Article(id=1238813309554708739, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1238813307784712441, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250809, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1761667200000, receivedDateStr=2025-10-29, revisedDate=null, revisedDateStr=null, acceptedDate=1766937600000, acceptedDateStr=2025-12-29, onlineDate=1773285709035, onlineDateStr=2026-03-12, pubDate=1772553600000, pubDateStr=2026-03-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1773285709035, onlineIssueDateStr=2026-03-12, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1773285709035, creator=13701087609, updateTime=1773285709035, updator=13701087609, issue=Issue{id=1238813307784712441, tenantId=1146029695717560320, journalId=1192105938417971205, year='2026', volume='66', issue='3', pageStart='961', pageEnd='1466', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1773285708614, creator=13701087609, updateTime=1773291912509, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1238839328915378858, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1238813307784712441, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1238839328915378859, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1238813307784712441, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=1045, endPage=1061, ext={EN=ArticleExt(id=1238813309948973323, articleId=1238813309554708739, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Nucleotide second messengers regulate metabolism, development, and environmental adaptation of actinobacteria: a review, columnId=1192149543727808575, journalTitle=Acta Microbiologica Sinica, columnName=Review, runingTitle=null, highlight=null, articleAbstract=

Actinobacteria are Gram-positive bacteria of major ecological and biotechnological importance, responsible for organic matter turnover in nature and serving as a primary source of antibiotics and other bioactive natural products. Their complex physiological adaptation and life cycles are regulated by sophisticated signal transduction networks. This review examines the metabolism, signaling, and regulatory networks of nucleotide second messengers including cyclic diadenosine monophosphate (c-di-AMP), cyclic di-guanosine monophosphate (c-di-GMP), adenosine monophosphate (cAMP), and (p)ppGpp in the morphologically complex Streptomyces with rich secondary metabolites and pathogenic Mycobacterium with host adaptation. We discuss how these second messengers interact with other signaling systems, such as two-component systems, quorum sensing, and protein acylation, to integrate environmental and developmental cues. This coordination regulates the growth, development, secondary metabolite biosynthesis, and environmental adaptation of actinobacteria. By synthesizing current knowledge, this review provides reference for understanding the integrity and dynamics of the signal transduction system of actinobacteria, as well as their potential applications in the basic research of life sciences and in the fields of biotechnology and medicine.

, correspAuthors=Di YOU, Bangce YE, authorNote=null, correspAuthorsNote=, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, authorCompany=null, fund=null, authors=, authorsList=Xiaohui TANG, Yifan LIANG, Jingqi LI, Di YOU, Bangce YE), CN=ArticleExt(id=1238813312473944391, articleId=1238813309554708739, tenantId=1146029695717560320, journalId=1192105938417971205, language=CN, title=核苷类第二信使与放线菌代谢、发育及环境适应的调控机制研究进展, columnId=1192149543882997826, journalTitle=微生物学报, columnName=综述, runingTitle=null, highlight=null, articleAbstract=

放线菌是一类在生态功能和生物技术领域均具有重要地位的革兰氏阳性细菌，它们不仅驱动自然界的有机质循环，还是抗生素及其他生物活性天然产物的主要来源。其复杂的生理适应与生命活动受到精密信号转导系统的调控。本文聚焦于2类代表性放线菌：形态复杂且次级代谢丰富的链霉菌(代表工业生产菌株)以及具备宿主适应性与致病性的分枝杆菌(代表病原类群)，系统综述了核苷类第二信使[包括环二腺嘌呤核苷酸(cyclic diadenosine monophosphate, c-di-AMP)、环二鸟嘌呤核苷酸(cyclic di-guanosine monophosphate, c-di-GMP)、环腺嘌呤核苷酸(adenosine monophosphate, cAMP)及(p)ppGpp]在细胞内的代谢、信号传递及其调控网络。进一步探讨了这些信号分子如何与双组分系统、群体感应及蛋白质酰化修饰等其他调控途径交互作用，从而整合环境与发育信号，协调细菌的生长、发育、次级代谢合成及环境适应性，为理解放线菌信号转导系统的整体性、动态性及其在基础生命科学研究和生物技术、医学等领域的潜在应用提供参考。

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A-D depict the synthesis and degradation processes of their corresponding signaling molecules, with orange circles representing the catalytic enzymes for the synthesis pathways and blue circles for the degradation pathways., figureFileSmall=WsL8Z+/C33buayE+o7gHvQ==, figureFileBig=PGQyJsM/YSv84/+YkCEvXg==, tableContent=null), ArticleFig(id=1238891101801730617, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1238813309554708739, language=CN, label=图2, caption=c-di-AMP (A)、c-di-GMP (B)、cAMP (C)和(p)ppGpp (D)合成与降解途径示意图。A-D分别展示对应信号分子的合成和降解过程，其中，橘色圆形代表合成途径的催化酶，蓝色圆形代表降解途径的催化酶。, figureFileSmall=WsL8Z+/C33buayE+o7gHvQ==, figureFileBig=PGQyJsM/YSv84/+YkCEvXg==, tableContent=null), ArticleFig(id=1238891101923365441, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1238813309554708739, language=EN, label=Figure 3, caption=c-di-AMP signaling and regulatory network. c-di-AMP binds to receptor proteins such as DarR and DasR to regulate the expression of genes involved in fatty acid transport, GlcNAc metabolism, secondary metabolism, and stress responses. In addition, nutrient stress induces acetyl phosphate (AcP)-dependent protein acetylation, which inhibits both the c-di-AMP synthase DisA and the receptor DasR, thereby perturbing intracellular c-di-AMP synthesis and homeostasis., figureFileSmall=MhWtSlNCpQABjpS+EgqKUA==, figureFileBig=IjEwe+bNGUY6xmwBkmiOvQ==, tableContent=null), ArticleFig(id=1238891103450092108, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1238813309554708739, language=CN, label=图3, caption=c-di-AMP信号转导途径与调控机制示意图。c-di-AMP通过结合转录因子DarR和DasR等受体蛋白，调控脂肪酸转运、GlcNAc代谢、次级代谢及应激响应相关基因的表达。在营养胁迫条件下，乙酰磷酸(AcP)依赖的蛋白质乙酰化作用可抑制c-di-AMP合成酶DisA及其受体DasR的活性，从而影响胞内c-di-AMP的合成与稳态。, figureFileSmall=MhWtSlNCpQABjpS+EgqKUA==, figureFileBig=IjEwe+bNGUY6xmwBkmiOvQ==, tableContent=null), ArticleFig(id=1238891103592698456, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1238813309554708739, language=EN, label=Figure 4, caption=c-di-GMP signaling and regulatory network. c-di-GMP regulates genes governing morphological differentiation and secondary metabolism by binding to the transcriptional regulator BldD and forming a complex with WhiG and RsiG. Under nutrient stress, acetyl phosphate (AcP)-dependent protein acetylation disrupts c-di-GMP-BldD signaling, thereby coordinating developmental and metabolic adaptation. In mycobacteria, c-di-GMP modulates biofilm formation through its interaction with the nucleoid-associated protein Lsr2., figureFileSmall=nIvvLgJP/OJiYdt2K0BY4Q==, figureFileBig=7F6C4ClljQZ29DxyXqkQ0g==, tableContent=null), ArticleFig(id=1238891103710138977, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1238813309554708739, language=CN, label=图4, caption=c-di-GMP信号转导途径与调控机制示意图。c-di-GMP通过结合转录调节因子BldD，并与WhiG、RsiG形成复合物，调控形态分化与次级代谢相关基因的表达。营养胁迫诱导的乙酰磷酸(AcP)依赖型蛋白乙酰化可阻断c-di-GMP与BldD的信号传导，从而在环境胁迫下协调细菌的形态分化与次级代谢过程。此外，在分枝杆菌中c-di-GMP与Lsr2结合，通过调控相关基因的转录影响生物膜的形成。, figureFileSmall=nIvvLgJP/OJiYdt2K0BY4Q==, figureFileBig=7F6C4ClljQZ29DxyXqkQ0g==, tableContent=null), ArticleFig(id=1238891103844356716, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1238813309554708739, language=EN, label=Figure 5, caption=cAMP signaling and regulatory network. Under nutrient deprivation and other environmental stresses, accumulation of cAMP binds to receptor proteins such as the cAMP receptor protein (CRP) and acetyltransferases. Through transcriptional control and posttranslational modification, cAMP thereby coordinates the expression and activity of genes involved in stress adaptation, host-immune interactions, and bacterial virulence, ultimately shaping mycobacterial fitness, and pathogenicity., figureFileSmall=++JWnpLGo6RLDS/aLSUfDw==, figureFileBig=7H95s3yuFkRUBQ77coyCEw==, tableContent=null), ArticleFig(id=1238891103957602932, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1238813309554708739, language=CN, label=图5, caption=cAMP信号转导途径与调控机制示意图。在营养匮乏等环境胁迫条件下，胞内cAMP水平升高，进而与CRP及乙酰转移酶等受体蛋白结合，通过转录调控和蛋白质翻译后修饰双重机制调控环境胁迫响应、宿主免疫互作及细菌毒力相关基因的表达与功能，影响分枝杆菌的环境适应性和致病性。, figureFileSmall=++JWnpLGo6RLDS/aLSUfDw==, figureFileBig=7H95s3yuFkRUBQ77coyCEw==, tableContent=null), ArticleFig(id=1238891104075043455, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1238813309554708739, language=EN, label=Figure 6, caption=(p)ppGpp signaling and regulatory network. (p)ppGpp cooperates with DksA to bind RNA polymerase (RNAP), repressing ribosomal and nitrogen-assimilation genes while activating those for secondary metabolism and morphological differentiation. This transcriptional shift reallocates cellular resources to coordinate growth, stress responses, development, and secondary metabolism in actinobacteria., figureFileSmall=y6H7iLbySGI4XyDi0B/ATA==, figureFileBig=R1B/8vUCa2Pbzl9qsancWw==, tableContent=null), ArticleFig(id=1238891104167318157, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1238813309554708739, language=CN, label=图6, caption=(p)ppGpp信号转导途径与调控机制示意图。(p)ppGpp与RNA聚合酶辅助因子DksA协同作用，通过结合RNA聚合酶(RNAP)形成调控复合物，特异性抑制核糖体生物合成及氮同化相关基因的转录，同时激活次级代谢与形态分化相关基因的表达。该转录重编程引导胞内资源重新分配，从而协调放线菌的生长、胁迫应答、形态分化及次级代谢过程。, figureFileSmall=y6H7iLbySGI4XyDi0B/ATA==, figureFileBig=R1B/8vUCa2Pbzl9qsancWw==, tableContent=null), ArticleFig(id=1238891104276370073, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1238813309554708739, language=EN, label=Table 1, caption=

Nucleotide second messengers, receptors, and functions

, figureFileSmall=null, figureFileBig=null, tableContent=

No.	Nucleotide second messenger	Receptor	Organism	Function	References
1	c-di-AMP	DarR, DasR	M. smegmatis, S. erythraea	DNA binding and transcriptional regulation	[30-31]
2	c-di-GMP	BldD, WhiG, RsiG, Lsr2	Streptomyces venezuelae, S. coelicolor, M. tuberculosis	Regulation of morphological differentiation, secondary metabolism, and biofilm formation	[37-38,44]
3	cAMP	CRP	M. tuberculosis	Regulation of bacterial environmental adaptability and pathogenicity	[45-47]
4	(p)ppGpp	DksA	Streptomyces diastatochromogenes 1628	Global regulation of resource allocation and bacterial growth	[66]

), ArticleFig(id=1238891104381227686, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1238813309554708739, language=CN, label=表1, caption=

核苷类第二信使、受体与功能

, figureFileSmall=null, figureFileBig=null, tableContent=

No.	Nucleotide second messenger	Receptor	Organism	Function	References
1	c-di-AMP	DarR, DasR	M. smegmatis, S. erythraea	DNA binding and transcriptional regulation	[30-31]
2	c-di-GMP	BldD, WhiG, RsiG, Lsr2	Streptomyces venezuelae, S. coelicolor, M. tuberculosis	Regulation of morphological differentiation, secondary metabolism, and biofilm formation	[37-38,44]
3	cAMP	CRP	M. tuberculosis	Regulation of bacterial environmental adaptability and pathogenicity	[45-47]
4	(p)ppGpp	DksA	Streptomyces diastatochromogenes 1628	Global regulation of resource allocation and bacterial growth	[66]

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