强对细菌病原体[如丁香假单胞菌(Pseudomonas syringae)]抗性的危险信号^[76]。AHLs还可影响植物体内激素水平，如调节ISR通路中的重要信号分子JA和ET等，从而优化植物的免疫反应^[77]。AHLs还具有增强植物防御反应的作用，如大蒜来源的群体感应抑制化合物可通过合成AHLs类似物抑制铜绿假单胞菌(Pseudomonas aeruginosa)的毒性，并阻断黄单胞菌(Xanthomonas)的信号接收，有效控制植物病害^[78]。也有研究表明，AHLs可通过激活PAL1基因促进植物防御物质苯丙素类的合成^[79]；而缺乏AHLs信号的细菌则会增强植物对蚜虫的抵抗力^[80]。

2.3 假单胞菌分泌物

假单胞菌作为一类重要的植物根际促生菌，能够产生多种铁载体(也称为嗜铁素)和抗生素，对药用植物健康具有多重益处^[81]。铁是植物生长必需的微量元素^[82]，但在土壤中通常以难溶形式存在，难以被植物直接吸收。铁载体是假单胞菌产生的一类小分子物质，能够特异性螯合铁离子，形成可溶性铁-铁载体复合物，提高铁的溶解度和吸收率，从而增强铁的生物有效性^[83]。药用植物可利用这些复合物获取铁元素，改善营养状况、促进生长。由于铁是病原菌生长繁殖所必需的元素，铁载体可与病原菌竞争铁资源，抑制其生长，从而减轻病害，因此铁载体还具有拮抗病原菌的作用^[66]。例如，荧光假单胞菌产生的铁载体能够有效抑制多种植物病原真菌和细菌的生长^[83]。此外，假单胞菌还能产生多种具有抗菌活性的抗生素，如吩嗪、2,4-二乙酰基根皮苷等，可直接抑制植物病原菌的生长繁殖，减轻病害^[84]。

3 双向信号

收起

3.1 共生关系的建立(菌根共生)

根际微生物组中某些真菌与药用植物根系可形成共生体，称为药用植物的菌根^[85]。菌根真菌的菌丝体既向根周土壤扩展，又与寄主植物组织相连通，一方面从寄主植物中吸收糖类等有机物质作为自身营养，另一方面从土壤中吸收养分和水分供给植物。菌根的存在有利于药用植物扩大根系吸收面积，增强对原根毛吸收范围外元素的吸收能力^[86]。菌根类型通常根据真菌在药用植物中的定殖方式分为丛枝菌根(arbuscular mycorrhiza, AM)、外生菌根(ectomycorrhiza, ECM)、类菌根(ericoid mycorrhiza, ERM)和兰花菌根(orchid mycorrhiza, ORM)^[87]。研究表明，不同菌根类型对不同药用植物的影响各异，如AM可促进墨旱莲地上部分药物活性成分的积累，ECM可提高油松的抗旱性等^[6]。

分布最广的菌根类型是AM，在药用植物保护中发挥着重要作用，包括改善药用植物营养、损害补偿、争夺定殖位点或光合产物、促进根系和根际微生物种群变化以及激活植物防御机制等^[88-89]。在共生关系形成前期，AMF与植物之间识别与联系的化学信号主要来源于植物根系分泌物中的SLs等信号因子，以及AMF分泌的结瘤因子(lipochitooligosaccharides, LCOs)和短链几丁质寡聚物(short-chain chitin oligomers, COs)等菌根因子^[90]。在植物生长发育过程中，根系分泌SLs，特异性吸引土壤中的AMF，诱导其孢子萌发和菌丝分枝，进而增加AMF与根系的接触概率^[19]。AMF感知SLs后会分泌LCOs、COs等信号分子，被植物根细胞表面的LysM受体样激酶3 (LysM receptor-like kinase, LYK3)/结瘤因子感知蛋白(nod factor perception, NFP)类受体识别并激活，随后触发细胞内核周钙离子周期性振荡，通过钙调蛋白互作蛋白(calcium/calmodulin-dependent protein kinase, CCaMK)解码信号^[91]，激活下游转录因子(如RAM1、DELLA)，直接调控菌丝附着体形成、根皮层细胞重编程及丛枝结构发育相关基因的表达，最终形成菌丝侵入植物细胞的共生关系，该过程无需其他辅助信号通路参与^[92]。

同时，AMF也是参与土壤磷活化的重要微生物类群之一^[37]。AMF与细菌共同参与凋落物降解，有助于增强土壤有机碳(soil organic carbon, SOC)的螯合能力并减少氮淋溶^[93]；AMF的存在还可减少养分损失，如降低硝酸盐浸出^[94]。此外，根系分泌物中多种次生代谢产物(如脱落酸、茉莉酸、水杨酸、油菜素内酯等)也被发现有助于植物与AMF建立共生关系，这些物质可与植物激素产生互作，协同调节植物生长发育及应对各类生物和非生物胁迫^[95]。AMF还可通过增加植物生物量直接影响次生代谢物的产生，或通过刺激次生代谢物生物合成途径间接调控其积累^[89]。

另一种被广泛研究的菌根类型是ECM，目前仅在约15种药用植物中发现^[96]。外生菌根真菌具有一定的宿主选择性，已知能够影响外生菌根形成的根系分泌物质包括酯类、黄酮类及植物激素^[97]。植物根系通过分泌这些物质作为招募ECMF的化学信号，ECMF则分泌COs，与植物根细胞表面高特异性的几丁质触发子受体激酶(chitin elicitor receptor kinase 1, CERK1)类受体结合并被识别^[98]，进而激活钙离子振荡-CCaMK保守通路。同时，ECMF分泌效应蛋白(如MiSSP7)进入植物细胞后与宿主转录因子(如JAZ)发生互作^[99]，启动宿主特异性调控分支通路，抑制植物产生防御反应以清除真菌。此过程还需生长素、茉莉酸等植物激素通路的协同调控^[100]。

本课题组Chen等^[101]通过比较转录组学分析铁皮石斛与2种菌根真菌[胶膜菌(Tulasnella sp.) S6和无孢蜡壳菌(Serendipita sp.) 12825]共生萌发的分子机制，发现共生萌发显著诱导宿主糖脂代谢、物质转运、激素信号及细胞壁重塑相关基因的表达，并伴随真菌碳水化合物活性酶(carbohydrate-active enzymes, CAZymes)和分泌相关基因的协同响应，表明宿主细胞壁重构与真菌定殖过程是兰科植物共生萌发建立的关键分子基础，为进一步探索植物共生萌发和菌根共生的分子机制提供了重要数据。

3.2 植物与有益/病原微生物互作过程中化学信号的异同

通过对大量研究的总结分析，植物-微生物互作过程中化学信号的差异性主要体现在性质、功能及通路等方面^[102-103]。植物与病原菌竞争互作过程中主要涉及植物防御素^[104]、病原菌毒素^[105]等具有毒性或抑制性的化学信号，驱动植物与病原菌之间的“化学竞争”；而植物与有益微生物的共生互作过程中主要涉及植物根系分泌物、微生物产生的生长素和细胞分裂素等具有营养或促进生长功能的化学信号，促进二者互利互惠共生关系的建立^[106]。

在不同的植物-微生物互作过程中，不同化学信号具有多样的功能。在竞争过程中，化学信号的主要功能是促进病原菌侵染和植物防御，病原菌利用化学信号寻找寄主、释放毒素，而植物则利用化学信号识别病原菌、激活防御反应；在共生互作过程中，化学信号的主要功能是促进双方识别、营养交换和协同生长。在信号通路方面，竞争过程中植物的防御反应通常涉及SA、JA、ET等信号通路，通过激活这些通路表达防御基因以增强抗病性；而共生互作过程中通常涉及生长素、细胞分裂素等植物激素的调控，通过复杂通路调控根际发育、细胞分裂与分化，提高养分吸收效率，协调植物与共生微生物的生长，建立稳定的共生关系。

植物-微生物互作过程中化学信号种类繁多，信号传递受环境因素(如光、温、水分、养分)的影响，且互作过程均涉及信号识别与响应的共性。因此，无论竞争互作还是共生互作，化学信号均是植物与微生物之间进行识别、通讯和互动的基础。

4 总结与展望

收起

植物与微生物之间的互作关系复杂，化学信号在其中扮演着至关重要的角色。本课题组前期围绕铁皮石斛、天麻等兰科药用植物，在内生及菌根真菌资源收集与鉴定、菌根共生专一性与植物发育阶段的关系、兰科种子-真菌互作理论、促进种子萌发的分子机制及关键信号分子赤霉素、优势菌群(Tulasnella、Serendipita)的鉴定等方面取得了系统成果，为理解植物-微生物互作关系提供了重要支撑，因此探究介导二者互作的化学信号具有重要意义。特别是在药用植物中，化学信号的调节直接影响植物健康、生长及药用成分的合成^[9]。植物来源的信号(如根系分泌物、挥发性化合物、防御相关次生代谢产物等)能影响根际微生物区系的组成与活性；微生物来源的信号(如激素类似物、群体感应分子、铁载体、抗生素等)则能调节药用植物的生理过程，诱导其产生抗性，增强对病原菌的防御能力；而双向信号能够促进植物与有益微生物的识别与共生，调控根际微生物群落结构与功能，激活植物免疫系统，使植物在受到病原菌侵染时释放化学信号以启动自身免疫反应。

尽管目前仍有许多化学信号及其作用机制有待进一步解析，但已可基于“信号-微生物-功能”模式尝试解决各类土壤健康及植物生长问题。未来可通过调控各类化学信号重塑健康的微生物区系；人为筛选具有降解自毒物质、抑制病原菌等功能的微生物，构建人工微生物群落，实现对根际微生态环境的精准调控；利用合成生物学等手段外源添加或遗传改良，调控植物根系分泌的化学信号，吸引有益微生物、抑制有害微生物；将微生物代谢产生的有益物质(如植物激素、维生素和有机酸等)应用于植物种植生产，促进植物生长并提高抗逆性。这将有助于实现中药材乃至农作物的生态种植，推动微生物肥料及生物农药的开发，同时为提高中药材质量和实现可持续农业生产提供参考。

基金

收起

国家自然科学基金(82173923)

参考文献

收起

文献

收起

参考文献引证文献

排序方式：

[1]

Nelson

, Gelambi

, Morales

, Whitehead

. Fruit secondary metabolites alter the quantity and quality of a seed dispersal mutualism[J]. Ecology, 2023, 104(5): e4032.

[2]

Zhan

, Chen

, Shen

. Environmental and genetic factors involved in plant protection-associated secondary metabolite biosynthesis pathways[J]. Frontiers in Plant Science, 2022, 13: 877304.

[3]

Musilova

, Ridl

, Polivkova

, Macek

, Uhlik

. Effects of secondary plant metabolites on microbial populations: changes in community structure and metabolic activity in contaminated environments[J]. International Journal of Molecular Sciences, 2016, 17(8): 1205.

[4]

Zheng

, Luo

, Zhou

, Chen

. Natural products from plants and microorganisms: novel therapeutics for chronic kidney disease via gut microbiota regulation[J]. Frontiers in Pharmacology, 2023, 13: 1068613.

[5]

Buenz

, Verpoorte

, Bauer

. The ethnopharmacologic contribution to bioprospecting natural products[J]. Annual Review of Pharmacology and Toxicology, 2018, 58: 509-530.

[6]

Weaver

. How taxol/paclitaxel kills cancer cells[J]. Molecular Biology of the Cell, 2014, 25(18): 2677-2681.

[7]

Zhou

, Hou

, Leng

, Li

, Xing

, Chen

. Deciphering the root microbiome and its relationship with active compound accumulation in medicinal Dendrobium officinale (Orchidaceae) from different regions[J]. Industrial Crops and Products, 2025, 226: 120692.

[8]

Abdul Hamid

, Nadarajah

. Microbe related chemical signalling and its application in agriculture[J]. International Journal of Molecular Sciences, 2022, 23(16): 8998.

[9]

Chagas

, de Cassia Pessotti

, Caraballo-Rodríguez

, Pupo

. Chemical signaling involved in plant-microbe interactions[J]. Chemical Society Reviews, 2018, 47(5): 1652-1704.

[10]

Zhang

, Wei

, Li

, Hu

, Zhao

, Wu

, Yang

, Li

, Zhou

. Rhizosphere microbiome and phenolic acid exudation of the healthy and diseased American ginseng were modulated by the cropping history[J]. Plants, 2023, 12(16): 2993.

[11]

, Cheng

, Luk

, Leung

, Chan

, Lam

. Deciphering metabolite signalling between plant roots and soil pathogens to design resistance[J]. BMC Plant Biology, 2025, 25: 308.

[12]

Jiang

, Li

, Wang

, Lv

, Jiang

, Cao

, Qu

, Ma

, Zhang

. Exogenous application of Bradyrhizobium japonicum AC20 enhances soybean tolerance to atrazine via regulating rhizosphere soil microbial community and amino acid, carbohydrate metabolism related genes expression[J]. Plant Physiology and Biochemistry, 2023, 196: 472-483.

[13]

Afridi

, Kumar

, Javed

, Dubey

, de Medeiros

FHV

, Santoyo

. Harnessing root exudates for plant microbiome engineering and stress resistance in plants[J]. Microbiological Research, 2024, 279: 127564.

[14]

. Integration of multiple volatile cues into plant defense responses[J]. New Phytologist, 2022, 233(2): 618-623.

[15]

Abbas

, O’Neill Rothenberg

, Zhou

, Ke

, Wang

. Volatile organic compounds as mediators of plant communication and adaptation to climate change[J]. Physiologia Plantarum, 2022, 174(6): e13840.

[16]

Janda

, Szalai

, Pál

. Salicylic acid signalling in plants[J]. International Journal of Molecular Sciences, 2020, 21(7): 2655.

[17]

Ruan

, Zhou

, Yan

, Khurshid

, Weng

, Cheng

, Zhang

. Jasmonic acid signaling pathway in plants[J]. International Journal of Molecular Sciences, 2019, 20(10): 2479.

[18]

Chen

, Zhang

, Chen

, Gao

, Xu

, Sun

, Huo

. The role of ethylene in the regulation of plant response mechanisms to waterlogging stress[J]. Plant Cell Reports, 2024, 43(12): 278.

[19]

Naseer

, Zhang

, Mukhtar

, Asad

, Wu

, Yang

, Zhou

. Strigolactones: a promising tool for nutrient acquisition through arbuscular mycorrhizal fungi symbiosis and abiotic stress tolerance[J]. Plant Physiology and Biochemistry, 2024, 215: 109057.

[20]

Kieber

, Schaller

. Cytokinin signaling in plant development[J]. Development, 2018, 145(4): dev149344.

[21]

Nguyen

, Yan

, Nambara

. Persistence of abscisic acid analogs in plants: chemical control of plant growth and physiology[J]. Genes, 2023, 14(5): 1078.

[22]

Gan

, Wang

, Yang

, Ma

. Molecular decoding of phytohormone crosstalk: JA-mediated key regulatory nodes and signal integration[J]. Plants, 2025, 14(17): 2647.

[23]

Wang

, Mo

, Tan

, Wu

, Pan

, Chen

. Effects of growing Coptis chinensis Franch in the natural understory vs. under a manmade scaffold on its growth, alkaloid contents, and rhizosphere soil microenvironment[J]. PeerJ, 2022, 10: e13676.

[24]

Javed

, Wang

, Yang

, Shen

, Sun

, Gao

, Zhang

. Pathogenesis related-1 proteins in plant defense: regulation and functional diversity[J]. Critical Reviews in Biotechnology, 2025, 45(2): 305-313.

[25]

Ali

, Ahmad Ganai

, Kamili

, Ali Bhat

, Ahmad Mir

, Bhat

, Tyagi

, Islam

, Mushtaq

, Yadav

, Rawat

, Grover

. Pathogenesis-related proteins and peptides as promising tools for engineering plants with multiple stress tolerance[J]. Microbiological Research, 2018, 212: 29-37.

[26]

李永芹, 周思彤, 许乐乐, 王锂韫. 健康与患病穿心莲根际土壤理化性质及微生物多样性分析[J]. 中国实验方剂学杂志, 2025, 31(7): 172-181.

, Zhou

, Xu

, Wang

. Microbial diversity and physicochemical properties of rhizosphere soil of healthy and diseased Andrographis paniculata [J]. Chinese Journal of Experimental Traditional Medical Formulae, 2025, 31(7): 172-181 (in Chinese).

[27]

吕柏辰, 孙海, 钱佳奇, 梁浩, 朱家鹏, 张强胜, 邵财, 张亚玉. 药用植物根系分泌物与根际微生物相互作用及其在中药材生态种植中的应用[J]. 中国中药杂志, 2024, 49(8): 2128-2137.

Lyu

, Sun

, Qian

, Liang

, Zhu

, Zhang

, Shao

, Zhang

. Interaction between root exudates of medicinal plants and rhizosphere microorganisms and its application in ecological planting of Chinese medicinal materials[J]. China Journal of Chinese Materia Medica, 2024, 49(8): 2128-2137 (in Chinese).

[28]

, Sun

, Zhu

. The role of artificial root exudate components in facilitating the degradation of pyrene in soil[J]. Scientific Reports, 2017, 7: 7130.

[29]

Jacoby

, Kopriva

. Metabolic niches in the rhizosphere microbiome: new tools and approaches to analyse metabolic mechanisms of plant-microbe nutrient exchange[J]. Journal of Experimental Botany, 2019, 70(4): 1087-1094.

[30]

Yang

, Chen

, Hong

, Wang

, Wu

, Tang

, Gao

, Hao

. Plant exudates-driven microbiome recruitment and assembly facilitates plant health management[J]. FEMS Microbiology Reviews, 2025, 49: fuaf008.

[31]

Luo

, Zhang

, Ye

, Li

, Duan

, Wang

, Huang

, Liu

, Deng

, Mei

, He

, Yang

, Zhu

. Foliar pathogen infection manipulates soil health through root exudate-modified rhizosphere microbiome[J]. Microbiology Spectrum, 2022, 10(6): e02418-22.

[32]

Guerrieri

, Dong

, Bouwmeester

. Role and exploitation of underground chemical signaling in plants[J]. Pest Management Science, 2019, 75(9): 2455-2463.

[33]

Jin

, Li

, Yu

, Zheng

. Plant Fe status affects the composition of siderophore-secreting microbes in the rhizosphere[J]. Annals of Botany, 2010, 105(5): 835-841.

[34]

Yang

, Fu

, Feng

, Jiang

, Finkel

, Sun

, Liu

, Huang

, Li

, Wang

, Yang

, Wang

, Xu

, Shen

, Friman

, Jousset

, Wei

. RIN enhances plant disease resistance via root exudate-mediated assembly of disease-suppressive rhizosphere microbiota[J]. Molecular Plant, 2023, 16(9): 1379-1395.

[35]

王雨菡, 陈莲, 张培珍, 王振江, 林森, 唐翠明, 罗国庆, 钟建武, 李智毅, 王圆. 根系分泌物与根际微生物对土壤重金属污染的响应与修复作用[J]. 江苏农业科学, 2024, 52(5): 19-27.

Wang

, Chen

, Zhang

, Wang

, Lin

, Tang

, Luo

, Zhong

, Li

. Response and remediation of root exudates and rhizosphere microorganisms to soil heavy metal pollution: a review[J]. Jiangsu Agricultural Sciences, 2024, 52(5): 19-27 (in Chinese).

[36]

沈川, 李夏. 根系分泌物对不同环境胁迫响应的研究进展[J]. 核农学报, 2024, 38(11): 2247-2257.

Shen

, Li

. Research progress on responses of root exudates to different environmental stresses[J]. Journal of Nuclear Agricultural Sciences, 2024, 38(11): 2247-2257 (in Chinese).

[37]

黄雨轩, 游欣, 张林平, 吴斐, 张扬, 黄绍华. 根系分泌物提高土壤磷有效性研究概述[J]. 林业科学研究, 2024, 37(4): 193-203.

Huang

, You

, Zhang

, Wu

, Zhang

, Huang

. Root exudates improving soil phosphorus availability: a review[J]. Forest Research, 2024, 37(4): 193-203 (in Chinese).

[38]

Bergman

, Davis

, Phillips

. Medically useful plant terpenoids: biosynthesis, occurrence, and mechanism of action[J]. Molecules, 2019, 24(21): 3961.

[39]

Brosset

, Blande

. Volatile-mediated plant-plant interactions: volatile organic compounds as modulators of receiver plant defence, growth, and reproduction[J]. Journal of Experimental Botany, 2022, 73(2): 511-528.

[40]

Abbas

, Wang

. Plant volatile organic compounds: revealing the hidden interactions[J]. Plants, 2025, 14(4): 507.

[41]

Abd-Elgawad

. Integrated nematode management strategies: optimization of combined nematicidal and multi-functional inputs[J]. Plants, 2025, 14(7): 1004.

[42]

D’Aquila

, Sena

, Crudo

, Passarino

, Bellizzi

. Effect of essential oils of Apiaceae, Lamiaceae, Lauraceae, Myrtaceae, and Rutaceae family plants on growth, biofilm formation, and quorum sensing in Chromobacterium violaceum, Pseudomonas aeruginosa, and Enterococcus faecalis [J]. Microorganisms, 2023, 11(5): 1150.

[43]

Kawasaki

, Dennis

, Forstner

, Raghavendra

AKH

, Mathesius

, Richardson

, Delhaize

, Gilliham

, Watt

, Ryan

. Manipulating exudate composition from root apices shapes the microbiome throughout the root system[J]. Plant Physiology, 2021, 187(4): 2279-2295.

[44]

Nakano

, Omae

, Tsuda

. Inter-organismal phytohormone networks in plant-microbe interactions[J]. Current Opinion in Plant Biology, 2022, 68: 102258.

[45]

, Duan

, Luo

, Dai

, Ding

, Xia

. How plant hormones mediate salt stress responses[J]. Trends in Plant Science, 2020, 25(11): 1117-1130.

[46]

Berens

, Berry

, Mine

, Argueso

, Tsuda

. Evolution of hormone signaling networks in plant defense[J]. Annual Review of Phytopathology, 2017, 55: 401-425.

[47]

Aerts

, Pereira Mendes

, Van Wees

SCM

. Multiple levels of crosstalk in hormone networks regulating plant defense[J]. The Plant Journal, 2021, 105(2): 489-504.

[48]

Schmelz

, Huffaker

, Sims

, Christensen

, Lu

, Okada

, Peters

. Biosynthesis, elicitation and roles of monocot terpenoid phytoalexins[J]. The Plant Journal, 2014, 79(4): 659-678.

[49]

Clark

, Bennett

. Cracking the Enigma: understanding strigolactone signalling in the rhizosphere[J]. Journal of Experimental Botany, 2024, 75(4): 1159-1173.

[50]

Chen

, Yan

, Tang

, Xing

, Li

, Zhou

, Guo

. Symbiotic and asymbiotic germination of Dendrobium officinale (Orchidaceae) respond differently to exogenous gibberellins[J]. International Journal of Molecular Sciences, 2020, 21(17): 6104.

[51]

刘思思, 张岗, 陈晓梅, 李淑超, 陈娟, 郭顺星. 铁皮石斛赤霉素3-氧化酶基因的克隆及表达分析[J]. 中草药, 2016, 47(6): 990-996.

Liu

, Zhang

, Chen

, Li

, Chen

, Guo

. Cloning and expression analysis of gibberellin 3-oxidase gene in Dendrobium officinale [J]. Chinese Traditional and Herbal Drugs, 2016, 47(6): 990-996 (in Chinese).

[52]

阎波, 陈娟, 郭顺星. 植物菌根共生中的激素调控作用研究进展[J]. 植物生理学报, 2017, 53(6): 916-924.

Yan

, Chen

, Guo

. Advances in regulating effects of hormones in mycorrhizal symbiosis[J]. Plant Physiology Journal, 2017, 53(6): 916-924 (in Chinese).

[53]

Yang

, Zhang

, Yang

, Xiang

, Yang

. Plant secondary metabolites-mediated plant defense against bacteria and fungi pathogens[J]. Plant Physiology and Biochemistry, 2024, 217: 109224.

[54]

Ketehouli

, Sossah

, Panwala

, Suazo Tejada

, Goss

, Garcia

FHS

, Vallad

, Martins

. Secondary metabolites in plant-microbe interactions[J]. Journal of Applied Microbiology, 2025, 136(6): lxaf124.

[55]

Natasha, Shahid

, Niazi

, Khalid

, Murtaza

, Bibi

, Rashid

. A critical review of selenium biogeochemical behavior in soil-plant system with an inference to human health[J]. Environmental Pollution, 2018, 234: 915-934.

[56]

, Guo

, Yu

, Du

. Application of microorganisms in Panax ginseng: cultivation of plants, and biotransformation and bioactivity of key component ginsenosides[J]. Archives of Microbiology, 2024, 206(11): 433.

[57]

Luo

, Wang

, Deng

, Mei

, Liu

, Huang

, Du

, Zhu

, Yang

. Enrichment of Burkholderia in the rhizosphere by autotoxic ginsenosides to alleviate negative plant-soil feedback[J]. Microbiology Spectrum, 2021, 9(3): e01400-21.

[58]

Qiao

, Tian

, Chen

, Liu

, Xu

, Wang

, Feng

, Chen

, Xu

. Antifungal mechanisms of amaryllidaceous alkaloids lycorine and narciclasine against the rice blast fungus Magnaporthe oryzae [J]. Pest Management Science, 2026, 82(2): 1470-1480.

[59]

Dang

, Van Damme

EJM

. Toxic proteins in plants[J]. Phytochemistry, 2015, 117: 51-64.

[60]

Deo

, Turton

, Kainth

, Kumar

, Wieden

. Strategies for improving antimicrobial peptide production[J]. Biotechnology Advances, 2022, 59: 107968.

[61]

Slavokhotova

, Shelenkov

, Andreev

, Odintsova

. Hevein-like antimicrobial peptides of plants[J]. Biochemistry (Moscow), 2017, 82(13): 1659-1674.

[62]

Singh

, Ma

, Shadan

. Perspective of ACC-deaminase producing bacteria in stress agriculture[J]. Journal of Biotechnology, 2022, 352: 36-46.

[63]

Tiwari

, Duraivadivel

, Sharma

, Hariprasad

. 1-aminocyclopropane-1-carboxylic acid deaminase producing beneficial rhizobacteria ameliorate the biomass characters of Panicum maximum Jacq. by mitigating drought and salt stress[J]. Scientific Reports, 2018, 8: 17513.

[64]

Etesami

, Glick

. Bacterial indole-3-acetic acid: a key regulator for plant growth, plant-microbe interactions, and agricultural adaptive resilience[J]. Microbiological Research, 2024, 281: 127602.

[65]

Liu

, Zhao

, Jia

, Song

, Huang

, Ma

, Song

. N-3-oxo-octanoyl-homoserine lactone-mediated priming of resistance to Pseudomonas syringae requires the salicylic acid signaling pathway in Arabidopsis thaliana [J]. BMC Plant Biology, 2020, 20: 38.

[66]

Hider

, Kong

. Chemistry and biology of siderophores[J]. Natural Product Reports, 2010, 27(5): 637.

[67]

Maillet

, Poinsot

, André

, Puech-Pagès

, Haouy

, Gueunier

, Cromer

, Giraudet

, Formey

, Niebel

, Martinez

, Driguez

, Bécard

, Dénarié

. Fungal lipochitooligosaccharide symbiotic signals in arbuscular mycorrhiza[J]. Nature, 2011, 469(7328): 58-63.

[68]

Volpe

, Chialva

, Mazzarella

, Crosino

, Capitanio

, Costamagna

, Kohlen

, Genre

. Long-lasting impact of chitooligosaccharide application on strigolactone biosynthesis and fungal accommodation promotes arbuscular mycorrhiza in Medicago truncatula [J]. New Phytologist, 2023, 237(6): 2316-2331.

[69]

Plett

, Daguerre

, Wittulsky

, Vayssières

, Deveau

, Melton

, Kohler

, Morrell-Falvey

, Brun

, Veneault-Fourrey

, Martin

. Effector MiSSP7 of the mutualistic fungus Laccaria bicolor stabilizes the Populus JAZ6 protein and represses jasmonic acid (JA) responsive genes[J]. Proceedings of the National Academy of Sciences of the United States of America, 2014, 111(22): 8299-8304.

[70]

Glick

, Nascimento

. Pseudomonas 1-aminocyclopropane-1-carboxylate (ACC) deaminase and its role in beneficial plant-microbe interactions[J]. Microorganisms, 2021, 9(12): 2467.

[71]

Gamalero

, Lingua

, Glick

. Ethylene, ACC, and the plant growth-promoting enzyme ACC deaminase[J]. Biology, 2023, 12(8): 1043.

[72]

Cao

, Narayanan

, Shi

, Chen

, Li

, Ma

. Optimistic contributions of plant growth-promoting bacteria for sustainable agriculture and climate stress alleviation[J]. Environmental Research, 2023, 217: 114924.

[73]

Tang

, Li

, Zhang

, Mu

, Jiang

, Fu

, Zhang

, Cui

, Yu

, Ye

. Biosynthetic pathways and functions of indole-3-acetic acid in microorganisms[J]. Microorganisms, 2023, 11(8): 2077.

[74]

Tena

. TIR1 and AFB1, so close yet so different[J]. Nature Plants, 2023, 9(8): 1171.

[75]

Joshi

, Khazanov

, Charkowski

, Faigenboim

, Senderowitz

, Yedidia

. Interkingdom signaling interference: the effect of plant-derived small molecules on quorum sensing in plant-pathogenic bacteria[J]. Annual Review of Phytopathology, 2021, 59: 153-190.

[76]

Zheng

, Liu

, Wang

. Quorum signaling molecules: interactions between plants and associated pathogens[J]. International Journal of Molecular Sciences, 2025, 26(11): 5235.

[77]

, Shao

, Zheng

, Sun

, Shi

, Yu

, Qi

, Shi

. Induction of systemic resistance in tomato against Botrytis cinerea by N-decanoyl-homoserine lactone via jasmonic acid signaling[J]. Planta, 2018, 247(5): 1217-1227.

[78]

Zhang

, Zhao

, Zhu

, Guan

, Xiong

, He

, Li

, Shi

, Xu

, Deng

, Cui

. Design, synthesis, and biological evaluation of asymmetrical disulfides based on garlic extract as Pseudomonas aeruginosa pqs quorum sensing inhibitors[J]. Journal of Agricultural and Food Chemistry, 2025, 73(10): 5850-5859.

[79]

Kavil

, Otti

, Bouvaine

, Armitage

, Maruthi

. PAL1 gene of the phenylpropanoid pathway increases resistance to the Cassava brown streak virus in cassava[J]. Virology Journal, 2021, 18: 184.

[80]

Sanchez-Mahecha

, Klink

, Heinen

, Rothballer

, Zytynska

. Impaired microbial N-acyl homoserine lactone signalling increases plant resistance to aphids across variable abiotic and biotic environments[J]. Plant, Cell & Environment, 2022, 45(10): 3052-3069.

[81]

Trivedi

, Leach

, Tringe

, Sa

, Singh

. Plant-microbiome interactions: from community assembly to plant health[J]. Nature Reviews Microbiology, 2020, 18(11): 607-621.

[82]

Soares

. Perspective on the biotechnological production of bacterial siderophores and their use[J]. Applied Microbiology and Biotechnology, 2022, 106(11): 3985-4004.

[83]

Luján

, Gómez

, Buckling

. Siderophore cooperation of the bacterium Pseudomonas fluorescens in soil[J]. Biology Letters, 2015, 11(2): 20140934.

[84]

Rani

, Rana

, Dhaka

, Singh

, Chahar

, Singh

, Nain

, Singh

, Minz

. Bacterial volatile organic compounds as biopesticides, growth promoters and plant-defense elicitors: current understanding and future scope[J]. Biotechnology Advances, 2023, 63: 108078.

[85]

Shi

, Wang

. Mycorrhizal symbiosis in plant growth and stress adaptation: from genes to ecosystems[J]. Annual Review of Plant Biology, 2023, 74: 569-607.

[86]

顾元钦, 郑健云, 黄瑾, 胡静雯, 丁妹, 杨帆, 杜勤. 丛枝菌根真菌对穿心莲生长及有效成分的影响[J]. 广西植物, 2025, 45(2): 377-388.

, Zheng

, Huang

, Hu

, Ding

, Yang

, Du

. Effects of arbuscular mycorrhizal fungi on growth and active ingredients of Andrographis paniculata [J]. Guihaia, 2025, 45(2): 377-388 (in Chinese).

[87]

Brundrett

, Tedersoo

. Evolutionary history of mycorrhizal symbioses and global host plant diversity[J]. New Phytologist, 2018, 220(4): 1108-1115.

[88]

Zeng

, Guo

, Chen

, Hao

, Wang

, Huang

, Yang

, Cui

, Yang

, Wu

, Chen

, Zhang

. Arbuscular mycorrhizal symbiosis and active ingredients of medicinal plants: current research status and prospectives[J]. Mycorrhiza, 2013, 23(4): 253-265.

[89]

Zhao

, Cartabia

, Lalaymia

, Declerck

. Arbuscular mycorrhizal fungi and production of secondary metabolites in medicinal plants[J]. Mycorrhiza, 2022, 32(3/4): 221-256.

[90]

侯时季, 陈保冬, 张莘. 丛枝菌根共生建成的信号识别机制[J]. 微生物学通报, 2016, 43(12): 2693-2699.

Hou

, Chen

, Zhang

. Signal recognition mechanism in establishing arbuscular mycorrhiza symbiosis[J]. Microbiology China, 2016, 43(12): 2693-2699 (in Chinese).

[91]

Shimoda

, Han

, Yamazaki

, Suzuki

, Hayashi

, Imaizumi-Anraku

. Rhizobial and fungal symbioses show different requirements for calmodulin binding to calcium calmodulin-dependent protein kinase in Lotus japonicus [J]. The Plant Cell, 2012, 24(1): 304-321.

[92]

MacLean

, Bravo

, Harrison

. Plant signaling and metabolic pathways enabling arbuscular mycorrhizal symbiosis[J]. The Plant Cell, 2017, 29(10): 2319-2335.

[93]

Gao

, Bezemer

, de Vries

, van Bodegom

. Trade-offs in soil microbial functions and soil health in agroecosystems[J]. Trends in Ecology & Evolution, 2024, 39(10): 895-903.

[94]

李彦林, 陈杨洋, 杨霜溶, 刘菊梅. 植物根系分泌的有机酸对土壤碳氮矿化的影响[J]. 生态环境学报, 2024, 33(9): 1362-1371.

, Chen

, Yang

, Liu

. Study on the effects of organic acids in plant root exudates on soil organic carbon and nitrogen mineralization[J]. Ecology and Environmental Sciences, 2024, 33(9): 1362-1371 (in Chinese).

[95]

廖德华, 刘俊丽, 刘健健, 杨晓峰, 陈潇, 顾冕, 陈爱群. 植物激素响应和调控丛枝菌根共生研究进展[J]. 植物营养与肥料学报, 2016, 22(6): 1679-1689.

Liao

, Liu

, Yang

, Chen

, Gu

, Chen

. Advances in the response and modulation of phytohormones on arbuscular mycorrhizal symbiosis[J]. Journal of Plant Nutrition and Fertilizers, 2016, 22(6): 1679-1689 (in Chinese).

[96]

Zhang

, Shi

, Zhang

, Gao

. A database on mycorrhizal traits of Chinese medicinal plants[J]. Frontiers in Plant Science, 2022, 13: 840343.

[97]

Garcia

, Delaux

, Cope

, Ané

. Molecular signals required for the establishment and maintenance of ectomycorrhizal symbioses[J]. New Phytologist, 2015, 208(1): 79-87.

[98]

Cope

, Bascaules

, Irving

, Venkateshwaran

, Maeda

, Garcia

, Rush

, Ma

, Labbé

, Jawdy

, Steigerwald

, Setzke

, Fung

, Schnell

, Wang

, Schleif

, Bücking

, Strauss

, Maillet

, Jargeat

, et al. The ectomycorrhizal fungus Laccaria bicolor produces lipochitooligosaccharides and uses the common symbiosis pathway to colonize Populus roots[J]. The Plant Cell, 2019, 31(10): 2386-2410.

[99]

Daguerre

, Basso

, Hartmann-Wittulski

, Schellenberger

, Meyer

, Bailly

, Kohler

, Plett

, Martin

, Veneault-Fourrey

. The mutualism effector MiSSP7 of Laccaria bicolor alters the interactions between the poplar JAZ6 protein and its associated proteins[J]. Scientific Reports, 2020, 10: 20362.

[100]

Basso

, Kohler

, Miyauchi

, Singan

, Guinet

, Šimura

, Novák

, Barry

, Amirebrahimi

, Block

, Daguerre

, Na

, Grigoriev

, Martin

, Veneault-Fourrey

. An ectomycorrhizal fungus alters sensitivity to jasmonate, salicylate, gibberellin, and ethylene in host roots[J]. Plant, Cell & Environment, 2020, 43(4): 1047-1068.

[101]

Chen

, Tang

, Kohler

, Lebreton

, Xing

, Zhou

, Li

, Martin

, Guo

. Comparative transcriptomics analysis of the symbiotic germination of D. officinale (Orchidaceae) with emphasis on plant cell wall modification and cell wall-degrading enzymes[J]. Frontiers in Plant Science, 2022, 13: 880600.

[102]

Tan

, Nian

, Tran

, Jin

, Lian

. Small peptides: novel targets for modulating plant-rhizosphere microbe interactions[J]. Trends in Microbiology, 2024, 32(11): 1072-1083.

[103]

Venturi

, Keel

. Signaling in the rhizosphere[J]. Trends in Plant Science, 2016, 21(3): 187-198.

[104]

Majumdar

, Sharma

, Belludi

. Natural and engineered resistance mechanisms in plants against phytoviruses[J]. Pathogens, 2023, 12(4): 619.

[105]

Khan

MSS

, Islam

, Chen

. A fungal effector hijacks a plastid protein to dampen plant immunity; PR1 is here for rescue[J]. Stress Biology, 2025, 5: 23.

[106]

Pieterse

CMJ

, Zamioudis

, Berendsen

, Weller

, Van Wees

SCM

, Bakker

PAHM

. Induced systemic resistance by beneficial microbes[J]. Annual Review of Phytopathology, 2014, 52: 347-375.

2026年第66卷第8期

PDF下载

230

109

引用本文

BibTeX

文章信息

doi: 10.13343/j.cnki.wsxb.20260109

接收时间：2026-02-05
首发时间：2026-08-21
出版时间：2026-08-04

补充材料

相关文章

文章信息

作者

出版历史

收稿日期：2026-02-05
录用日期：2026-04-13

基金

National Natural Science Foundation of China(82173923)

国家自然科学基金(82173923)

作者信息

中国医学科学院北京协和医学院，药用植物研究所，道地药材品质保障与资源持续利用全国重点实验室，北京

通讯作者:

参考文献

分享链接

https://castjournals.cast.org.cn/joweb/wswxb/CN/10.13343/j.cnki.wsxb.20260109

分享至

全文二维码

扫描看全文

引用本文

BibTeX

本文的引用情况

2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

关闭全屏

BibTeX
EndNote
RefWorks
TxT