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Article(id=1241045265763856485, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1239895163967959761, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20230368, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1685116800000, receivedDateStr=2023-05-27, revisedDate=null, revisedDateStr=null, acceptedDate=1694966400000, acceptedDateStr=2023-09-18, onlineDate=1773817848864, onlineDateStr=2026-03-18, pubDate=1704297600000, pubDateStr=2024-01-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1773817848864, onlineIssueDateStr=2026-03-18, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1773817848864, creator=13701087609, updateTime=1773817848864, updator=13701087609, issue=Issue{id=1239895163967959761, tenantId=1146029695717560320, journalId=1192105938417971205, year='2024', volume='64', issue='1', pageStart='1', pageEnd='322', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=0, createTime=1773543643228, creator=13701087609, updateTime=1773820020328, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1241054373594320900, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1239895163967959761, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1241054373598515205, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1239895163967959761, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=208, endPage=219, ext={EN=ArticleExt(id=1241045266724352120, articleId=1241045265763856485, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Isolation and identification of a tomato endophyte FQ-G3 and its disease-resistant and growth-promoting properties, columnId=1241045257748533520, journalTitle=Acta Microbiologica Sinica, columnName=Research Articles, runingTitle=null, highlight=null, articleAbstract=

Gray mold caused byBotrytis cinerea is one of the major diseases affecting tomato production. The currently used fungicides are gradually restricted due to residues, pathogen resistance, and food safety. Therefore, screening out antagonistic microorganisms has gradually become an effective approach for the biocontrol of gray mold. [Objective] We screened out endophytic strains capable of endowing plants with disease resistance and promoting plant growth from tomato plants and evaluated their biocontrol potential, aiming to provide a theoretical basis for developing a new approach for the biocontrol of tomato gray mold.[Methods] The endophytic bacteria and fungi were isolated from different parts of tomato plants by the tissue culture method, and the candidate strains were preliminarily identified by 16S rRNA and ITS sequence analysis. The endophytes with antagonistic activity againstB.cinerea were screened by confrontation culture and fruit inoculationin vitro. Furthermore, we examined the abilities of the strain to secrete indole-3 acetic acid (IAA), protease, and siderophores and the promoting effects on the growth ofArabidopsis thaliana and tomato seedlings.[Results] A total of 72 endophytic bacterial strains and 31 endophytic fungal strains were isolated from different parts of tomato plants. An endophytic bacterium FQ-G3 with strong inhibitory activities against several pathogens was screened out and was identified asBacillus velezensis. FQ-G3 showed the inhibition rate of 80.93% againstB.cinereain vitro and inhibited the mycelial expansion on tomato fruitsin vivo. The strain could secrete IAA, protease, and siderophores, and promote the growth ofA.thaliana and tomato seedlings.[Conclusion] The endophytic strain FQ-G3 isolated from tomato plants endows plants with disease resistance and promotes plant growth, and thus can serve as a candidate for the prevention and control of gray mold. The findings enrich the tomato endophyte resources and provide support for the control of gray mold and the growth promotion of tomato.

, correspAuthors=Peiqian LI, authorNote=null, correspAuthorsNote=
*LI Peiqian, E-mail:
, copyrightStatement=Copyright ©2024 Acta Microbiologica Sinica. All rights reserved., 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=null, authorsList=Baozhen FENG, Peiqian LI, Jin LIU, Yanli YANG, Xiaojing WANG), CN=ArticleExt(id=1241045272852230425, articleId=1241045265763856485, tenantId=1146029695717560320, journalId=1192105938417971205, language=CN, title=番茄内生菌的分离鉴定及菌株FQ-G3抗病促生特性, columnId=1192149544164012138, journalTitle=微生物学报, columnName=研究报告, runingTitle=null, highlight=null, articleAbstract=

由灰葡萄孢(Botrytis cinerea)引起的灰霉病是番茄生产中最重要的病害之一，当前使用的杀菌剂因药物残留、病原菌抗药性及食品安全等原因逐渐受到限制。因此，利用拮抗微生物的生物防治逐渐成为灰霉病防控的有效策略。【目的】从番茄植株体内筛选具有抗病促生特性内生菌株并对其生防潜力进行评估，为开发番茄灰霉病生物防治新策略提供理论依据。【方法】采用组织分离法在番茄植株不同部位分离出内生细菌、真菌，结合16S rRNA和ITS序列分析，对候选菌株进行初步鉴定；通过菌株对峙培养、果实离体接种筛选对灰葡萄孢具有拮抗活性的内生菌；进一步测定菌株分泌生长素、嗜铁素的能力及其对拟南芥和番茄幼苗生长的促生特性。【结果】从番茄植株不同部位共分离出72株内生细菌和31株内生真菌，通过平板对峙法筛选出1株对多种病原菌具有较好抑菌活性的内生细菌FQ-G3，分子鉴定为Bacillus velezensis。FQ-G3对灰葡萄孢抑菌率达80.93%，并显著抑制灰葡萄孢在番茄果实上的扩展。该菌株能够分泌生长素、蛋白酶和嗜铁素，且对拟南芥、番茄幼苗具有明显的促生效果。【结论】本研究表明分离自番茄植株的内生菌FQ-G3具有抗病促生特性，对番茄灰霉病的防控具有潜在应用价值。研究结果丰富了番茄内生菌资源，为番茄灰霉病的防病促生提供技术和理论支持。

, correspAuthors=李培谦, authorNote=null, correspAuthorsNote=null, copyrightStatement=版权所有©《微生物学报》编辑部2024, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=vap+kqo5/keBbHn6fqReeg==, magXml=vP5nyrWeiLezBVIxwXAP7g==, pdfUrl=null, pdf=ZAbJoxXvdzlQYq7rm6+cuA==, pdfFileSize=1146654, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=5YcW9zrF73PUZ8HNqUmuYw==, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=hpm0pn6QM8dmsnBOU/mgMg==, mapNumber=null, authorCompany=null, fund=null, authors=null, authorsList=冯宝珍, 李培谦, 刘缙, 杨燕丽, 王晓静)}, authors=[Author(id=1241084432967324140, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, 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, 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inArabidopsis[J].Plant Physiology,2004,134(3):1017-1026., articleTitle=Bacterial volatiles induce systemic resistance inArabidopsis, refAbstract=null), Reference(id=1241084446456205381, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, doi=null, pmid=null, pmcid=null, year=2003, volume=100, issue=8, pageStart=4927, pageEnd=4932, url=null, language=null, rfNumber=[32], rfOrder=36, authorNames=null, journalName=Proceedings of the National Academy of Sciences of the United States of America, refType=null, unstructuredReference=RYU CM, FARAG MA, HU CH, REDDY MS, WEI HX, PARÉ PW, KLOEPPER JW.Bacterial volatiles promote growth inArabidopsis[J].Proceedings of the National Academy of Sciences of the United States of America,2003,100(8):4927-4932., articleTitle=Bacterial volatiles promote growth inArabidopsis, refAbstract=null), Reference(id=1241084446552674376, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, 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S, SHAHZAD R, HUSSAIN J, LEE IJ, AL-HARRASI A.Endophytes from medicinal plants and their potential for producing indole acetic acid, improving seed germination, and mitigating oxidative stress[J].Journal of Zhejiang University Science B,2017,18(2):125-137., articleTitle=Endophytes from medicinal plants and their potential for producing indole acetic acid, improving seed germination, and mitigating oxidative stress, refAbstract=null), Reference(id=1241084446733029457, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, doi=10.1007/s10327-016-0648-9, pmid=null, pmcid=null, year=2016, volume=82, issue=3, pageStart=165, pageEnd=173, url=null, language=null, rfNumber=[35], rfOrder=39, authorNames=null, journalName=Journal of General Plant Pathology, refType=null, unstructuredReference=LIU XY, DOU GM, MA YC.Potential of endophytes from medicinal plants for biocontrol and plant growth promotion[J].Journal of General Plant Pathology,2016,82(3):165-173., 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figs=[ArticleFig(id=1241084435802673858, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=EN, label=Figure 1, caption=Antagonistic effect of FQ-G3 on four kinds of plant pathogens. A: Q-G3 againstBotrytiscinerea. B: FQ-G3 againstCladosporium sp.. C: FQ-G3 againstFusarium pemambucanum. D: FQ-G3 againstAlternaria alternata., figureFileSmall=BUKqYTmniolaTpgsQzDXHA==, figureFileBig=jB18qCuLNrPnqAL67gGjGA==, tableContent=null), ArticleFig(id=1241084437287457481, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=CN, label=图1, caption=菌株FQ-G3与四种病原菌对峙培养, figureFileSmall=BUKqYTmniolaTpgsQzDXHA==, figureFileBig=jB18qCuLNrPnqAL67gGjGA==, tableContent=null), ArticleFig(id=1241084437409092310, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=EN, label=Figure 2, caption=Phylogenetic trees of FQ-G3 based on 16S rRNA gene. The ex-type strains are noted using T. GenBank accession numbers are in parentheses. Numbers at the nodes indicate the level of bootstrap values based on 1 000 replications. The scale bar indicates 0.002 substitutions per nucleotide position., figureFileSmall=/rHO3ALgmS/GVzkr5ZEUkw==, figureFileBig=hDhBEfgPkeFe78FTyI4b6w==, tableContent=null), ArticleFig(id=1241084437505561312, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=CN, label=图2, caption=基于16S rRNA基因构建的系统发生树, figureFileSmall=/rHO3ALgmS/GVzkr5ZEUkw==, figureFileBig=hDhBEfgPkeFe78FTyI4b6w==, tableContent=null), ArticleFig(id=1241084437623001832, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=EN, label=Figure 3, caption=Phylogenetic trees of FQ-G3 based ongyrA (A) andgyrB (B) genes. GenBank accession numbers are in parentheses. Numbers at the nodes indicate the level of bootstrap values based on 1 000 replications. The scale bar indicates 0.01 substitutions per nucleotide position., figureFileSmall=Y/EQKgQwJg+F7BZbhIAgzw==, figureFileBig=4Ce38+iaCVMHsNKf/3dyag==, tableContent=null), ArticleFig(id=1241084437702693614, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=CN, label=图3, caption=基于gyrA和gyrB基因构建的系统发生树, figureFileSmall=Y/EQKgQwJg+F7BZbhIAgzw==, figureFileBig=4Ce38+iaCVMHsNKf/3dyag==, tableContent=null), ArticleFig(id=1241084437773996790, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=EN, label=Figure 4, caption=In vivo inhibition of strain FQ-G3 againstBotrytis cinerea. A: Cherry tomatoes were mock inoculated with FQ-Q3 andB.cinerea, cherry tomatoes inoculated withB.cinerea as control. B: Average diameters of lesions of treated tomatoes. The bar and error bar represented the mean and standard error, respectively. Different letters indicated significant differences among groups (P < 0.05)., figureFileSmall=sTGkW41xk83CzKS3qMeBpA==, figureFileBig=ksc/PZiVLtWeHMlYFTQuJw==, tableContent=null), ArticleFig(id=1241084437862077181, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=CN, label=图4, caption=菌株FQ-G3对灰葡萄孢的活体抑制作用, figureFileSmall=sTGkW41xk83CzKS3qMeBpA==, figureFileBig=ksc/PZiVLtWeHMlYFTQuJw==, tableContent=null), ArticleFig(id=1241084437958546178, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=EN, label=Figure 5, caption=Promotion effect of endophytic strain FQ-G3 toArabidopsis thaliana seedlings. A: The NA medium was used as the blank control. B: FQ-G3 was streaked on NA agar. C:A.thaliana root hairs in control group. D:A.thaliana root hairs in test group., figureFileSmall=RSkjryGsuu+PS50V4SVs4g==, figureFileBig=VRCQ4oJ4MyoRlcjvvomwlw==, tableContent=null), ArticleFig(id=1241084438067598089, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=CN, label=图5, caption=内生菌FQ-G3对拟南芥的促生作用, figureFileSmall=RSkjryGsuu+PS50V4SVs4g==, figureFileBig=VRCQ4oJ4MyoRlcjvvomwlw==, tableContent=null), ArticleFig(id=1241084438155678478, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=EN, label=Figure 6, caption=Physiological index ofArabidopsis thaliana seedlings. A: Fresh weight ofA.thaliana seedlings (n=30). B: The average length of theA.thaliana roots (n=30). C: Number ofA.thaliana root hairs (n=30). The bar and error bar represented the mean and standard error, respectively (n=30). Different letters indicated significant differences among groups (P < 0.05)., figureFileSmall=dpboOI3zuVN8Ua8KMvTv3Q==, figureFileBig=+J1ZhB7IX9m5hxDDG1CAoA==, tableContent=null), ArticleFig(id=1241084438226981652, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=CN, label=图6, caption=拟南芥幼苗生理指标, figureFileSmall=dpboOI3zuVN8Ua8KMvTv3Q==, figureFileBig=+J1ZhB7IX9m5hxDDG1CAoA==, tableContent=null), ArticleFig(id=1241084438298284824, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=EN, label=Figure 7, caption=Growth-promoting effect of the strain FQ-G3 to tomato seedlings. A: FQ-G3 treated tomato plant, seedling treated with sterile water used as control. B: Number of tomato compound leaves (n=24). C: Height of tomato seedlings (n=24). D: Length of tomato seedlings taproots (n=24). E: Number of tomato seedlings lateral roots (n=24). The bar and error bar represented the mean and standard error, respectively (n=24). Different letters indicated significant differences among groups (P < 0.05)., figureFileSmall=Jr9WggSY+B3tYka7uePgAQ==, figureFileBig=ZSBGuaXxOVOAnJnJFqkzLg==, tableContent=null), ArticleFig(id=1241084438419919649, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=CN, label=图7, caption=菌株FQ-G3对番茄苗的促生作用, figureFileSmall=Jr9WggSY+B3tYka7uePgAQ==, figureFileBig=ZSBGuaXxOVOAnJnJFqkzLg==, tableContent=null), ArticleFig(id=1241084438516388650, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=EN, label=Table 1, caption=

Primary identification of endophytes from tomato

, figureFileSmall=null, figureFileBig=null, tableContent=
ClassificationIsolates (accession No.)Identity to the closest species (accession No.)
BacteriaFQ-G3 (MZ827471)99% toBacillus velezensis (MT271916)
FQ-G4 (MZ827472)99% toBacillus velezensis (MK310268)
FQ-G5 (MZ827473)99% toBacillus amyloliquefaciens (MT685260)
FQ-G6 (MZ827474)99% toBacillus velezensis (KY887762)
FQ-G8 (MZ827475)99% toBacillus subtilis (MG751326)
FQ-G12 (MZ827476)99% toBacillus subtilis (MF136610)
FQ-G13 (MZ827477)99% toBacillus amyloliquefaciens (MH265996)
FQ-G17 (MZ827478)99% toEscherichia fergusonii (MN208072)
FQ-J2 (MZ827479)99% toBacillus subtilis (MF136610)
FungiFQ-B (ON045560)99% toAspergillus niger (MZ819922)
FQ-G10 (ON045555)99% toDidymella macrostoma (MN420662)
FQ-14 (ON045556)99% toDidymella microchlamydospora (MH862504)
FQ-G16 (ON045557)99% toAspergillus sydowii (KU687805)
FQ-J1 (ON045558)99% toCladosporium cladosporioides (MT367253)
FQ-J4 (ON045559)99% toAlternaria alternata (MN856355)
), ArticleFig(id=1241084438596080430, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=CN, label=表1, caption=

番茄内生菌的初步鉴定结果

, figureFileSmall=null, figureFileBig=null, tableContent=
ClassificationIsolates (accession No.)Identity to the closest species (accession No.)
BacteriaFQ-G3 (MZ827471)99% toBacillus velezensis (MT271916)
FQ-G4 (MZ827472)99% toBacillus velezensis (MK310268)
FQ-G5 (MZ827473)99% toBacillus amyloliquefaciens (MT685260)
FQ-G6 (MZ827474)99% toBacillus velezensis (KY887762)
FQ-G8 (MZ827475)99% toBacillus subtilis (MG751326)
FQ-G12 (MZ827476)99% toBacillus subtilis (MF136610)
FQ-G13 (MZ827477)99% toBacillus amyloliquefaciens (MH265996)
FQ-G17 (MZ827478)99% toEscherichia fergusonii (MN208072)
FQ-J2 (MZ827479)99% toBacillus subtilis (MF136610)
FungiFQ-B (ON045560)99% toAspergillus niger (MZ819922)
FQ-G10 (ON045555)99% toDidymella macrostoma (MN420662)
FQ-14 (ON045556)99% toDidymella microchlamydospora (MH862504)
FQ-G16 (ON045557)99% toAspergillus sydowii (KU687805)
FQ-J1 (ON045558)99% toCladosporium cladosporioides (MT367253)
FQ-J4 (ON045559)99% toAlternaria alternata (MN856355)
), ArticleFig(id=1241084438679966518, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=EN, label=Table 2, caption=

Determination of bacteriostatic spectrum of strain FQ-G3

, figureFileSmall=null, figureFileBig=null, tableContent=
Plant pathogensColony diameter after treatment (cm)Colony diameter of control (cm)Inhibition rate (%)
Different letters indicated significant differences among values atP < 0.05.
Botrytis cinerea1.23±0.08a4.33±0.12b80.93
Alternaria alternata2.21±0.22a4.20±0.13b53.78
Fusarium pemambucanum1.97±0.45a4.24±0.12b60.69
Cladosporium sp.1.54±0.21a4.28±0.10b72.48
), ArticleFig(id=1241084438772241215, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=CN, label=表2, caption=

菌株FQ-G3抑菌率的测定

, figureFileSmall=null, figureFileBig=null, tableContent=
Plant pathogensColony diameter after treatment (cm)Colony diameter of control (cm)Inhibition rate (%)
Different letters indicated significant differences among values atP < 0.05.
Botrytis cinerea1.23±0.08a4.33±0.12b80.93
Alternaria alternata2.21±0.22a4.20±0.13b53.78
Fusarium pemambucanum1.97±0.45a4.24±0.12b60.69
Cladosporium sp.1.54±0.21a4.28±0.10b72.48
), ArticleFig(id=1241084438856127300, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=EN, label=Table 3, caption=

Enzyme-producing and growth-promoting characteristics of strain FQ-G3

, figureFileSmall=null, figureFileBig=null, tableContent=
StrainEnzyme producing and plant growth-promoting attributes
IAAProteaseChitinaseSiderophoresPhosphate solubilization
+: Positive; ++: Strong positive; –: Negative.
CK–––––
FQ-G3+++++++–
), ArticleFig(id=1241084438969373513, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1241045265763856485, language=CN, label=表3, caption=

菌株FQ-G3的产酶及促生特性

, figureFileSmall=null, figureFileBig=null, tableContent=
StrainEnzyme producing and plant growth-promoting attributes
IAAProteaseChitinaseSiderophoresPhosphate solubilization
+: Positive; ++: Strong positive; –: Negative.
CK–––––
FQ-G3+++++++–
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番茄内生菌的分离鉴定及菌株FQ-G3抗病促生特性
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冯宝珍 , 李培谦 * , 刘缙 , 杨燕丽 , 王晓静
微生物学报 | 研究报告 2024,64(1): 208-219
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微生物学报 | 研究报告 2024, 64(1): 208-219
番茄内生菌的分离鉴定及菌株FQ-G3抗病促生特性
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冯宝珍, 李培谦* , 刘缙, 杨燕丽, 王晓静
作者信息
  • 运城学院生命科学系, 山西 运城 044000
Isolation and identification of a tomato endophyte FQ-G3 and its disease-resistant and growth-promoting properties
Baozhen FENG, Peiqian LI* , Jin LIU, Yanli YANG, Xiaojing WANG
Affiliations
  • Life Sciences Department, Yuncheng University, Yuncheng 044000, Shanxi, China
出版时间: 2024-01-04 doi: 10.13343/j.cnki.wsxb.20230368
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摘要
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由灰葡萄孢(Botrytis cinerea)引起的灰霉病是番茄生产中最重要的病害之一，当前使用的杀菌剂因药物残留、病原菌抗药性及食品安全等原因逐渐受到限制。因此，利用拮抗微生物的生物防治逐渐成为灰霉病防控的有效策略。【目的】从番茄植株体内筛选具有抗病促生特性内生菌株并对其生防潜力进行评估，为开发番茄灰霉病生物防治新策略提供理论依据。【方法】采用组织分离法在番茄植株不同部位分离出内生细菌、真菌，结合16S rRNA和ITS序列分析，对候选菌株进行初步鉴定；通过菌株对峙培养、果实离体接种筛选对灰葡萄孢具有拮抗活性的内生菌；进一步测定菌株分泌生长素、嗜铁素的能力及其对拟南芥和番茄幼苗生长的促生特性。【结果】从番茄植株不同部位共分离出72株内生细菌和31株内生真菌，通过平板对峙法筛选出1株对多种病原菌具有较好抑菌活性的内生细菌FQ-G3，分子鉴定为Bacillus velezensis。FQ-G3对灰葡萄孢抑菌率达80.93%，并显著抑制灰葡萄孢在番茄果实上的扩展。该菌株能够分泌生长素、蛋白酶和嗜铁素，且对拟南芥、番茄幼苗具有明显的促生效果。【结论】本研究表明分离自番茄植株的内生菌FQ-G3具有抗病促生特性，对番茄灰霉病的防控具有潜在应用价值。研究结果丰富了番茄内生菌资源，为番茄灰霉病的防病促生提供技术和理论支持。

灰霉病  /  内生菌  /  鉴定  /  生物防治  /  促生作用

Gray mold caused byBotrytis cinerea is one of the major diseases affecting tomato production. The currently used fungicides are gradually restricted due to residues, pathogen resistance, and food safety. Therefore, screening out antagonistic microorganisms has gradually become an effective approach for the biocontrol of gray mold. [Objective] We screened out endophytic strains capable of endowing plants with disease resistance and promoting plant growth from tomato plants and evaluated their biocontrol potential, aiming to provide a theoretical basis for developing a new approach for the biocontrol of tomato gray mold.[Methods] The endophytic bacteria and fungi were isolated from different parts of tomato plants by the tissue culture method, and the candidate strains were preliminarily identified by 16S rRNA and ITS sequence analysis. The endophytes with antagonistic activity againstB.cinerea were screened by confrontation culture and fruit inoculationin vitro. Furthermore, we examined the abilities of the strain to secrete indole-3 acetic acid (IAA), protease, and siderophores and the promoting effects on the growth ofArabidopsis thaliana and tomato seedlings.[Results] A total of 72 endophytic bacterial strains and 31 endophytic fungal strains were isolated from different parts of tomato plants. An endophytic bacterium FQ-G3 with strong inhibitory activities against several pathogens was screened out and was identified asBacillus velezensis. FQ-G3 showed the inhibition rate of 80.93% againstB.cinereain vitro and inhibited the mycelial expansion on tomato fruitsin vivo. The strain could secrete IAA, protease, and siderophores, and promote the growth ofA.thaliana and tomato seedlings.[Conclusion] The endophytic strain FQ-G3 isolated from tomato plants endows plants with disease resistance and promotes plant growth, and thus can serve as a candidate for the prevention and control of gray mold. The findings enrich the tomato endophyte resources and provide support for the control of gray mold and the growth promotion of tomato.

Botrytis cinerea  /  endophyte  /  identification  /  biocontrol  /  growth promotion
冯宝珍, 李培谦, 刘缙, 杨燕丽, 王晓静. 番茄内生菌的分离鉴定及菌株FQ-G3抗病促生特性. 微生物学报, 2024 , 64 (1) : 208 -219 . DOI: 10.13343/j.cnki.wsxb.20230368
Baozhen FENG, Peiqian LI, Jin LIU, Yanli YANG, Xiaojing WANG. Isolation and identification of a tomato endophyte FQ-G3 and its disease-resistant and growth-promoting properties[J]. Acta Microbiologica Sinica, 2024 , 64 (1) : 208 -219 . DOI: 10.13343/j.cnki.wsxb.20230368
正文
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灰葡萄孢(Botrytis cinerea)引起的灰霉病是造成番茄采后腐烂的重要原因[1]。目前，对灰霉病的控制仍然是以化学防治为主，由此带来的食品安全和环境安全等问题已成为人们关注的重要问题[2]。生物防治已经成为植物病害控制的重要途径。大量的研究已经证实了植物内生菌有防病促生的作用。植物内生菌(endophytes)是指生活史的全部或部分阶段在植物组织内部，并且不引起植物发生明显组织病害症状的微生物。内生菌普遍存在于水生和陆生植物中，在植物的生命活动中发挥着重要作用[3]。与植物病原菌不同，植物内生菌普遍且长期栖息在宿主植物体内，并与宿主长期协同进化成为了植物微生物环境的重要组成部分[4]。
植物内生菌在促进植物生长、增强植物抗逆能力等方面都有着无限的潜力，同时，植物内生菌还可以通过生物固氮、分泌激素类物质等促进宿主植物的生长和产量。大量研究表明，内生菌对植物的促进作用主要是通过分泌生长素、赤霉素等直接促进植物生长，同时还能通过固氮、溶磷作用、产生嗜铁素等间接促进生长[5-6]。内生菌还能通过调节植物的免疫系统或者直接产生活性物质抑制病原菌来增强植物的抗病性[7-8]。植物内生细菌不仅种类多、繁殖快、代谢产物丰富，而且易于培养，现如今化学农药引起的一系列环境污染情况下，植物内生菌的研发有利于病害的生物防治、污染土壤的修复，同时还有利于减少农药的使用，更有利于促进全球农业的绿色可持续发展，因此植物内生菌的研究和开发具有非常重要的应用前景[9]。
本研究从番茄组织中分离筛选出抗病促生功能的贝莱斯芽孢杆菌，其在体内外均对灰葡萄孢具有强烈的抑制作用，并对拟南芥和番茄幼苗生长表现出促生作用。本研究丰富了番茄内生菌资源，也为灰霉病防治奠定了理论基础。
1 材料与方法
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1.1 主要材料、试剂和仪器
本研究中所用到的植物病原真菌灰葡萄孢B.cinerea LK7、Cladosporium sp.、Fusarium pemambucanum及Alternaria alternata等均分离于腐烂果蔬的表面，经纯化鉴定后保存于本实验室[10-11]。
试剂：细菌DNA快速提取试剂盒、真菌基因组DNA快速提取试剂盒，北京艾德莱生物科技有限公司；引物对27F/1492R、ITS1/ITS4，生工生物工程(上海)股份有限公司；2×Taq PCR MasterMix Ⅱ，TIANGEN；牛肉膏、酵母膏、蛋白胨、葡萄糖和琼脂粉等常规试剂均为国产分析纯。
仪器：BMJ-250型培养箱，上海博迅医疗生物仪器股份有限公司；Olympus CX43显微镜，奥林巴斯公司；T100TM Thermal Cycler PCR仪，伯乐公司。
1.2 番茄组织内生菌的分离
从山西运城、吕梁等地采集健康番茄果实、根、茎和叶样本。参照之前的消毒方法处理标本[10]，具体如下：先分别称取15 g番茄果实、根、茎和叶，经自来水冲洗晾干，再用75%酒精充分浸泡3−5 min (叶片浸泡时间偏短，根、茎和果实处理时间偏长)，然后用无菌水冲洗3−5次；再用5.4%次氯酸钠溶液浸泡2−5 min，用无菌水冲洗4−5次，晾干；研钵内碾碎后移入试管中加10 mL无菌水，静止放置15−20 min，吸取100 μL研磨液，将其分别涂布于potato dextrose agar (PDA)和nutrient agar (NA)平板。平板置于25黑暗培养20 d，每天观察菌落的生长情况；最后根据菌落的颜色、形态、光泽以及大小差异等挑取单菌落，将菌落纯化后统一编号置于NA或者PDA斜面培养基中于4保存备用。消毒可靠性检验参照前人研究方法[10]进行。
1.3 内生菌株的初步鉴定
选择菌落形态明显不同的菌株继续培养，提取基因组DNA。细菌用细菌基因组DNA提取试剂盒提取DNA，以细菌16S rRNA通用引物对27F/1492R[11]作为扩增引物进行PCR扩增，反应条件参照文献[11]进行。真菌用真菌基因组试剂盒，以真菌ITS通用引物ITS1/ITS4[10]进行PCR扩增，反应条件参照以报道文献[10]进行。PCR扩增产物于陕西杨凌天润奥科生物科技有限公司进行测序，序列上传GenBank。菌株序列通过NCBI BLAST (https://blast.ncbi.nlm.nih.gov/Blast)在线比对分析，进行初步鉴定。
1.4 抑菌内生菌株的筛选和鉴定
初筛：将病原菌接种在PDA培养基24 ℃培养5 d，用直径5 mm的打孔器在菌落边缘打取菌饼备用。内生细菌在NA平板28 ℃培养48 h，打菌饼备用。采用平板对峙法，先将病原菌饼置于PDA平板中心，在距中心2 cm的上下左右位置分别放置4块内生细菌菌饼，对峙平板置于24 ℃培养5 d。每天观察抑菌带情况，初步筛选出具有抑菌活性的候选菌株。
复筛：在PDA培养基中间划线法接种候选细菌，在距细菌2.0 cm的两侧放置病原菌菌饼，以只接病原菌菌饼为对照组，24 ℃培养5 d，测量病原菌菌落直径，待对照组病原菌菌丝长满平板后停止测量。抑菌率参照文献[7]计算，每组处理重复3次。
PCR扩增：为进一步明确候选菌株分类地位，用细菌基因组DNA提取试剂盒提取DNA，扩增DNA旋回酶亚基A基因(gyrA)和DNA拓扑异构酶亚基B基因(gyrB)并测序。gyrA和gyrB扩增引物及PCR反应条件均参照文献[11]。PCR扩增产物于陕西杨凌天润奥科生物科技有限公司进行测序，序列上传GenBank。
进化树构建：使用MEGA-X软件以邻接法(neighbour-joining, NJ)对ITS序列、gyrA和gyrB区序列构建系统发生树，分析该菌与其他菌株的亲缘关系，以确定其分类地位[12]。
1.5 菌株FQ-G3抑制灰葡萄孢侵染果实
将菌株FQ-G3用NB (nutrient broth)液体培养基，28 ℃、130 r/min培养48 h得到新鲜菌液。取大小均匀，成熟度一致的健康樱桃番茄，用75%酒精消毒5 min后，用无菌水冲洗2遍，置于无菌操作台内晾干备用。用打孔器在果实赤道处打孔(深度约2 mm，直径5 mm)，将20 µL FQ-G3的新鲜菌液和20 µL灰葡萄孢子悬浮液(1×106)接种至伤口处。每个处理接种10个果实，重复3次。对照组仅接种灰葡萄孢。处理后的果实置于无菌保鲜盒内，于24 ℃黑暗环境放置，观察果实发病情况，测量病斑大小。
1.6 菌株FQ-3的生理生化特性
吲哚乙酸(indole acetic acid, IAA)测定：将菌株FQ-G3按2%的接种量分别接种至含5 mmol/L色氨酸的LB液体培养基中，37 ℃、130 r/min摇床培养24 h，随后将菌液于离心机中8 000 r/min离心15 min后，取上清液1 mL，加入聚合氯化硫酸铁(polyferric chloride sulfate, PC)显色剂(1 mL 0.5 mol FeCl3+80 mL 37.5% H2SO4) 4 mL混匀，避光静置30 min进行显色反应后观察颜色变化。以加入50 μL浓度为10 mg/L的植物生长激素的LB培养基为阳性对照，以含有色氨酸的LB培养基为阴性对照，每组处理重复3次。观察颜色变化，若能产生IAA其混合液在静置后会变为粉红或红色。
蛋白酶测定：将菌株FQ-G3按2%的接种量接种至LB液体培养基中，37 ℃、130 r/min摇床培养24 h，分别用三点法或稀释涂布法平板法接种至脱脂牛奶培养基[13]平板，以涂布LB培养基为对照，每组处理重复3次。将平板封口后放置于28 ℃恒温培养箱中，每天观察培养基变化。观察菌落周围是否有透明圈形成，有透明圈出现则说明菌株具有产蛋白酶能力，通过计算透明圈直径与菌落直径的比值确定菌株产蛋白酶能力大小。
几丁质酶测定：将菌株FQ-G3按2%的接种量接种至LB液体培养基中，37 ℃、130 r/min摇床培养24 h，将菌株分别用三点法和稀释涂布法接种至几丁质酶诱导固体培养基[(NH4)2SO4 3 g、K2HPO4 0.7 g、KH2PO4 0.3 g、MgSO4·7H2O 0.5 g、MgSO4·7H2O 0.1 g、胶体几丁质3 g、酵母粉3 g、琼脂13 g、蒸馏水1 000 mL，pH 7.2]，以接种LB培养基为对照，每组处理重复3次，将平板封口后放置于28 ℃恒温培养箱中培养，每天观察培养基变化。观察菌落周围是否有透明圈形成，有透明圈出现则说明菌株具有产几丁质酶能力。
嗜铁素测定：将菌株FQ-G3按2%的接种量接种至LB液体培养基中，37 ℃、130 r/min摇床培养24 h，将培养好菌液在高速离心机中8 000 r/min离心15 min，取离心后的上清液和铬奥醇(chrome azurols, CAS)[14]按1:1等比例混合，避光显色2 h，观察颜色变化并记录。
溶磷测定：将菌株FQ-G3按2%的接种量接种至LB液体培养基中，37 ℃、130 r/min摇床培养24 h，将菌株分别用三点法和稀释涂布法接种至固体培养基[葡萄糖10 g、(NH4)2SO4 0.5 g、NaCl 0.3 g、KCl 0.3 g、MgSO4·7H2O 0.3 g、FeSO4·7H2O 0.03 g、MnSO4·4H2O 0.03 g、酵母粉0.4 g、Ca3PO4 10 g、琼脂20 g、蒸馏水1 000 mL，pH 7.0]，以接种LB培养基为对照，每组处理重复3次，将平板封口后放置于28 ℃恒温培养箱中培养，每天观察菌落周围能否形成透明圈。
1.7 拟南芥培养试验
二分隔培养皿法：参照前人研究方法[15]略加改进，分别将10 mL MS (Murashige and Skoog)和NA培养基分别倒入二分隔培养皿中，冷却后将消毒的拟南芥种子轻轻点在MS培养基上，同时用灭菌牙签将菌株GQ-G3接种到NA培养基上。以空白的MS/NA二分隔培养为对照，试验设3个重复。将处理好的培养皿置于20 ℃光照培养箱中，设置16 h/8 h (光照/黑暗)培养条件。培养21 d，测量拟南芥幼苗的鲜重、根长度以及侧根数量等参数。
1.8 番茄幼苗盆栽试验
将“粉冠”番茄种子在30 ℃的水浴锅处理2 h，再用75%乙醇浸泡5 min，无菌水清洗5−6次，然后将种子置于铺有浸润的无菌滤纸的培养皿中，于28 ℃的培养箱中进行催芽。
将灭菌育苗基质装至直径16 cm，高14 cm的花盆中，每个花盆种植一颗上述方法催芽的番茄种子，覆盖基质后放入智能温室出苗。智能温室昼/夜光照为12 h/12 h，温度26−30 ℃，光照强度60 klx，相对湿度65%。播种15 d番茄长出真叶时，第一次浇灌FQ-G3发酵液(每盆10 mL，浓度108 CFU/mL)。每隔7 d浇灌1次，连续3次。每组处理8棵幼苗，重复3次。对照组浇灌无菌水，幼苗长至60 d时，将番茄苗拔出，统计株高、叶片数量、根毛数量及根长。
1.9 数据分析
所有测量数据均使用SPSS 27.0进行统计分析。采用t检验比较不同处理组之间平均值，P < 0.05。
2 结果与分析
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2.1 内生菌的分离鉴定
从番茄植株不同部位共分离得到72株内生细菌和31株内生真菌，挑选菌落形态不同的菌株进行测序鉴定。将获得的细菌16S rRNA和真菌ITS序列通过NCBI在线比对，获得番茄内生菌株初步鉴定结果。如表1所示，番茄内生细菌主要为芽孢杆菌属(Bacillus)和埃希菌属(Escherichia)。而内生真菌多为亚隔孢壳属(Didymella)、曲霉属(Aspergillus)、枝孢霉属(Cladosporium)和链格孢属(Alternaria)。
2.2 抑菌内生菌的筛选及鉴定
通过初筛和复筛，得到一株对灰葡萄孢具有强烈拮抗作用的内生细菌FQ-G3，抑菌率达80.93% (表2)。如图1所示，FQ-G3对果蔬贮藏期病原菌Cladosporium sp.、Fusarium pemambucanum及Alternaria alternata也表现出不同程度的抑制作用，抑菌率分别为72.48%、60.69%及53.78%，说明FQ-G3具有开发为果蔬采后病害生物防治菌剂的潜力。
2.3 菌株FQ-G3的分子鉴定
扩增测序结果表明菌株FQ-G3的gyrA基因序列全长951 bp，GenBank No.: OP867063；gyrB基因序列全长1 209 bp, GenBank No.: OP867064。使用MEGA-X软件以NJ法分别对菌株16S rRNA、gyrA和gyrB序列构建系统发生树。16S rRNA序列比对，发现FQ-G3的16S rRNA (MZ827471)与模式菌株Bacillus velezensis CR-502相似度最高，为99.64%。下载相似性高的模式菌株序列与FQ-G3的16S rRNA (MZ827471)构建系统发育树，结果显示FQ-G3与Bacillus velezensis CR-502聚为一支(图2)。如图3所示，基于gyrA和gyrB基因的系统发生树FQ-G3均与B.velezensis聚为一支。以上结果表明FQ-G3属于B.velezensis。
2.4 菌株FQ-G3对灰葡萄孢的体内抑制作用
接种处理结果表明，菌株FQ-G3能够抑制灰葡萄孢对番茄果实的侵染(图4A)。在接种5 d后，对照组果实布满灰霉菌丝，发病率100%，而处理组番茄果实依然保持完好；同时处理组果实病斑直径明显小于对照组(图4B)。该结果说明菌株FQ-G3可以明显抑制灰葡萄孢在番茄果实上的扩展，具有活体生防潜力。
2.5 菌株FQ-G3生理生化特性分析
如表3所示，参试菌株FQ-G3具有产生IAA、蛋白酶、嗜铁素、蛋白酶和几丁质酶的能力。菌株FQ-G3能够产生IAA、嗜铁素，说明该菌株对植物生长具有一定促进作用。此外，该菌株能够产生蛋白酶和几丁质酶，这可能与抑菌作用有关。
2.6 菌株FQ-G3产生挥发性物质(volatile organic compounds, VOCs)对拟南芥的促生作用
利用二分隔培养皿将拟南芥与内生菌株共培养21 d，进一步验证B.velezensis FQ-G3的促生特性。结果显示处理组的拟南芥长势明显优于对照组，幼苗鲜重、根长及根毛数量等指标均高于对照组(P < 0.05)，说明FQ-G3能够产生挥发性促生物质(图5、图6)。
2.7 番茄苗盆栽实验
经过FQ-G3发酵液灌根处理的番茄苗长势优于对照组，其株高、叶片数量、根长、侧根数量与对照组相比，均有显著提高(P < 0.05) (图7)。
3 讨论与结论
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本研究利用组织分离法对番茄内生细菌和真菌进行了分离鉴定，并筛选出一株具有抗病促生潜力的细菌B.velezensis FQ-G3。所分离的内生菌中细菌的数量居多，说明细菌是番茄内生菌的优势菌群，这与前人研究结果类似。番茄内生菌种类丰富，包括贝莱斯芽孢杆菌(B.velezensis)、枯草芽孢杆菌(B.subtilis)、解淀粉酶芽孢杆菌(B.amyloliquefaciens)、亚隔孢壳属(Didymella)、曲霉属(Aspergillus)、枝孢霉属(Cladosporium)和链格孢属(Alternaria)等。王梅菊等[16]发现，油菜内生细菌中芽孢杆菌属(Bacillus)、假单胞菌属(Pseudomonas)和寡养单胞菌属(Stenotrophomonas)所占比例较高。对甘蔗根茎中内生菌分离鉴定，发现其内生菌株属于6个种属[17]，说明了植物内生菌多样性。
平板对峙和果实接种结果证明B.velezensis FQ-G3对灰葡萄孢表现出强烈的拮抗作用，而且FQ-G3对Cladosporium sp.、Fusarium pemambucanum及Alternaria alternata也有不同程度的抑制作用。研究表明内生芽孢杆菌属(Bacillus)菌株对很多病原菌具有抑制作用，如B.amyloliquefaciens对Fusarium oxysporum、Penicillium spp.、Colletotrichum spp.、B.cinerea和Sclerotinia sclerotiorum[18-19]。B.halotolerans能够抑制B.cinerea和F.oxysporum菌丝生长[20-21]。分离自葡萄的内生菌株Bacillus sp. K1对葡萄采后灰葡萄孢具有良好的防控效果[22]。
很多研究表明微生物通过氮素固定、ACC脱氨酶合成、激素合成(IAA、2, 3-丁二醇等)和铁载体生物合成等途径促进植物生长[7]。王红珠等[23]从植物中分离筛选出链球菌菌株GZ01，经鉴定其能够通过分泌IAA、产铁载体和溶磷等功能来促进宿主植物生长。Sukhada[24]从番石榴根际分离筛选到一株放线菌可以产生IAA和铁载体来促进植物的生长，并且能够抑制番石榴和番茄枯萎病的病原菌。Zhang等[25]从木薯根部分离出一株内生菌A02，能产生IAA，具有溶磷固氮能力，对木薯的根茎叶生长具有促进作用。从黄鸢尾叶片中获得的4株内生细菌株Pseudomonas gessardii HRT18、Brevibacterium frigoritolerans HRT8、Streptomyces atratus HRT13和Bacillus toyonensis HST13，不仅对植物病原真菌Fusarium、Rhizoctonia、Botrytis、Pythium和Alternaria表现出抗菌活性，还能产生几丁质酶、蛋白酶、葡聚糖酶、脂肪酶和IAA等物质[26]。本研究中FQ-G3能够分泌IAA、蛋白酶、嗜铁素、蛋白酶和几丁质酶等植物促生抑菌相关物质，番茄盆栽试验进一步证明了其促生效果。
二分隔培养试验表明FQ-G3菌株能够产生VOCs，并且对拟南芥和番茄幼苗的生长具有明显的促进作用。据报道很多细菌能够产生具有抗病促生活性的VOCs，主要活性物质包括2, 3-丁二醇(2, 3-butanediol)[27-28]、乙缩醛(acetoin)[27]、2-庚酮(2-heptanone)[29]和2-壬酮(2-nonnanone)[15]等。解淀粉芽孢杆菌(B.amyloliquefaciens)菌株FZB42释放的挥发性有机化合物具有抗菌活性，也能促进植物生长和增强系统抗性[30]。枯草芽孢杆菌GB03释放的VOCs主成分为乙偶姻(acetoin)和2, 3-丁二醇，能够增强拟南芥幼苗的系统抗性，并能够降低幼苗软腐病发病率[31-32]。菌株FQ-G3能够产生植物促生活性物质，但需要通过固相微萃取气相色谱-质谱法(solid-phase micro-extraction gas chromatography mass spectrometry, SPME-GC-MS)进一步鉴定该菌株主要活性成分。
内生菌在促进植物生长和生物控制剂方面具有非常大的潜力，它能够促进植物的生长和正常发育[7]，还可以有效的抑制果蔬病害[33-35]。本研究分离得到的菌株FQ-G3可以作为拮抗菌预防番茄由灰葡萄孢引起的腐烂损失，为果蔬采后绿色环保的生物防治剂和植物促生剂研发提供一种可参考的新资源。但本研究对于菌株FQ-G3产生的挥发性气体成分、促生长机制和田间促生效果还待进一步的试验研究。
基金
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  • 山西省自然科学基金(20210302123084)
  • 山西省自然科学基金(20210302123087)
文献
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参考文献 引证文献
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2024年第64卷第1期
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doi: 10.13343/j.cnki.wsxb.20230368
  • 接收时间：2023-05-27
  • 首发时间：2026-03-18
  • 出版时间：2024-01-04
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  • 收稿日期：2023-05-27
  • 录用日期：2023-09-18
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Natural Science Foundation of Shanxi Province(20210302123084)
山西省自然科学基金(20210302123084)
Natural Science Foundation of Shanxi Province(20210302123087)
山西省自然科学基金(20210302123087)
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    运城学院生命科学系, 山西 运城 044000

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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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