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Article(id=1226296954582381485, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226296952975966478, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20240584, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1727020800000, receivedDateStr=2024-09-23, revisedDate=null, revisedDateStr=null, acceptedDate=1731254400000, acceptedDateStr=2024-11-11, onlineDate=1770301577468, onlineDateStr=2026-02-05, pubDate=1738598400000, pubDateStr=2025-02-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1770301577468, onlineIssueDateStr=2026-02-05, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1770301577468, creator=13701087609, updateTime=1770301577468, updator=13701087609, issue=Issue{id=1226296952975966478, tenantId=1146029695717560320, journalId=1192105938417971205, year='2025', volume='65', issue='2', pageStart='421', pageEnd='861', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1770301577085, creator=13701087609, updateTime=1770353593135, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1226515124169650204, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226296952975966478, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1226515124173844509, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226296952975966478, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=597, endPage=613, ext={EN=ArticleExt(id=1226296956138468279, articleId=1226296954582381485, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Isolation and identification of actinomycetes from rhizosphere soil samples of mangrove forests in Quanzhou Bay and secondary metabolites of Streptomyces sp. W444, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

[Objective] To isolate and identify actinomycetes from two mangrove soil samples in Quanzhou Bay as well as secondary metabolites from the target strain Streptomyces sp. W444 with antifungal activity. [Methods] The rhizosphere soil samples of two different mangrove plants were collected from Luoyang River in Quanzhou Bay. Actinomycetes were isolated from the soil samples by the dilution plating method. The isolates were classified by the phylogenetic tree constructed based on 16S rRNA gene sequences. The antifungal activities of the isolates were examined by the agar diffusion method. The target strain Streptomyces sp. W444 was subjected to fermentation scale-up for the isolation of secondary metabolites. Furthermore, the biosynthetic gene clusters were analyzed to deduce the biosynthetic pathway of staurosporine. [Results] A total of 56 strains of actinomycetes were isolated from mangrove soil samples and categorized into 8 genera belonging to 6 families of 6 orders. Among them, Streptomyces and Micromonospora were dominant, with the relative abundance of 41.0% and 33.9%, respectively. Strain Streptomyces sp. W444 exhibited excellent antifungal activity, and three indoles (staurosporine, K252c, and streptochlorin) were isolated and identified from this strain. The biosynthetic gene cluster of staurosporine was localized in the genome of Streptomyces sp. W444 by bioinformatics analysis. The biosynthetic pathway of staurosporine was then proposed. [Conclusion] Actinomycetes in the rhizosphere soil of mangrove plants in Quanzhou Bay had high diversity and contained potential natural product resources. Staurosporine, K252c, and streptochlorin were isolated from strain W444. These findings lay a solid foundation for studying the diversity and secondary metabolites of cultivable actinomycetes in mangrove forests in Quanzhou Bay.

, correspAuthors=Fei PENG, authorNote=null, correspAuthorsNote=
*E-mail:
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【目的】 从泉州湾2种红树林的根际土壤中分离并鉴定放线菌，进行抗真菌活性初筛以获取目标菌株，并对链霉菌属(Streptomyces) W444的次级代谢产物进行分离与鉴定。 【方法】 从泉州湾洛阳江采集2种不同红树林植物的根际土壤样品，采用稀释涂布法对土样中的放线菌进行分离，并构建16S rRNA基因系统发育树。采用琼脂打孔扩散法进行抗真菌活性筛选，对获取的目标菌株Streptomyces sp. W444进行放大规模发酵，并对其次级代谢产物进行了分离与鉴定。根据生物合成基因簇定位和分析，推导星孢菌素的生物合成途径。 【结果】 从根际土样中分离得到56株放线菌，分布于6目6科8属，其中链霉菌属和小单孢菌属为优势菌，占比分别为41.0%和33.9%。通过抗真菌筛选获得抑菌活性较好的菌株Streptomyces sp. W444，并从中分离鉴定了3个吲哚类化合物：staurosporine、K252c和streptochlorin。此外，还从菌株Streptomyces sp. W444基因组中定位了星孢菌素生物合成基因簇，并对其生物合成途径进行了推导。 【结论】 泉州湾红树林根际土壤中的放线菌具有多样性，并蕴含着丰富的潜在天然产物资源，从菌株Streptomyces sp. W444中分离鉴定了吲哚咔唑类生物碱staurosporine、K252c和streptochlorin，这为研究泉州湾红树林可培养放线菌的多样性和次级代谢产物研究提供了坚实的基础。

, correspAuthors=彭飞, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=Lrx8+xRP7SgfNXmU0guEuw==, magXml=P0ud4IzCjgtfyGBaRj/Ydw==, pdfUrl=null, pdf=YTbyadJG5b51+anBzcj/yg==, pdfFileSize=3764371, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=VdmVB+Hcsx5LXzMX4z6s8Q==, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=mOq5buOxY3aGy06PnG9SWQ==, mapNumber=null, authorCompany=null, fund=null, authors=

作者贡献声明

张文州：实验设计与执行，撰写初稿；陈琳琳：数据收集与整理，抗菌实验操作；林水森：数据收集与整理，协助撰写初稿；林阳君：数据收集与整理，数据分析与解释；庄月娥：数据收集与整理，协助撰写初稿；谢志新：技术支持；江红：稿件修订；连云阳：稿件修订；骆祝华：稿件修订；彭飞：概念与设计，数据分析与解释，稿件修订。

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European Journal of Medicinal Chemistry, 2012, 53: 283-291., articleTitle=Synthesis and fungicidal activity of novel pimprinine analogues, refAbstract=null), Reference(id=1226514053619696199, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, doi=null, pmid=null, pmcid=null, year=2018, volume=13, issue=2, pageStart=137, pageEnd=144, url=null, language=null, rfNumber=[37], rfOrder=43, authorNames=王冕, 郝晓萌, 李娇, 胡媛媛, 关艳, 王以光, 甘茂罗, 肖春玲, journalName=中国医药生物技术, refType=null, unstructuredReference=王冕, 郝晓萌, 李娇, 胡媛媛, 关艳, 王以光, 甘茂罗, 肖春玲. 海洋链霉菌IMB3-202产生的吲哚咔唑生物碱[J]. 中国医药生物技术, 2018, 13(2): 137-144., articleTitle=海洋链霉菌IMB3-202产生的吲哚咔唑生物碱, refAbstract=null), Reference(id=1226514053724553803, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, doi=null, pmid=null, pmcid=null, year=2018, volume=13, issue=2, pageStart=137, pageEnd=144, url=null, language=null, rfNumber=[37], rfOrder=44, authorNames=WANG M, HAO XM, LI J, HU YY, GUAN Y, WANG YG, GAN ML, XIAO CL, journalName=Chinese Medicinal Biotechnology, refType=null, unstructuredReference=WANG M, HAO XM, LI J, HU YY, GUAN Y, WANG YG, GAN ML, XIAO CL. Identification of indolocarbazoles from the marine-derived Streptomyces sp. IMB3-202[J]. 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journalId=1192105938417971205, articleId=1226296954582381485, language=CN, label=图1, caption=红树林来源部分代表性分离菌株通过16S rRNA基因序列构建的系统发育树, figureFileSmall=JBInf/YBU7Pva67ASAj4tw==, figureFileBig=iHiiSJB8sb44UjGQzuqz9w==, tableContent=null), ArticleFig(id=1226514041489768638, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, language=EN, label=Figure 2, caption=The proportion and classification of cultivable actinobacteria isolated from rhizosphere soil samples of two mangrove plants. 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Soil sample from Mangrove Nature Reserve of Luoyang River in Quanzhou Bay

, figureFileSmall=null, figureFileBig=null, tableContent=

样品编号

Sample number

植物根际

Plant rhizosphere

采样地经纬度

Latitude and longitude of the sampling site

S1互花米草Spartina alterniflora24°55′14″N, 118°39′49″E
S2秋茄Kandelia candel24°54′0″N, 118°40′8″E
), ArticleFig(id=1226514044127985933, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, language=CN, label=表1, caption=

泉州湾洛阳江红树林自然保护区土壤样品

, figureFileSmall=null, figureFileBig=null, tableContent=

样品编号

Sample number

植物根际

Plant rhizosphere

采样地经纬度

Latitude and longitude of the sampling site

S1互花米草Spartina alterniflora24°55′14″N, 118°39′49″E
S2秋茄Kandelia candel24°54′0″N, 118°40′8″E
), ArticleFig(id=1226514044262203665, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, language=EN, label=Table 2, caption=

Summary of actinomycete strains isolated from rhizosphere soil samples of two different

, figureFileSmall=null, figureFileBig=null, tableContent=
NamesMangrove plantsTop-hit taxonAccession numberSimilarity (%)
W1-3Kandelia candelMicromonospora provocatoris (AY894337)PP20417498.93
W4-1Kandelia candelMicromonospora aurantiaca (CP002162)PP20417599.93
W3-1-1Kandelia candelMicromonospora taraxaci (VIWZ01000001)PP20417799.30
W3-7Kandelia candelMicromonospora solifontis (LC383890)PP20417899.09
W112Kandelia candelMicromonospora humida (MT907442)PP20418199.26
W126Kandelia candelMicromonospora fluminis (LR130241)PP20418299.78
W321Kandelia candelMicromonospora globbae (LC177396)PP20418399.78
W152Kandelia candelMicromonospora fluminis (LR130241)PP204184100.00
W153Kandelia candelMicromonospora endophytica (EU560726)PP20418599.77
W156Kandelia candelMicromonospora maritima (HQ704071)PP204186100.00
W343Kandelia candelMicromonospora aurantiaca (CP002162)PP20418799.85
W2-1Kandelia candelMicromonospora aurantiaca (CP002162)PP20419199.41
W3-4Kandelia candelMicromonospora taraxaci (VIWZ01000001)PP20419298.69
W3-6Kandelia candelMicromonospora terminaliae (KX394339)PP20419398.37
W3-8Kandelia candelMicromonospora aurantiaca (CP002162)PP20419499.49
W136Kandelia candelMicromonospora fluminis (LR130241)PP204196100.00
W145Kandelia candelMicromonospora fluminis (LR130241)PP20419799.27
W131Kandelia candelStreptomyces sundarbansensis (AY550275)PP21192199.42
W221Kandelia candelStreptomyces niveus (DQ442532)PP21192299.50
W141Kandelia candelStreptomyces sundarbansensis (AY550275)PP21192399.85
W147Kandelia candelStreptomyces spirodelae (MW602308)PP21192599.48
W162Kandelia candelStreptomyces olivaceus (JOFH01000101)PP21192699.78
W171Kandelia candelStreptomyces badius (AY999783)PP21192799.78
W262Kandelia candelStreptomyces mayteni (EU200683)PP21192899.48
W272Kandelia candelStreptomyces badius (AY999783)PP21192999.78
W273Kandelia candelStreptomyces mayteni (EU200683)PP21193099.48
W372Kandelia candelStreptomyces badius (AY999783)PP21193299.78
W444Kandelia candelStreptomyces fradiae (MIFZ01000280)PP21193399.93
W462Kandelia candelStreptomyces olivaceus (JOFH01000101)PP21193499.93
W471Kandelia candelStreptomyces sundarbansensis (AY550275)PP21193599.85
W442Kandelia candelStreptomyces endocoffeicus (MN116545)PP21194099.41
W132Kandelia candelAmycolatopsis lurida (AJ577997)PP21197099.41
W233Kandelia candelAmycolatopsis lurida (AJ577997)PP21197199.34
W334Kandelia candelAmycolatopsis lurida (AJ577997)PP21197398.69
W242Kandelia candelSaccharopolyspora shandongensis (EF104116)PP211985100.00
W251Kandelia candelGordonia terrae (BAFD01000032)PP211986100.00
W6-2Spartina alternifloraMicromonospora fluminis (LR130241)PP20417999.92
W6-4Spartina alternifloraMicromonospora aurantiaca (CP002162)PP204180100.00
W5-4Spartina alternifloraMicromonospora aurantiaca (CP002162)PP204195100.00
W5-1Spartina alternifloraMicromonospora provocatoris (AY894337)PP20417699.35
W743Spartina alternifloraVerrucosispora rhizosphaerae (HQ123438)PP20418899.55
W751Spartina alternifloraMicromonospora globispora (KF818390)PP20418999.62
W763Spartina alternifloraMicromonospora krabiensis (LT598496)PP20419099.85
W671Spartina alternifloraStreptomyces badius (AY999783)PP21193699.78
W742Spartina alternifloraStreptomyces spirodelae (MW602308)PP21193799.48
W744Spartina alternifloraStreptomyces spirodelae (MW602308)PP21193899.26
W761Spartina alternifloraStreptomyces olivaceus (JOFH01000101)PP21193999.93
W572Spartina alternifloraStreptomyces mayteni (EU200683)PP21194199.48
W531Spartina alternifloraAmycolatopsis lurida (AJ577997)PP21197599.40
W723Spartina alternifloraGordonia didemnid (JN615417)PP211976100.00
W731Spartina alternifloraNocardia rhizosphaerae (KP972639)PP21197799.78
W733Spartina alternifloraAmycolatopsis keratiniphila (LQMT01000206)PP21197899.48
W735Spartina alternifloraNocardioides luteus (AF005007)PP21197999.48
W822Spartina alternifloraGordonia terrae (BAFD01000032)PP21198099.93
W831Spartina alternifloraAmycolatopsis keratiniphila (LQMT01000206)PP21198199.48
W745Spartina alternifloraSaccharopolyspora spongiae (KX037095)PP21198898.66
), ArticleFig(id=1226514044388032790, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, language=CN, label=表2, caption=

两种不同红树林植物根际土样品分离的放线菌概况 (mangrove plants)

, figureFileSmall=null, figureFileBig=null, tableContent=
NamesMangrove plantsTop-hit taxonAccession numberSimilarity (%)
W1-3Kandelia candelMicromonospora provocatoris (AY894337)PP20417498.93
W4-1Kandelia candelMicromonospora aurantiaca (CP002162)PP20417599.93
W3-1-1Kandelia candelMicromonospora taraxaci (VIWZ01000001)PP20417799.30
W3-7Kandelia candelMicromonospora solifontis (LC383890)PP20417899.09
W112Kandelia candelMicromonospora humida (MT907442)PP20418199.26
W126Kandelia candelMicromonospora fluminis (LR130241)PP20418299.78
W321Kandelia candelMicromonospora globbae (LC177396)PP20418399.78
W152Kandelia candelMicromonospora fluminis (LR130241)PP204184100.00
W153Kandelia candelMicromonospora endophytica (EU560726)PP20418599.77
W156Kandelia candelMicromonospora maritima (HQ704071)PP204186100.00
W343Kandelia candelMicromonospora aurantiaca (CP002162)PP20418799.85
W2-1Kandelia candelMicromonospora aurantiaca (CP002162)PP20419199.41
W3-4Kandelia candelMicromonospora taraxaci (VIWZ01000001)PP20419298.69
W3-6Kandelia candelMicromonospora terminaliae (KX394339)PP20419398.37
W3-8Kandelia candelMicromonospora aurantiaca (CP002162)PP20419499.49
W136Kandelia candelMicromonospora fluminis (LR130241)PP204196100.00
W145Kandelia candelMicromonospora fluminis (LR130241)PP20419799.27
W131Kandelia candelStreptomyces sundarbansensis (AY550275)PP21192199.42
W221Kandelia candelStreptomyces niveus (DQ442532)PP21192299.50
W141Kandelia candelStreptomyces sundarbansensis (AY550275)PP21192399.85
W147Kandelia candelStreptomyces spirodelae (MW602308)PP21192599.48
W162Kandelia candelStreptomyces olivaceus (JOFH01000101)PP21192699.78
W171Kandelia candelStreptomyces badius (AY999783)PP21192799.78
W262Kandelia candelStreptomyces mayteni (EU200683)PP21192899.48
W272Kandelia candelStreptomyces badius (AY999783)PP21192999.78
W273Kandelia candelStreptomyces mayteni (EU200683)PP21193099.48
W372Kandelia candelStreptomyces badius (AY999783)PP21193299.78
W444Kandelia candelStreptomyces fradiae (MIFZ01000280)PP21193399.93
W462Kandelia candelStreptomyces olivaceus (JOFH01000101)PP21193499.93
W471Kandelia candelStreptomyces sundarbansensis (AY550275)PP21193599.85
W442Kandelia candelStreptomyces endocoffeicus (MN116545)PP21194099.41
W132Kandelia candelAmycolatopsis lurida (AJ577997)PP21197099.41
W233Kandelia candelAmycolatopsis lurida (AJ577997)PP21197199.34
W334Kandelia candelAmycolatopsis lurida (AJ577997)PP21197398.69
W242Kandelia candelSaccharopolyspora shandongensis (EF104116)PP211985100.00
W251Kandelia candelGordonia terrae (BAFD01000032)PP211986100.00
W6-2Spartina alternifloraMicromonospora fluminis (LR130241)PP20417999.92
W6-4Spartina alternifloraMicromonospora aurantiaca (CP002162)PP204180100.00
W5-4Spartina alternifloraMicromonospora aurantiaca (CP002162)PP204195100.00
W5-1Spartina alternifloraMicromonospora provocatoris (AY894337)PP20417699.35
W743Spartina alternifloraVerrucosispora rhizosphaerae (HQ123438)PP20418899.55
W751Spartina alternifloraMicromonospora globispora (KF818390)PP20418999.62
W763Spartina alternifloraMicromonospora krabiensis (LT598496)PP20419099.85
W671Spartina alternifloraStreptomyces badius (AY999783)PP21193699.78
W742Spartina alternifloraStreptomyces spirodelae (MW602308)PP21193799.48
W744Spartina alternifloraStreptomyces spirodelae (MW602308)PP21193899.26
W761Spartina alternifloraStreptomyces olivaceus (JOFH01000101)PP21193999.93
W572Spartina alternifloraStreptomyces mayteni (EU200683)PP21194199.48
W531Spartina alternifloraAmycolatopsis lurida (AJ577997)PP21197599.40
W723Spartina alternifloraGordonia didemnid (JN615417)PP211976100.00
W731Spartina alternifloraNocardia rhizosphaerae (KP972639)PP21197799.78
W733Spartina alternifloraAmycolatopsis keratiniphila (LQMT01000206)PP21197899.48
W735Spartina alternifloraNocardioides luteus (AF005007)PP21197999.48
W822Spartina alternifloraGordonia terrae (BAFD01000032)PP21198099.93
W831Spartina alternifloraAmycolatopsis keratiniphila (LQMT01000206)PP21198199.48
W745Spartina alternifloraSaccharopolyspora spongiae (KX037095)PP21198898.66
), ArticleFig(id=1226514044522250524, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, language=EN, label=Table 3, caption=

Primary results of the antifungal activities of crude extract of Streptomyces sp. W444

, figureFileSmall=null, figureFileBig=null, tableContent=

样品

Sample

白色念珠菌ATCC 90028

Candida albicans ATCC 90028

黑曲霉FJAT-31087

Aspergillus niger FJAT-31087

茄病镰刀菌QY22

Fusarium solanaceae QY22

W444201924
多菌灵Carbendazim353732
), ArticleFig(id=1226514044648079648, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, language=CN, label=表3, caption=

Streptomyces sp. W444发酵粗提物的抗真菌活性初筛结果

, figureFileSmall=null, figureFileBig=null, tableContent=

样品

Sample

白色念珠菌ATCC 90028

Candida albicans ATCC 90028

黑曲霉FJAT-31087

Aspergillus niger FJAT-31087

茄病镰刀菌QY22

Fusarium solanaceae QY22

W444201924
多菌灵Carbendazim353732
), ArticleFig(id=1226514044778103076, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, language=EN, label=Table 4, caption=

1H and 13C NMR data of compound F4-1 and F5-1

, figureFileSmall=null, figureFileBig=null, tableContent=
PositionCompound 4-1δ H values for compound 5-1
δ C, typeδ Hδ C, type
1112.5, CH7.63, 1H, d108.9, CH7.56, 1H, d
2126.3, C7.44, 1H, t125.7, CH7.45, 1H, td
3119.4, CH7.24, 1H, t120.4, C7.28, 1H, t
4127.0, CH9.20, 1H, d127.0, CH9.27, 1H, d
4a124.5, C-124.4, C-
4b117.7, C-115.4, C-
4c120.3, C-119.5, C-
5176.0, C-175.6, C-
6----
747.1, CH25.03, 2H, s45.4, CH24.98, 2H, s
7a134.7, C-133.9, C-
7b115.9, C-115.8, C-
7c124.3, C-125.6, C-
8122.0, CH8.04, 1H, d121.9, CH7.96, 1H, d
9121.3, CH7.33, 1H, t121.3, CH7.28, 1H, t,
10126.4, CH7.48, 1H, t126.3, CH7.41, 1H, td,
11111.8, CH7.69, 1H, t116.3, CH8.00, 1H, d
11a141.1, C-140.9, C-
12a130.8, C-131.9, C-
12b126.5, C-128.2, C-
13a141.2, C-138.1, C-
2′93.3, C-
3′84.5, CH4.04, 1H, d
4′52.7, CH3.25, 1H, m
5′31.1, CH2252, 2.62, 2H, m
6′82.1, CH6.62, 1H, dd
7′30.1, CH32.39, 3H, s
8′58.5, CH33.14, 3H, s
9′334, CH31.79, 3H, s
), ArticleFig(id=1226514044908126509, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, language=CN, label=表4, caption=

化合物F4-1和F5-1的氢谱和碳谱数据

, figureFileSmall=null, figureFileBig=null, tableContent=
PositionCompound 4-1δ H values for compound 5-1
δ C, typeδ Hδ C, type
1112.5, CH7.63, 1H, d108.9, CH7.56, 1H, d
2126.3, C7.44, 1H, t125.7, CH7.45, 1H, td
3119.4, CH7.24, 1H, t120.4, C7.28, 1H, t
4127.0, CH9.20, 1H, d127.0, CH9.27, 1H, d
4a124.5, C-124.4, C-
4b117.7, C-115.4, C-
4c120.3, C-119.5, C-
5176.0, C-175.6, C-
6----
747.1, CH25.03, 2H, s45.4, CH24.98, 2H, s
7a134.7, C-133.9, C-
7b115.9, C-115.8, C-
7c124.3, C-125.6, C-
8122.0, CH8.04, 1H, d121.9, CH7.96, 1H, d
9121.3, CH7.33, 1H, t121.3, CH7.28, 1H, t,
10126.4, CH7.48, 1H, t126.3, CH7.41, 1H, td,
11111.8, CH7.69, 1H, t116.3, CH8.00, 1H, d
11a141.1, C-140.9, C-
12a130.8, C-131.9, C-
12b126.5, C-128.2, C-
13a141.2, C-138.1, C-
2′93.3, C-
3′84.5, CH4.04, 1H, d
4′52.7, CH3.25, 1H, m
5′31.1, CH2252, 2.62, 2H, m
6′82.1, CH6.62, 1H, dd
7′30.1, CH32.39, 3H, s
8′58.5, CH33.14, 3H, s
9′334, CH31.79, 3H, s
), ArticleFig(id=1226514045050732855, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, language=EN, label=Table 5, caption=

Deduced functions of ORFs in cluster 3 in Streptomyces sp. W444 from rhizosphere soil of

, figureFileSmall=null, figureFileBig=null, tableContent=
OrfProduct size (aa)Protein homologIdentity (%)/Similarity (%)Proposed functionAccession number
1957StaR68/72Transcriptional activatorBAC55205
2408StaN85/92Cytochrome P450BAC55208
3436StaG79/87N-glycosyltransferaseBAC55209
4504StaO82/87l-amino acid oxidaseBAC55210
51 154StaD73/79Chromopyrrolic acid synthaseBAC15759
6418StaP73/80Cytochrome P450BAC55212
7274StaMA73/83MethyltransferaseBAC55213
8288StaMB82/89MethyltransferaseBAC55218
9545StaC80/88MonooxygenaseBAF47693
1028089/92Recombinase family proteinMET7710960
1129596/97Aminoglycoside phosphotransferase family proteinWP_093549433
1242797/98Helix-turn-helix domain-containing proteinSCD65229
13157100/100ATP-binding proteinWP_283298866
1415095/96DUF6415 family natural product biosynthesis proteinWP_283298867
1515796/96DUF6302 family proteinWP_229870986
166697/96DUF5999 family proteinWP_229901893
1722197/99Hypothetical proteinWP_337674500
1841198/99Methyltransferase, FxLD systemWP_283298873
196597/98FxLD family lanthipeptideWP_093549423
201 00697/97Lantibiotic dehydrataseWP_337674501
2140797/98Lanthionine synthetase C family proteinWP_189716992
2232997/98Thiopeptide-type bacteriocin biosynthesis proteinWP_308435751
2324182/85Hypothetical proteinMER5525945
2427897/97SGNH/GDSL hydrolase family proteinWP_031128207
2533899/99Hemolysin family proteinWP_359285438
2651696/97Hemolysin family proteinWP_359388163
2714294/95GNAT family N-acetyltransferaseAYA81838
288396/96AntitoxinWP_315879421
2912399/99Fic family toxin-antitoxin system, toxin componentWP_317433620
3012687/88Hypothetical proteinOSY52980
3122697/98Class I SAM-dependent methyltransferaseMEV2200025
3211498/98VOC family proteinMEV2200026
3324096/96Dethiobiotin synthaseWP_317433624
), ArticleFig(id=1226514045172367673, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, language=CN, label=表5, caption=

红树林根际土壤来源的 Streptomyces sp. W444中cluster 3各开放阅读框的功能预测 (mangrove plant)

, figureFileSmall=null, figureFileBig=null, tableContent=
OrfProduct size (aa)Protein homologIdentity (%)/Similarity (%)Proposed functionAccession number
1957StaR68/72Transcriptional activatorBAC55205
2408StaN85/92Cytochrome P450BAC55208
3436StaG79/87N-glycosyltransferaseBAC55209
4504StaO82/87l-amino acid oxidaseBAC55210
51 154StaD73/79Chromopyrrolic acid synthaseBAC15759
6418StaP73/80Cytochrome P450BAC55212
7274StaMA73/83MethyltransferaseBAC55213
8288StaMB82/89MethyltransferaseBAC55218
9545StaC80/88MonooxygenaseBAF47693
1028089/92Recombinase family proteinMET7710960
1129596/97Aminoglycoside phosphotransferase family proteinWP_093549433
1242797/98Helix-turn-helix domain-containing proteinSCD65229
13157100/100ATP-binding proteinWP_283298866
1415095/96DUF6415 family natural product biosynthesis proteinWP_283298867
1515796/96DUF6302 family proteinWP_229870986
166697/96DUF5999 family proteinWP_229901893
1722197/99Hypothetical proteinWP_337674500
1841198/99Methyltransferase, FxLD systemWP_283298873
196597/98FxLD family lanthipeptideWP_093549423
201 00697/97Lantibiotic dehydrataseWP_337674501
2140797/98Lanthionine synthetase C family proteinWP_189716992
2232997/98Thiopeptide-type bacteriocin biosynthesis proteinWP_308435751
2324182/85Hypothetical proteinMER5525945
2427897/97SGNH/GDSL hydrolase family proteinWP_031128207
2533899/99Hemolysin family proteinWP_359285438
2651696/97Hemolysin family proteinWP_359388163
2714294/95GNAT family N-acetyltransferaseAYA81838
288396/96AntitoxinWP_315879421
2912399/99Fic family toxin-antitoxin system, toxin componentWP_317433620
3012687/88Hypothetical proteinOSY52980
3122697/98Class I SAM-dependent methyltransferaseMEV2200025
3211498/98VOC family proteinMEV2200026
3324096/96Dethiobiotin synthaseWP_317433624
), ArticleFig(id=1226514045302391104, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, language=EN, label=Table 6, caption=

Minimum inhibitory concentration (MIC) test of compounds (μg/mL)

, figureFileSmall=null, figureFileBig=null, tableContent=

样品

Sample

白色念珠菌ATCC 90028

Candida albicans

黑曲霉 FJAT-31087

Aspergillus niger

茄病镰刀菌QY22

Fusariums olanaceae QY22

Compound F2-1>200>200200
Compound F4-1>200100100
Compound F5-1502550
Carbendazim2512.550
), ArticleFig(id=1226514045390471493, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226296954582381485, language=CN, label=表6, caption=

化合物的最小抑菌浓度(MIC)测定

, figureFileSmall=null, figureFileBig=null, tableContent=

样品

Sample

白色念珠菌ATCC 90028

Candida albicans

黑曲霉 FJAT-31087

Aspergillus niger

茄病镰刀菌QY22

Fusariums olanaceae QY22

Compound F2-1>200>200200
Compound F4-1>200100100
Compound F5-1502550
Carbendazim2512.550
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泉州湾红树林根际土壤放线菌的分离鉴定及链霉菌W444次级代谢产物的鉴定
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张文州 1 , 陈琳琳 1 , 林水森 1 , 林阳君 1 , 庄月娥 1 , 谢志新 1 , 江红 2 , 连云阳 2 , 骆祝华 3 , 彭飞 1, *
微生物学报 | 研究报告 2025,65(2): 597-613
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微生物学报 | 研究报告 2025, 65(2): 597-613
泉州湾红树林根际土壤放线菌的分离鉴定及链霉菌W444次级代谢产物的鉴定
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张文州1, 陈琳琳1, 林水森1, 林阳君1, 庄月娥1, 谢志新1, 江红2, 连云阳2, 骆祝华3, 彭飞1, *
作者信息
  • 1 泉州医学高等专科学校，福建 泉州
  • 2 福建省微生物研究所，福建 福州
  • 3 自然资源部第三海洋研究所，福建 厦门
Isolation and identification of actinomycetes from rhizosphere soil samples of mangrove forests in Quanzhou Bay and secondary metabolites of Streptomyces sp. W444
Wenzhou ZHANG1, Linlin CHEN1, Shuisen LIN1, Yangjun LIN1, Yue'e ZHUANG1, Zhixin XIE1, Hong JIANG2, Yunyang LIAN2, Zhuhua LUO3, Fei PENG1, *
Affiliations
  • 1 Quanzhou Medical College, Quanzhou, Fujian, China
  • 2 Fujian Institute of Microbiology, Fuzhou, Fujian, China
  • 3 Third Institute of Oceanography, Ministry of Natural Resources, Xiamen, Fujian, China
出版时间: 2025-02-04 doi: 10.13343/j.cnki.wsxb.20240584
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摘要
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【目的】 从泉州湾2种红树林的根际土壤中分离并鉴定放线菌，进行抗真菌活性初筛以获取目标菌株，并对链霉菌属(Streptomyces) W444的次级代谢产物进行分离与鉴定。 【方法】 从泉州湾洛阳江采集2种不同红树林植物的根际土壤样品，采用稀释涂布法对土样中的放线菌进行分离，并构建16S rRNA基因系统发育树。采用琼脂打孔扩散法进行抗真菌活性筛选，对获取的目标菌株Streptomyces sp. W444进行放大规模发酵，并对其次级代谢产物进行了分离与鉴定。根据生物合成基因簇定位和分析，推导星孢菌素的生物合成途径。 【结果】 从根际土样中分离得到56株放线菌，分布于6目6科8属，其中链霉菌属和小单孢菌属为优势菌，占比分别为41.0%和33.9%。通过抗真菌筛选获得抑菌活性较好的菌株Streptomyces sp. W444，并从中分离鉴定了3个吲哚类化合物：staurosporine、K252c和streptochlorin。此外，还从菌株Streptomyces sp. W444基因组中定位了星孢菌素生物合成基因簇，并对其生物合成途径进行了推导。 【结论】 泉州湾红树林根际土壤中的放线菌具有多样性，并蕴含着丰富的潜在天然产物资源，从菌株Streptomyces sp. W444中分离鉴定了吲哚咔唑类生物碱staurosporine、K252c和streptochlorin，这为研究泉州湾红树林可培养放线菌的多样性和次级代谢产物研究提供了坚实的基础。

泉州湾红树林  /  放线菌  /  吲哚类化合物  /  生物合成基因簇

[Objective] To isolate and identify actinomycetes from two mangrove soil samples in Quanzhou Bay as well as secondary metabolites from the target strain Streptomyces sp. W444 with antifungal activity. [Methods] The rhizosphere soil samples of two different mangrove plants were collected from Luoyang River in Quanzhou Bay. Actinomycetes were isolated from the soil samples by the dilution plating method. The isolates were classified by the phylogenetic tree constructed based on 16S rRNA gene sequences. The antifungal activities of the isolates were examined by the agar diffusion method. The target strain Streptomyces sp. W444 was subjected to fermentation scale-up for the isolation of secondary metabolites. Furthermore, the biosynthetic gene clusters were analyzed to deduce the biosynthetic pathway of staurosporine. [Results] A total of 56 strains of actinomycetes were isolated from mangrove soil samples and categorized into 8 genera belonging to 6 families of 6 orders. Among them, Streptomyces and Micromonospora were dominant, with the relative abundance of 41.0% and 33.9%, respectively. Strain Streptomyces sp. W444 exhibited excellent antifungal activity, and three indoles (staurosporine, K252c, and streptochlorin) were isolated and identified from this strain. The biosynthetic gene cluster of staurosporine was localized in the genome of Streptomyces sp. W444 by bioinformatics analysis. The biosynthetic pathway of staurosporine was then proposed. [Conclusion] Actinomycetes in the rhizosphere soil of mangrove plants in Quanzhou Bay had high diversity and contained potential natural product resources. Staurosporine, K252c, and streptochlorin were isolated from strain W444. These findings lay a solid foundation for studying the diversity and secondary metabolites of cultivable actinomycetes in mangrove forests in Quanzhou Bay.

mangrove in Quanzhou Bay  /  actinomycetes  /  indoles  /  biosynthetic gene cluster
张文州, 陈琳琳, 林水森, 林阳君, 庄月娥, 谢志新, 江红, 连云阳, 骆祝华, 彭飞. 泉州湾红树林根际土壤放线菌的分离鉴定及链霉菌W444次级代谢产物的鉴定. 微生物学报, 2025 , 65 (2) : 597 -613 . DOI: 10.13343/j.cnki.wsxb.20240584
Wenzhou ZHANG, Linlin CHEN, Shuisen LIN, Yangjun LIN, Yue'e ZHUANG, Zhixin XIE, Hong JIANG, Yunyang LIAN, Zhuhua LUO, Fei PENG. Isolation and identification of actinomycetes from rhizosphere soil samples of mangrove forests in Quanzhou Bay and secondary metabolites of Streptomyces sp. W444[J]. Acta Microbiologica Sinica, 2025 , 65 (2) : 597 -613 . DOI: 10.13343/j.cnki.wsxb.20240584
正文
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近年来，致病性真菌感染的致病率和死亡率呈上升趋势，严重威胁人类的健康[1]。目前临床上治疗真菌感染的药物主要包括唑类、棘白菌素类、嘧啶类似物和多烯类等[2]，但是真菌通常采用改变药物靶标、过度表达外排泵和细胞应激反应等手段来实现对药物的耐受，从而形成耐药性[3]。这种持续存在的严重威胁，促使科技工作者努力寻找新的药物以应对真菌感染。
放线菌产生的天然产物及其衍生物是新药先导化合物的重要来源[4-6]。早期，放线菌主要从陆地生境中筛选获得。然而，随着筛选研究的深入，从普通陆地生境的放线菌中获取新活性产物变得越来越难。因此，更多的研究者将目光聚焦于高原、沙漠、海洋、潮间带、红树林、温泉等特殊生境，以期在这些环境中寻找新的放线菌资源[7-9]，并发现新的活性代谢产物。红树林生长于入海口湿地，具有高盐、高渗透压、低氧等特点，其所处的海水与淡水交汇的特殊生境，使得该地区蕴藏着极为丰富且独特的放线菌资源[10]。例如，Wang等[11]从海南西海岸红树林沉积物中分离的广岛链霉菌(Streptomyces hiroshimensis) GXIMD 06359的粗提物中，获得4个新的大环多烯内酯类化合物antifungalmycin B-E。除了antifungalmycin C外，其他化合物均对马尔尼菲篮状真菌(Talaromyces marneffei)具有较强的抑制活性，其最低抑菌浓度(minimum inhibitory concentration, MIC)为2-128 μg/mL。Wang等[12]从广东珠海淇澳岛红树林土样中分离的链霉菌属(Streptomyces) ZQ4BG中，得到9个32元环多烯内酯类化合物flavofungins I-IX，其中flavofungins III-IX为新化合物，而flavofungins I和II均对白色念珠菌(Candida albicans)具有较强的抑制活性。Che等[13]从广东红树林芦苇根际土壤中分离的链霉菌Streptomyces sp. CHQ-64中，获得6个新的大环多烯内酯化合物reedsmycins A-F，其中reedsmycin A对白色念珠菌(C. albicans)具有较强的抑制活性，其MIC范围为25-50 µmol/L。Zeng等[14]从海南东寨港红树林沉积物分离的白色链霉菌(Streptomyces albogriseolus) HA10002中，得到一类新的抗根结线虫活性化合物fungichromin B，该化合物对酵母菌(Saccharomyces cerevisiae)、尖孢镰孢菌(Fusarium oxysporum)和黑曲霉(Aspergillus niger)均具有较强的抑菌活性。综上所述，从红树林放线菌资源中寻找抗真菌活性先导化合物，仍然是天然药物化学研究的重要方向。
洛阳江红树林湿地位于泉州湾河口湿地自然保护核心区，对于该区域微生物的研究主要集中于微生物群落结构变迁、分布和种类鉴定等方面[15-17]，而关于泉州湾红树林放线菌资源的研究则鲜有报道。本研究报道了泉州湾洛阳江红树林根际土壤中放线菌的分离与鉴定，初步分析了红树林植物根际土壤中可培养放线菌的多样性；结合初步的抗真菌活性筛选，选取Streptomyces sp. W444作为目标菌株进行次级代谢产物研究。从中分离鉴定了3个吲哚类化合物：staurosporine、K252c和streptochlorin，定位了这些化合物生物合成相关的基因簇，并推导了其生物合成途径。本研究揭示了泉州湾红树林放线菌资源的多样性，为后续抗真菌次级代谢产物的研究奠定了基础，同时拓宽了星孢菌素类化合物产生菌株的来源，为该类化合物的生物合成研究提供参考。
1 材料与方法
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1.1 土壤样品
植物根际土壤样品于2022年6月采自泉州湾洛阳江红树林自然保护区，样品及采集信息如表1所示。采集土样的深度为根系以下10 cm，样品用灭菌采样袋收集，于超净台中自然晾干，4 ℃待用。
1.2 培养基
分离培养基W1：可溶性淀粉20 g，CaCO3 1 g，KH2PO4·3H2O 0.5 g，KNO3 0.1 g，MgSO4·7H2O 0.5 g，NaCl 0.5 g，L-天冬素1 g。W2：酪蛋白0.4 g，淀粉10 g，CaCO3 0.1 g，KH2PO4·3H2O 0.2 g。W3：燕麦片40 g。W4：淀粉10 g，酵母提取物3 g，蛋白胨1.5 g。W5：甘油6 mL，精氨酸1 g，KH2PO4·3H2O 1 g，MgSO4·7H2O 1 g。W6：腐殖酸1 g，Na2HPO4 0.5 g，KCl 1.7 g，MgSO4·7H2O 0.05 g，FeSO4·7H2O 0.01 g，CaCO3 0.02 g，复合维生素母液1 mL。W7：葡萄糖4 g，酵母粉4 g，麦芽浸粉10 g，复合维生素母液1 mL。其中复合维生素母液的配方参照梁效伟[18]的研究。以上培养基分别加入15 g的琼脂粉和10 g的海盐，加水定容至1 L，调pH 7.2-7.4。杂菌抑制剂选取放线菌酮、制霉菌素、新生霉素和重铬酸钾，用以抑制细菌和真菌的生长，终浓度分别为25、10、25、25 mg/L，抑制剂待培养基冷却至50 ℃左右时加入。
发酵培养基(g/L)：可溶性淀粉40.0，葡萄糖5.0，花生饼粉20.0，蛋白胨5.0， MgSO4·7H2O 0.5，K2HPO4 0.5，CaCO3 1.0，海盐10，调pH 7.5。
检定菌培养基：真菌培养采用PDA培养基。
1.3 主要试剂和仪器
pEASY-T1克隆试剂盒及感受态细胞，北京全式金生物技术股份有限公司；2×Es Taq MasterMix，北京康为世纪生物科技有限公司；Chelex-100，Bio-Rad公司；AxyPrep PCR纯化试剂盒，爱思进生物技术(杭州)有限公司；PCR引物由苏州金唯智生物科技有限公司合成；IPTG、氨苄西林，北京绿泽森生物技术有限公司；其他试剂均为分析纯或色谱纯。
超净工作台，苏州安泰空气技术有限公司；高速台式离心机，上海安亭科学仪器厂；PCR仪，国力天(深圳)科技有限公司；立式压力蒸汽灭菌器，厦门柏嘉生物科技有限公司；恒温振荡培养箱，上海福玛实验设备有限公司；恒温培养箱，上海一恒科学仪器有限公司；Agilent-1260高效液相色谱仪，色谱柱为Agilent Eclipse XDB-C18柱(5 μm，250 mm×4.6 mm)；核磁共振波谱采用Bruker DRX2500核磁共振仪(600 MHz)测定，以TMS为内标，MeOD为溶剂；ESI-MS采用Agilent 6460液质联用仪，以CH3OH为溶剂测定；凝胶为Sephadex LH-20 (40-70 μm，Amersham Pharmacia Biotech AB，Uppsala，Sweden)；反相硅胶为YMC* GEL ODS-A (12 nm，S-50 μm)。
1.4 检定菌
真菌白色念珠菌(ATCC 90028)购自广东环凯微生物科技有限公司，黑曲霉(FJAT-31087)由福建省农业科学院生物资源研究所农业微生物研究中心惠赠，茄病镰刀菌(QY22)购自中国海洋微生物菌种保藏管理中心。
1.5 土壤样品的处理
将2份土壤样品于无菌超净台中自然风干后，分别置于灭过菌的研钵研碎。称取研碎后的土样3 g，悬浮于30 mL 100%无菌海水中，置于250 mL三角瓶(内含10个小玻璃珠)，充分振荡，混合均匀后加入蛋白胨和十二烷基硫酸钠母液至终浓度分别为6%和0.05%，于50 ℃水浴中10 min，之后超声波处理15 s，用无菌海水稀释至最终浓度分别为10-3 g/mL和10-4 g/mL。将各样品稀释液150 µL分别涂布于7种分离培养基上，每一个稀释度样品重复涂布3个分离平板，28 ℃静置培养7 d。
1.6 菌株的分离纯化与保存
肉眼观察上述已培养的分离平板，挑取放线菌菌落于相应的琼脂斜面，28 ℃培养5-7 d；转接于斜面的分离菌株挑出进一步纯化，采用平板划线法进行单菌落分离，挑取纯化后的单菌落接种于相应的斜面培养基；将纯化平板上的菌落刮下，保存于50%灭菌甘油中，-80 ℃保藏。
1.7 基于16S rRNA基因序列的分子鉴定
菌株的基因组DNA提取、PCR反应体系和条件均参照梁效伟[18]的方法，采用细菌通用引物27F (5′-AGAGTTTGATCCTGGCTCAG-3′)和1492R (5′-GGTTACCTTGTTACGACTT-3′)进行16S rRNA基因的PCR扩增。PCR产物经0.8%的琼脂糖凝胶电泳检测验证后，其测序工作由苏州金唯智生物科技有限公司完成。测序结果于NCBI进行BLAST比对，选取近缘模式菌的16S rRNA基因序列，用MEGA 7.0的Neighbor-joining法构建系统发育树。
1.8 发酵粗提物抗真菌活性检测
1.8.1 放线菌发酵液粗提物的制备
将目标菌株接种于相应的培养基，于28 ℃恒温箱培养5 d，取菌丝体或者孢子接入3瓶装有150 mL发酵培养基的500 mL锥形瓶中，于28 ℃、180 r/min培养3-5 d。发酵液于5 000 r/min离心20 min，取菌渣浸泡于甲醇中1 d，再次离心获取甲醇上清液，之后于旋转蒸发仪浓缩成膏状；取上清液，等体积乙酸乙酯萃取3次，有机相同法浓缩至膏状；将上述2种浸膏合并待用。称取少量粗提物用甲醇溶解，配成浓度为20 mg/mL的母液，4 ℃冰箱中备用
1.8.2 待测液抗真菌活性的检测
采用琼脂打孔扩散试验，通过测量抑菌圈的大小直观地考察发酵液的抑菌性能。取制备好的107 CFU/mL菌悬液100 μL (白色念珠菌、茄病镰刀菌和黑曲霉孢子)均匀涂布到PDA琼脂平板上，将制备好的30 mg/mL发酵液100 µL滴加至直径为9 mm圆形孔洞上，置于培养箱中28 ℃ (真菌)培养，分别于24 h (白色念珠菌)和5 d (茄病镰刀菌和黑曲霉)后测量其抑菌圈直径。每个试验平行进行3次，取平均值。
1.8.3 化合物最小抑菌浓度(MIC)的测定
对化合物F2-1、F4-1和F5-1进行抗真菌活性测试，阳性对照为多菌灵。指示真菌为白色念珠菌ATCC 90028、茄病镰刀菌QY22和黑曲霉FJAT-31087。化合物和多菌灵DMSO溶解，采用试管稀释法，用培养基将化合物稀释为200、100、50、25、12.5、6.25 μg/mL的质量浓度，无菌试管为空白对照，加入100 μL菌悬液(1×106 CFU/mL)，28 ℃恒温培养72 h，每组平行实验3次，取平均值，以无菌生长的最低浓度作为化合物对该菌的MIC。
1.9 阳性菌株发酵粗提物的HPLC-DAD分析
取1.8制备的发酵粗提物，使用0.45 μm的滤膜过滤，色谱柱：Eclipse XDB-C18，4.6 mm×250 mm，进样量10 μL，流动相为甲醇-水(10%-100%，60 min)，流速为1 mL/min，检测采用二极管阵列检测器(diode array detector, DAD)检测器，波长扫描范围190-800 nm。
1.10 菌株W444的大规模摇瓶发酵
将菌株W444接种于W7固体培养基上，28 ℃培养4 d，刮取孢子接种于W7液体培养基中，28 ℃、250 r/min培养2 d获取种子液，将其按体积分数为5%的接种量接入20 L发酵培养基中，28 ℃、250 r/min培养4 d获取发酵液。发酵粗提物的制备同方法1.8.1。
1.11 化合物的分离和纯化
粗提物(8.5 g)经C18反相硅胶柱层析粗分，甲醇水梯度(10%-100%)洗脱，收集馏分，HPLC检测合并得馏分Fr.1-Fr.7。馏分Fr.2上样于Sephadex LH-20，甲醇洗脱得馏分Fr.2-11，Fr.2-11进行高压液相制备，48%甲醇水洗脱得到化合物F2-1 (10.5 mg)；馏分Fr.4上样于Sephadex LH-20，甲醇洗脱得馏分Fr.4-3，Fr.4-3进行高压液相制备，65%甲醇水洗脱得到化合物F4-1 (6.5 mg)；馏分Fr.5上样于Sephadex LH-20，甲醇洗脱得馏分Fr.5A-Fr.5C，Fr.5A进行高压液相制备，70%甲醇水洗脱，制备得到化合物F5-1 (8 mg)。化合物经质谱(ESI-MS)和NMR等波谱分析及与文献比较确定结构。
2 结果与分析
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2.1 菌株分离鉴定及不同红树林植物根际沉积物中的放线菌多样性分析
本研究通过16S rRNA基因序列的BLAST比对去重，并将结果提交至EzBioCloud数据库进行鉴定。表2中的“Similarity”列显示了分离菌株与数据库中相似度最高的参考菌株之间的相似性百分比，该数据通过16S rRNA基因序列比对得出。较高的相似度表明分离菌株与已知菌株在基因序列上具有密切的亲缘关系，为菌株的准确分类和鉴定提供了可靠依据。基于16S rRNA基因序列构建的系统发育树(图1)和表2的数据表明，从2种土壤样品中分离出56株放线菌，隶属于6目6科8属，包括链霉菌23株、小单孢菌19株、拟无枝酸菌6株、糖多孢菌2株，戈登氏菌3株、类诺卡氏菌1株、诺卡氏菌1株和疣孢菌1株。在所有分离的放线菌中，小单孢菌为主要类群，占总分离菌株的41.0%，其次是链霉菌属，占33.9%，其他属放线菌占总分离菌株的25.0%。
对2种红树林植物根际沉积物样品的放线菌进行多样性分析，发现从秋茄和互花米草的根际土样中获得的放线菌数目有一定差异，分别为36株(占比64.3%)和20株(占比35.7%) (表2和图2)。秋茄根际土样中小单孢菌属和链霉菌属分别占47%和39%，拟无枝酸属仅占8%，其他放线菌占6%。互花米草根际土样中放线菌的多样性最丰富，共分离到8个不同属的20株放线菌，链霉菌属占30%，小单孢菌属占25%，其他放线菌占45% (表2和图2)。综上所述，2种不同红树林植物根际土样中的放线菌数目和种属都有一定的差异。在秋茄根际土样中，链霉菌属和小单孢菌属为优势种群，占比为86%；而在互花米草根际土样中，优势种群为稀有放线菌，占比为75%。
2.2 菌株 Streptomycessp. W444发酵粗提物HPLC分析和抗真菌活性测试
本研究根据系统发育树的分类特征和菌株的摇瓶长势，筛选出21株放线菌进行小规模发酵，经HPLC分析发现菌株Streptomyces sp. W444在发酵培养基中发酵的次级代谢产物较为丰富(图3)。采用琼脂扩散法对该菌株的发酵粗提物进行抗真菌活性测试，特别是测试了Streptomyces sp. W444的发酵粗提物对白色念珠菌ATCC 90028、茄病镰刀菌QY22和黑曲霉FJAT-31087的抗真菌活性。测试结果如表3和图4所示，Streptomyces sp. W444的发酵粗提物对这3种指示菌均表现出较强的抑制活性。鉴于Streptomyces sp. W444发酵产物的高丰度及其良好的抗真菌活性，本研究决定将其作为后续次级代谢产物研究的候选菌株。
2.3 菌株 Streptomycessp. W444中化合物的分离和结构鉴定
对菌株Streptomyces sp. W444进行20 L扩大发酵，经乙酸乙酯萃取得8.5 g粗浸膏，通过中压反相色谱、Sephadex LH-20凝胶色谱和HPLC半制备等分离手段获得3个吲哚类化合物(F2-1、F4-1和F5-1)。利用质谱ESI-MS、1H NMR、13C NMR及与相关文献[19-22]比较分析，确定它们分别为streptochlorin、K252c和staurosporine (图5)。
化合物F2-1为白色粉末。UV (MeOH) λmax 221、271和286 nm处有特征吸收峰。ESI-MS m/z 219 [M+H]+。1H-NMR (MeOD，600 MHz) δ: 8.20 (1H, s, H-2), 8.02 (1H, d, H-4′), 7.84 (1H, s, H-2′), 7.46 (1H, d, H-7′), 7.22 (1H, t, H-6′), 7.16 (1H, t, H-5′)。13C-NMR (MeOD, 150 MHz) δ: 149.9 (d, C-2), 145.2 (s, C-5), 137.6 (s, C-7′a), 125.8 (s, C-3′a), 125.2 (d, C-2′), 123.7 (d, C-6′), 121.6 (d, C-5′), 121.4 (s, C-4), 121.2 (d, C-4′), 112.8 (d, C-7′), 103.4 (s, C-3′)。综合分析紫外、质谱及核磁等数据，表明该化合物与文献[19]报道的streptochlorin一致，确定它们同质。结构如图5A所示。
化合物4-1为淡黄色粉末状。UV (MeOH)λmax 240、288、333、360 nm处有特征吸收峰。ESI-MS m/z 310 [M-H]-。1H-NMR (MeOD，600 MHz)和13C-NMR (MeOD，150 MHz)数据见表4。综合上述数据表明，该化合物的理化性质、波谱性质与K252c[20]数据一致，结构如图5B所示。
化合物F5-1为淡黄色固体(MeOH)，在240、290和336 nm处有较强的紫外吸收。ESI-MS给出分子离子峰m/z 467.1 [M+H]+。1H-NMR (MeOD，600 MHz)和13C-NMR (MeOD，150 MHz)数据见表4。综合上述数据表明，该化合物的理化性质、波谱性质与化合物星孢菌素[21-22]数据一致，结构如图5C所示。
2.4 Staurosporine生物合成基因簇定位分析与生物合成途径推导
利用antiSMASH 7.1[23]对Streptomyces sp. W444的全基因组(该基因组已提交至GenBank，登录号为CP169412、CP169413和CP169414)进行比较分析，结果表明cluster 3与吲哚咔唑类化合物星形孢菌素(GenBank登录号为AB088119)[24]的生物合成基因簇具有60%的相似性(表5，图6A)。生物信息学分析表明，cluster 3含有33个基因，与星孢菌素合成相关的基因只有9个，其中负责吲哚[2,3-α]-吡咯并[3,4-c]咔唑母核结构合成相关的基因为orf4-6和orf9，它们分别与Streptomyces sp. TP-A0274的星孢菌素合成基因簇[25]中的staO (编码L-氨基酸氧化酶)、staD (编码色吡咯酸合成酶)、staP (编码细胞色素P450)以及staC (编码单氧化酶)编码的蛋白具有较高的相似性(表5)；基因orf7和orf8编码的N-甲基转移酶和O-甲基转移酶分别负责催化星孢菌素氨基糖苷(L-ristosamine)的 4′-NH和3-OH甲基化反应[26]，分别与星孢菌素合成基因簇中的staMA和staMB编码的蛋白具有较高的相似性(表5)；orf2和orf3分别编码细胞色素P450和N-糖基转移酶，这2个蛋白主要负责催化吲哚咔唑母核与糖基的偶联反应，分别与星孢菌素合成基因簇中的staN和staG编码的蛋白具有较高的相似性(表5)；orf1则负责编码转录激活蛋白，与星孢菌素合成基因簇中的staR编码的蛋白具有较高的相似性(表5)；此外，进一步分析发现该基因簇缺少糖基合成相关的基因，如staA (编码glucose-1-phosphate thymidyltransferase)、staB (编码dTDP-glucose 4,6-dehydratase)等。最后结合文献[25,27]对2个吲哚咔唑类化合物的生物合成途径进行了推导，结果表明星孢菌素母核的合成主要由L-色氨酸修饰、色吡咯酸的合成以及氧化闭环生成色咔唑3个步骤构成(图5)。L-色氨酸在L-氨基酸氧化酶(StaO)催化下生成吲哚-3-丙酮酸(indole-3-propionic acid, IPA)亚胺衍生物。之后，色吡咯酸合成酶(StaD)催化2分子IPA亚胺产生色吡咯酸(chromopyrrolic acid, CPA)。在糖基转移酶(StaG)催化下，K252c (F4-1)的N-13位与氨基糖的C-6′位发生脱水缩合反应，产生吲哚咔唑氮苷中间体holyrineA；在细胞色素P450酶(StaN)催化下holyrineA再次发生脱水缩合反应，使糖基的C-2′位和N-12位连接形成第二个C-N键，产生O-去甲基-N-去甲基星孢菌素，随后在甲基转移酶StaMA和StaMB的作用下糖基上-OH和-NH3分别发生甲基化反应，最终形成星形孢菌素(F5-1)，其生物合成途径如图6B所示。综合产物和基因簇分析表明，Streptomyces sp. W444 中cluster3主要负责星孢菌素类化合物的合成。
2.5 化合物的MIC值测定
通过MIC的测定发现，3个化合物对真菌均具有较好的抑制活性，结果如表6所示。其中，化合物F5-1对黑曲霉FJAT-31087的抑制活性最强，MIC值为25 μg/mL，且对3株测试菌均具有一定程度的抑制作用。此外，化合物F4-1对黑曲霉FJAT-31087和茄病镰刀菌QY22都具有一定的抑制活性，MIC值均为100 μg/mL。相比之下，化合物F2-1对测试真菌的抑菌活性较弱，仅表现出对茄病镰刀菌QY22的抑制活性，其MIC值为200 μg/mL。
3 讨论与结论
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红树林生态系统表现为生物群落复杂多样，其生物量占全球热带和亚热带沿海地区的60%-75%[28]，其中孕育着丰富独特的微生物资源。泉州湾洛阳江红树林主要分布于洛阳江、晋江出海口沿岸，此区域潮流作用强，东海暖流、南海暖流和黑潮分支等多种水系交汇，受此影响该区域蕴含着丰富的放线菌资源和潜在的结构新颖的代谢产物[29]。基于此，本研究选取了7种分离培养基，从泉州湾洛阳江不同区域采集2份红树林根际土壤样品，并从中分离鉴定了放线菌56株，隶属6目6科8属，其中链霉菌21株、小单孢菌23株、拟无枝酸菌6株、糖多孢菌2株，戈登氏菌3株、类诺卡氏菌1株和诺卡氏菌1株。从多样性分布可以看出，链霉菌属和小单孢菌属为优势菌群，该优势类群分布与何洁等[30]报道的印度洋红树林沉积物放线菌，以及何子邦等[31]报道的广西沿海红树林土壤放线菌的优势类群一致。从分菌土样来源不同分析，2种不同红树林植物根际土样中可培养放线菌的数目和多样性均呈现出一定差异，其中从互花米草根际土样中分离得到的稀有放线菌种属更为丰富，约占该样品中可培养放线菌总数的75%，表明互花米草根际土样中富含稀有放线菌，可作为稀有放线菌分离的重要资源。
结合抑菌活性筛选，选取Streptomyces sp. W444作为目标菌株，从中分离鉴定了3个吲哚类化合物，分别为streptochlorin、K252c和staurosporine。结构上，星孢菌素类化合物(K252c和staurosporine)作为吲哚咔唑类生物碱，表现出广泛的生物活性，如Streptomyces sp. ZS-A121产生了5个星孢菌素衍生物，其中一个新的星孢菌素衍生物streptomholyrine A对白色念珠菌(C. albicans)具有较强抑制活性，其MIC为12.5 μg/mL[32]，此外，本研究分离得到的K252c对大肠杆菌(Escherichia coli)以及金黄色葡萄球菌(Staphylococcus aureus)也具有较显著的抑制活性，其MIC均为50 μg/mL。同时，研究还表明星孢菌素对蛋白激酶C具有极强的抑制活性，其IC50值达到2.7 nmol/L[33]，因此其被作为科研工具化合物，用于细胞生命活动的追踪，如细胞信号传导以及细胞凋亡[34]；此外，星孢菌素衍生物也具有较强的细胞毒活性。例如Xiao等[35]通过异源表达从基因工程菌Streptomyces coelicolor M1146分离得到2个新的星孢菌素衍生物(staurosporines M1和M2)，生物活性分析表明staurosporines M1和M2对肿瘤细胞(HCT-116，K562和Huh 7.5)的生长具有显著的抑制活性。同时，研究还发现吲哚类化合物streptochlorin对植物病原性真菌Pythium dissimile、Botrytis cinerea、Zymoseptoria tritici、Pyriculariaory zae、Fusarium culmorum和Rhizoctonia solani等均具有较强的抑制活性[36]，显示出良好的生防菌潜能。基于此，抗真菌分析结果表明，Streptomyces sp. W444粗提物展示的优良抗真菌活性可能与该菌株所产的吲哚类化合物密切相关，分离得到的3个吲哚类化合物的抗真菌MIC值的测定结果进一步验证了上述结论。基因组分析表明，其合成基因簇cluster 3与星孢菌素的生物合成基因簇高度相似，主要包括负责吲哚[2,3-α]-吡咯并[3,4-c]咔唑母核合成相关的基因，进一步分析还发现该基因簇缺少糖基合成相关的基因，这与王冕等[37]报道的海洋链霉菌IMB3-202中的星孢菌素合成基因簇的构成一致，该发现也预示着cluster 3中的orf2和orf3所编码的细胞色素P450和N-糖基转移酶具有较宽的底物识别能力，能从初级糖代谢中识别底物，进而催化吲哚咔唑母核与糖基的偶联反应，该发现也为研究这2个酶的催化机制和合成新的星孢菌素衍生物奠定了基础，也预示着泉州湾红树林放线菌中蕴含着丰富的基因资源，值得进一步开发利用。
综上所述，本研究从泉州湾的2种红树林植物根际土样中分离鉴定了56株放线菌，并基于样品来源初步探讨了可培养放线菌的多样性，结果表明泉州湾红树林的放线菌资源丰富，发掘潜力巨大，为今后进一步开发利用泉州湾红树林药用放线菌资源奠定了基础。同时，综合菌株代谢产物的抗真菌活性，从Streptomyces sp. W444中鉴定了3个吲哚类化合物，定位了staurosporine类化合物的生物合成基因簇，并推导了其生物合成途径，拓宽了staurosporine类化合物的菌株来源，为其生物合成研究奠定了基础。
基金
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  • 泉州市科技计划(2023NS082)
  • 泉州医学高等专科学校校级课题重点科技项目(XJK2210A)
  • 自然资源部第三海洋研究所开放基金(HY202304)
  • 泉州医学高等专科学校研究型教学项目(XJJK2308)
文献
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2025年第65卷第2期
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doi: 10.13343/j.cnki.wsxb.20240584
  • 接收时间：2024-09-23
  • 首发时间：2026-02-05
  • 出版时间：2025-02-04
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  • 收稿日期：2024-09-23
  • 录用日期：2024-11-11
基金
Science and Technology Project of Quanzhou City(2023NS082)
泉州市科技计划(2023NS082)
Key Science and Technology Project of Quanzhou Medical College(XJK2210A)
泉州医学高等专科学校校级课题重点科技项目(XJK2210A)
Scientific Research Foundation of Third Institute of Oceanography, Ministry of Natural Resources(HY202304)
自然资源部第三海洋研究所开放基金(HY202304)
Research-based Teaching Project of Quanzhou Medical College(XJJK2308)
泉州医学高等专科学校研究型教学项目(XJJK2308)
作者信息
    1 泉州医学高等专科学校，福建 泉州
    2 福建省微生物研究所，福建 福州
    3 自然资源部第三海洋研究所，福建 厦门

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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
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