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Article(id=1226554100536492302, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226554095926952065, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20240818, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1734364800000, receivedDateStr=2024-12-17, revisedDate=null, revisedDateStr=null, acceptedDate=1739548800000, acceptedDateStr=2025-02-15, onlineDate=1770362885841, onlineDateStr=2026-02-06, pubDate=1751558400000, pubDateStr=2025-07-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1770362885841, onlineIssueDateStr=2026-02-06, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1770362885841, creator=13701087609, updateTime=1770362885841, updator=13701087609, issue=Issue{id=1226554095926952065, tenantId=1146029695717560320, journalId=1192105938417971205, year='2025', volume='65', issue='7', pageStart='2771', pageEnd='3233', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1770362884741, creator=13701087609, updateTime=1770363575040, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1226556991309529548, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226554095926952065, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1226556991309529549, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226554095926952065, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=2965, endPage=2975, ext={EN=ArticleExt(id=1226554102172270982, articleId=1226554100536492302, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Identification of an antifungal compound produced by Pseudomonas protegens FD6 and determination of its inhibitory effect against Botrytis cinerea, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

[Objective] To explore the antifungal activity and inhibitory mechanism and identify the main antagonistic substances of the crude extract of Pseudomonas protegens FD6 in fermentation. [Methods] The plate confrontation test was employed to screen the optimal medium, extractant, and solvent and determine the antifungal activity stability of the crude extract. The main antifungal substances in the crude extract were identified by MALDI-TOF/TOF mass spectrometry, and the antifungal spectrum of the crude extract was explored. The inhibition mechanism of the crude extract against Botrytis cinerea was investigated by mycelial morphology observation, spore germination test, PI staining, and conductivity determination. [Results] The crude extract of FD6 in the potato juice medium without any carbon source displayed the strongest antifungal activity. The methanol-solved crude extract obtained with ethyl acetate as the extractant had the strongest inhibitory activity. The main antifungal substance of the crude extract was identified as a cyclic lipopeptide, orfamide A, based on mass spectrometry data. In addition, the crude extract displayed broad-spectrum inhibitory activities against Phomopsis amygdali, Fusarium graminearum, Colletotrichum gloeosporioides, and Monilinia fructicola. The antifungal activity of the crude extract was stable after treatment at 121 ℃ for 20 min, within the range of pH 4.0-10.0, and with protease K and trypsin. The mycelial morphology of B. cinerea became abnormal after treatment with the crude extract. The treatment with 2.0 mg/mL crude extract completely inhibited the conidial germination of B. cinerea, caused a 50% damage of the cell membrane, increased the conductivity, and led to leakage of intracellular nucleic acids and proteins. [Conclusion] The metabolites produced by P. protegens FD6 have broad-spectrum and stable antifungal activities, serving as lead compounds for exploration of new green fungicides.

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【目的】 研究防御假单胞菌(Pseudomonas protegens) FD6发酵粗提物的抑菌活性及其作用机制，解析粗提物中主要的活性物质。 【方法】 通过平板对峙试验筛选发酵最佳的培养基、萃取剂、溶剂，以及发酵粗提物抑菌活性的稳定性。利用MALDI-TOF/TOF质谱鉴定粗提物中主要抗菌物质，并探究粗提物的抑菌谱。通过菌丝形态观察、孢子萌发试验、PI染色及电导率测定，探究粗提物对番茄灰霉病菌(Botrytiscinerea)的抑菌作用机制。 【结果】 平板对峙试验表明，菌株FD6以不添加碳源马铃薯汁培养基进行发酵的粗提物抑菌活性最强。以乙酸乙酯作为萃取剂，通过甲醇溶解所得的粗提物抑菌活性最强。质谱分析表明，粗提物中含有环脂肽orfamide A。进一步检测粗提物抑菌谱，发现其对桃枝枯病菌(Phomopsis amygdali)、小麦赤霉病菌(Fusarium graminearum)、梨炭疽病菌(Colletotrichum gloeosporioides)和桃褐腐病菌(Monilinia fructicola)均具有抑制作用。稳定性试验发现，粗提物在121 ℃处理20 min以及pH 4.0-10.0条件下抑菌活性稳定，且蛋白酶K和胰蛋白酶处理不影响其抑菌活性。粗提物处理B. cinerea菌丝可导致其形态异常，B. cinerea分生孢子经2.0 mg/mL粗提物处理后萌发被完全抑制，50%的细胞膜受到破坏，电导率增加，细胞内核酸与蛋白外渗。 【结论】 防御假单胞菌FD6产生的代谢物抑菌谱广且稳定性强，为发掘新型绿色农药提供先导化合物。

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作者贡献声明

杨晶龙：实验操作和论文撰写；宋琦：协助实验操作；胡婕妤：数据收集；焦永鑫：数据处理；吴涛：文章修改及论文润色；张清霞：研究构思和设计并参与论文讨论。

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防御假单胞菌FD6抗菌代谢物鉴定及其对灰霉病菌的抑制作用
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杨晶龙 , 宋琦 , 胡婕妤 , 焦永鑫 , 吴涛 , 张清霞
微生物学报 | 研究报告 2025,65(7): 2965-2975
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微生物学报 | 研究报告 2025, 65(7): 2965-2975
防御假单胞菌FD6抗菌代谢物鉴定及其对灰霉病菌的抑制作用
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杨晶龙, 宋琦, 胡婕妤, 焦永鑫, 吴涛, 张清霞
作者信息
  • 扬州大学 植物保护学院，江苏 扬州
Identification of an antifungal compound produced by Pseudomonas protegens FD6 and determination of its inhibitory effect against Botrytis cinerea
Jinglong YANG, Qi SONG, Jieyu HU, Yongxin JIAO, Tao WU, Qingxia ZHANG
Affiliations
  • College of Plant Protection, Yangzhou University, Yangzhou, Jiangsu, China
出版时间: 2025-07-04 doi: 10.13343/j.cnki.wsxb.20240818
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【目的】 研究防御假单胞菌(Pseudomonas protegens) FD6发酵粗提物的抑菌活性及其作用机制，解析粗提物中主要的活性物质。 【方法】 通过平板对峙试验筛选发酵最佳的培养基、萃取剂、溶剂，以及发酵粗提物抑菌活性的稳定性。利用MALDI-TOF/TOF质谱鉴定粗提物中主要抗菌物质，并探究粗提物的抑菌谱。通过菌丝形态观察、孢子萌发试验、PI染色及电导率测定，探究粗提物对番茄灰霉病菌(Botrytiscinerea)的抑菌作用机制。 【结果】 平板对峙试验表明，菌株FD6以不添加碳源马铃薯汁培养基进行发酵的粗提物抑菌活性最强。以乙酸乙酯作为萃取剂，通过甲醇溶解所得的粗提物抑菌活性最强。质谱分析表明，粗提物中含有环脂肽orfamide A。进一步检测粗提物抑菌谱，发现其对桃枝枯病菌(Phomopsis amygdali)、小麦赤霉病菌(Fusarium graminearum)、梨炭疽病菌(Colletotrichum gloeosporioides)和桃褐腐病菌(Monilinia fructicola)均具有抑制作用。稳定性试验发现，粗提物在121 ℃处理20 min以及pH 4.0-10.0条件下抑菌活性稳定，且蛋白酶K和胰蛋白酶处理不影响其抑菌活性。粗提物处理B. cinerea菌丝可导致其形态异常，B. cinerea分生孢子经2.0 mg/mL粗提物处理后萌发被完全抑制，50%的细胞膜受到破坏，电导率增加，细胞内核酸与蛋白外渗。 【结论】 防御假单胞菌FD6产生的代谢物抑菌谱广且稳定性强，为发掘新型绿色农药提供先导化合物。

防御假单胞菌  /  抑菌活性  /  番茄灰霉病菌  /  作用机制  /  orfamide A

[Objective] To explore the antifungal activity and inhibitory mechanism and identify the main antagonistic substances of the crude extract of Pseudomonas protegens FD6 in fermentation. [Methods] The plate confrontation test was employed to screen the optimal medium, extractant, and solvent and determine the antifungal activity stability of the crude extract. The main antifungal substances in the crude extract were identified by MALDI-TOF/TOF mass spectrometry, and the antifungal spectrum of the crude extract was explored. The inhibition mechanism of the crude extract against Botrytis cinerea was investigated by mycelial morphology observation, spore germination test, PI staining, and conductivity determination. [Results] The crude extract of FD6 in the potato juice medium without any carbon source displayed the strongest antifungal activity. The methanol-solved crude extract obtained with ethyl acetate as the extractant had the strongest inhibitory activity. The main antifungal substance of the crude extract was identified as a cyclic lipopeptide, orfamide A, based on mass spectrometry data. In addition, the crude extract displayed broad-spectrum inhibitory activities against Phomopsis amygdali, Fusarium graminearum, Colletotrichum gloeosporioides, and Monilinia fructicola. The antifungal activity of the crude extract was stable after treatment at 121 ℃ for 20 min, within the range of pH 4.0-10.0, and with protease K and trypsin. The mycelial morphology of B. cinerea became abnormal after treatment with the crude extract. The treatment with 2.0 mg/mL crude extract completely inhibited the conidial germination of B. cinerea, caused a 50% damage of the cell membrane, increased the conductivity, and led to leakage of intracellular nucleic acids and proteins. [Conclusion] The metabolites produced by P. protegens FD6 have broad-spectrum and stable antifungal activities, serving as lead compounds for exploration of new green fungicides.

Pseudomonas protegens  /  antifungal activity  /  Botrytis cinerea  /  mechanism  /  orfamide A
杨晶龙, 宋琦, 胡婕妤, 焦永鑫, 吴涛, 张清霞. 防御假单胞菌FD6抗菌代谢物鉴定及其对灰霉病菌的抑制作用. 微生物学报, 2025 , 65 (7) : 2965 -2975 . DOI: 10.13343/j.cnki.wsxb.20240818
Jinglong YANG, Qi SONG, Jieyu HU, Yongxin JIAO, Tao WU, Qingxia ZHANG. Identification of an antifungal compound produced by Pseudomonas protegens FD6 and determination of its inhibitory effect against Botrytis cinerea[J]. Acta Microbiologica Sinica, 2025 , 65 (7) : 2965 -2975 . DOI: 10.13343/j.cnki.wsxb.20240818
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由无性菌类灰葡萄孢(Botrytis cinerea)引起的灰霉病在番茄棚室栽培中发生面积逐年增加，已严重影响番茄的产量和品质[1-2]。B. cinerea主要危害果实，每年造成产量损失高达50%，严重威胁我国番茄产业的可持续发展[3-4]。B. cinerea的寄主范围广且致病性强，据报道约有1 400多种植物可被B. cinerea侵染[5]。当前灰霉病的防治主要依靠化学农药，然而化学农药的过度使用不仅影响人类健康、引发食品安全问题，还会造成环境污染，并且易导致病原菌产生抗药性。因此，寻找绿色高效的灰霉病防治方法已成为当前农业生产亟待解决的问题[6]。
假单胞菌属(Pseudomonas)在自然界中分布广泛，具有广谱的植物病原菌抑制作用。它们不仅能在植物根际定殖并促进植物生长，还能通过生态位竞争和利用植物分泌物作为自身营养物质来抑制病原物的生长[7-8]。此外，假单胞菌能产生多种天然抗菌物质，如2,4-二乙酰基间苯三酚(2,4-diacetylphloroglucinol, 2,4-DAPG)、硝吡咯菌素(pyrrolnitrin, PRN)、藤黄绿脓菌素(pyoluteorin, PLT)、氢氰酸(hydrogen cyanide, HCN)、环脂肽(cyclic lipopeptides, CLP)、吩嗪- 1-羧酸(phenazine-1-carboxylic acid, PCA)等[9]。其中，CLP是一类由细菌通过非核糖体肽合成酶合成的脂肽类化合物，主要由假单胞菌、芽孢杆菌(Bacillus)和链霉菌(Streptomyces)产生[10-11]。CLP作为一种新型表面活性剂，对部分病原菌具有较好的生防活性。宋晓雅等[12]发现，贝莱斯芽孢杆菌(B. velezensis) QZ-1产生的伊枯草菌素可以有效抑制番茄灰霉菌的扩展。Nielsen等[13]发现，荧光假单胞菌(P. fluorescens) 96.578产生的环脂肽类抗生素tensin能够显著抑制立枯丝核菌菌丝的延伸，可用于防治水稻纹枯病。P. fluorescens SS101产生的环脂肽massetolide A可以裂解腐霉属(Pythium)真菌产生的游动孢子[14]。此外，环脂肽orfamide能够诱导植物产生系统获得抗性(systemic acquired resistance, SAR)，帮助植物抵御多种病原菌的侵染[15]。目前，已有假单胞菌商品化制剂登记，如在美国环境保护署注册的Bio-Save 10 LP、Bio-Save 11 LP和BlightBan A506[16]。
防御假单胞菌(P. protegens) FD6分离自青菜根际土壤，可有效抑制番茄灰霉病菌、桃褐腐病菌(Monilinia fructicola)、小麦根腐病菌等多种病原物的生长[17-19]。前期研究表明FD6可产生多种抗菌代谢产物，如PLT、2,4-DAPG、PRN、HCN和嗜铁素等[17]。通过antiSMASH预测，FD6全基因组可能还含有orfamide类环脂肽、甲烷氧化菌素及3种未知次生代谢产物合成基因簇[20]。本研究分析鉴定P. protegens FD6产生的新型抗菌物质，明确其对B. cinerea的作用机制，旨在为利用生防菌绿色高效防控植物病害奠定基础。
1 材料与方法
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1.1 菌株
P. protegens FD6 (GenBank登录号为CP031396.1)分离自福建闽侯青口青菜根围[17]。所使用的植物病原真菌包括桃枝枯病菌(Phomopsis amygdali)、小麦赤霉病菌(Fusarium graminearum)、梨炭疽病菌(Colletotrichum gloeosporioides)、桃褐腐病菌(M. fructicola)、番茄灰霉病菌(B. cinerea)、水稻恶苗病菌(F. moniliforme)、茭白黑粉病菌(Ustilago esculenta)、小麦全蚀病菌(Gaeumannomyces graminis)和水稻纹枯病菌(Rhizoctonia solani)。小麦全蚀病菌由南京农业大学伍辉军课题组提供，其余菌株由本课题组保存。
1.2 培养基
马铃薯汁培养基(PA) (g/L)：马铃薯200.0，琼脂20.0。
马铃薯葡萄糖培养基(PDA) (g/L)：马铃薯200.0，葡萄糖20.0，琼脂20.0。
马铃薯蔗糖培养基(PSA) (g/L)：马铃薯200.0，蔗糖20.0，琼脂20.0。
1.3 不同碳源对菌株FD6抑菌活性和抗生素产量的影响
采用平板对峙法测定菌株FD6在PA、PDA和PSA上对B. cinerea和R. solani的抑菌活性，每个处理重复3次。
取100 μL FD6菌液涂布在20 mL上述3种培养基平板上，28 ℃培养5 d后切碎并添加等体积的乙酸乙酯，28 ℃、180 r/min避光振荡90 min，蒸干有机相，用1 mL甲醇溶解后经0.22 μm滤膜过滤得到粗提物。利用HPLC进行抗生素定量检测。2,4-DAPG的HPLC条件：乙腈的体积分数为55%，水的体积分数为44.9%，甲酸的体积分数为0.1%，流速为1.0 mL/min，紫外检测器波长270 nm；PLT的HPLC条件：甲醇的体积分数为70%，水的体积分数为30%，流速为1.0 mL/min，紫外检测器波长308 nm。
1.4 抗菌物质萃取剂及溶剂的筛选及其鉴定
参照1.3节粗提物提取方法，以三氯甲烷、乙酸乙酯和正丁醇3种有机溶剂分别萃取FD6产生的代谢物再用甲醇溶解；将乙酸乙酯萃取物蒸干后，分别用0.5 mL 1‰ DMSO、甲醇、ddH2O溶解获得粗提物。取100 μL不同处理的粗提物，以100 μL甲醇作为阴性对照，采用打孔法检测抑菌活性。待对照组病原真菌长满平板后，分别测量其抑菌带宽，每组设3个重复。
将上述粗提物送至中国科学院生物物理研究所蛋白质组学技术实验室进行质谱分析。将样品溶液与CHCA基质饱和溶液等体积充分混合，取0.7 μL样品与基质混合液滴加至不锈钢MALDI靶板上，并在室温下自然干燥。样品用MALDI-TOF MS (Bruker公司)进行质谱分析，激光源为200 Hz (紫外波长355 nm)的Nd:YAG激光器，加速电压为20 kV，MS采用反射正离子模式，质量范围800-4 000 Da，每个图谱采集4 000 shots。
1.5 抗菌粗提物抑菌谱及稳定性的研究
利用1.3节方法制备的粗提物，采用打孔法检测对供试8种植物病原真菌的抑菌活性。
以B. cinerea为指示菌，测定粗提物对温度、pH和蛋白酶的敏感性，每组处理设3次重复。设置粗提物在20、40、60、80、100和121 ℃条件下孵育20 min，室温未处理为空白对照；将粗提物加入0.2 mol/L Na2HPO4- 0.1 mol/L柠檬酸缓冲液(pH 4.0-10.0)中，置于37 ℃孵育2 h，pH 7.0为空白对照，测定其抑菌活性；将蛋白酶K和胰蛋白酶的浓度定为2 mg/mL，分别以1:1的比例与粗提物混合，37 ℃孵育2 h后测定抑菌活性，以未添加蛋白酶为空白对照。
1.6 抗菌粗提物对 B. cinerea 抑菌机理的研究
1.6.1 抗菌粗提物对 B. cinerea 菌丝形态的影响
分别挑取未经粗提物处理和经粗提物处理后的B. cinerea菌丝体，置于已滴加无菌水的载玻片上，使菌丝体展开后，通过显微镜观察菌丝体的形态。
1.6.2 抗菌粗提物对 B. cinerea 分生孢子萌发和芽管伸长的影响
在PDB培养基中分别添加浓度为0、0.3、0.6、1.0、1.5和2.0 mg/mL的粗提物，再加入孢子悬浮液，使最终孢子浓度为105个/mL。取20 μL滴于凹玻片上，在25 ℃条件下培养至对照组孢子完全萌发，测定处理组孢子萌发率和芽管伸长情况[21]。当芽管长度等于或大于孢子直径时，表明孢子已萌发。使用显微镜测微计测定芽管长度。每组随机观察约200个孢子，试验重复3次。
1.6.3 抗菌粗提物对 B. cinerea 分生孢子细胞膜完整性的影响
参考Jeong等[22]的方法，采用碘化丙啶(propidium iodide, PI)染色法检测孢子细胞膜是否受到破坏。取B. cinerea分生孢子悬浮液240 μL (105个/mL)，加入到240 μL PDB中，再加入20 μL不同浓度的粗提物，在25 ℃、180 r/min振荡培养4 h。8 000 r/min离心5 min后去掉上清液，加入1 mL PI染色液(终浓度10 μg/mL)室温染色30 min，用PBS缓冲液洗涤3次后定容至240 μL。在荧光显微镜下随机观察约100个孢子，显示红色荧光的孢子即为细胞膜受损。设置空白对照组，每组处理3次重复。
1.6.4 抗菌粗提物对 B. cinerea 分生孢子细胞质渗漏的影响
参考Tao等[23]方法测定细胞渗漏。取B. cinerea分生孢子悬浮液100 μL (105个/mL)，加入到20 mL PDB中，置于25 ℃、180 r/min条件下振荡培养3 d。4 000 r/min离心30 min，用无菌蒸馏水洗涤3次后定容至20 mL。将粗提物分别以0、0.3、1.0和2.0 mg/mL的浓度加入上述4 mL孢子悬浮液中。测定处理后2 h内细胞外电导率、OD260以及OD280。设置空白对照组，每组处理3次重复。
1.7 统计分析
所有试验处理均设3个重复。数据显著性分析采用方差分析(IBM SPSS Statistics 25，ANOVA)进行，使用GraphPad Prism 8.4.3软件对平均值进行Tukey检验(P<0.05)并绘图。
2 结果与分析
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2.1 不同碳源对菌株FD6抑菌能力及抗生素产量的影响
在PA培养基中，FD6对B. cinerea和R. solani的抑菌活性最强，抑菌带宽分别为1.4 cm和1.3 cm；其次为PSA培养基，抑菌带宽分别为0.6 cm和0.5 cm；而在PDA培养基中，FD6的抑菌活性最弱(图1A、1B)。FD6主要产生抗生素2,4-DAPG和PLT。通过HPLC定量分析FD6在不同培养基上2,4-DAPG和PLT的产生情况，结果显示FD6在PDA和PSA培养基上均可产生较高水平的2,4-DAPG，但在PA培养基上，2,4-DAPG的产量显著低于PSA和PDA (图1C)。在这3种培养基中均未检测到PLT (数据未显示)。然而，在PA培养基中，FD6的抑菌活性最强，推测FD6在PA培养基中可能产生其他新型抗菌物质。
2.2 抗菌粗提物提取条件优化及活性物质鉴定
使用不同萃取剂提取FD6抑菌粗提物后检测其对B. cinerea的平板抑制效果。结果显示，3种有机溶剂均能萃取到抗菌物质，其中三氯甲烷提取物的抑菌活性最强，抑菌带宽为1.57 cm，其次是乙酸乙酯和正丁醇(图2A)。由于三氯甲烷属于管控化学品，因此后续研究选择乙酸乙酯作为萃取剂。经乙酸乙酯萃取的粗提物旋蒸后，使用不同溶剂溶解粗提物并检测其对B. cinerea的平板抑制效果。结果显示，甲醇溶解的粗提物对B. cinerea的抑制效果最显著，其次为DMSO，而无菌水处理组未表现出抑菌效果(图2B)。因此，后续研究选定甲醇溶解FD6粗提物。对上述所得粗提物进行MALDI-TOF MS检测，结果显示粗提物中可检测到多个质谱峰，主峰为1 317.906 m/z ([M+Na]+)，特征峰为1 295.914 m/z ([M+H]+)和1 333.886 m/z ([M+K]+)，结合orfamide A的分子量大小可以确定抗菌粗提物中含有脂肽类化合物orfamide A (图2C)。
2.3 粗提物抑菌谱测定及稳定性研究
粗提物对桃枝枯病菌和小麦赤霉病菌的抑菌能力最强，抑菌带宽分别达2.68 cm和2.36 cm；其次为梨炭疽病菌、桃褐腐病菌、水稻恶苗病菌和番茄灰霉病菌；对茭白黑腐病菌和小麦全蚀病菌的抑菌作用较弱(图3)。稳定性分析结果表明，pH在4.0-10.0 (图4A)、121 ℃高温处理(图4B)、胰蛋白酶及蛋白酶K处理(图4C)对粗提物的抑菌活性均无显著影响。
2.4 粗提物对灰霉病菌抑菌机理的研究
2.4.1 粗提物对菌丝形态的影响
与对照组(图5A)相比，FD6粗提物处理B. cinerea菌丝体后，菌丝体和细胞质均发生明显变化。处理组的菌丝体出现无规则分枝(图5B)、细胞内原生质体皱缩(图5C)及菌丝体顶端膨大(图5D)。
2.4.2 粗提物对孢子萌发和芽管伸长的影响
结果显示，FD6粗提物处理B. cinerea分生孢子4 h后，对分生孢子的萌发具有明显的抑制作用。使用0.6 mg/mL粗提物处理B. cinerea分生孢子时，能显著抑制孢子的萌发和芽管伸长；而使用2.0 mg/mL粗提物处理时，完全抑制了B. cinerea分生孢子的萌发和芽管伸长(图6)。
2.4.3 粗提物对孢子细胞膜完整性的影响
PI染色结果显示，当用1.0 mg/mL粗提物处理B. cinerea分生孢子时，约33.34%的分生孢子细胞膜完整性被破坏；而当用2.0 mg/mL粗提物处理时，50%左右的B. cinerea分生孢子可以被染色(图7)。这表明FD6粗提物能够破坏B. cinerea分生孢子的细胞膜完整性。
2.4.4 粗提物对孢子细胞质渗漏的影响
如图8A所示，1.0 mg/mL和2.0 mg/mL FD6粗提物处理B. cinerea分生孢子30 min后，细胞电导率显著升高；处理90 min后，电导率开始下降。细胞外核酸和蛋白质检测结果显示，粗提物处理后，胞外核酸和蛋白质逐渐增多(图8B、8C)。这表明FD6粗提物能够破坏B. cinerea分生孢子的细胞膜完整性，导致细胞膜通透性增强，从而促使胞内离子、核酸和蛋白质渗漏。
3 讨论与结论
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分泌抗菌物质是生防细菌防治植物病害的主要方式之一。抗生素的产生通常受到环境因子的影响，其中碳源对生防菌抗生素的产生影响较大。例如，葡萄糖可以促进P. fluorescens CHA0中2,4-DAPG的产生，但抑制PLT的合成[24]。本研究发现，使用蔗糖作为碳源更有利于P. protegens FD6中2,4-DAPG的产生。然而，在不添加任何碳源的PA培养基中，P. protegens FD6产生的代谢物对B. cinerea和R. solani的拮抗能力最强。前期报道显示，PLT和2,4-DAPG是P. protegens FD6产生的主要活性抑菌物质[20]。然而本研究发现，在PA培养基中P. protegens FD6不产生2,4-DAPG和PLT，说明P. protegens FD6在碳源匮乏的PA培养基中可 能产生其他抗菌物质。James等[25]报道P. fluorescens HV37a在PDA培养基中代谢物的抑菌活性要高于PA培养基，说明以葡萄糖作为培养基碳源对不同假单胞菌的抗生素产生具有差异性。
环脂肽类通常由8-25个氨基酸的环状寡肽链和一个脂肪酸链组成，是具有双亲功能的大分子化合物。假单胞菌产生的环脂肽根据寡肽链的长度和特点可以分为14个组，其中包括orfamide[26]。截至目前，从不同细菌中已经鉴定到8种orfamide同源物(A-H)，这些orfamides在肽序列中第4位氨基酸(Ile或Val)的组成上或脂肪酸部分的长度(或饱和度)上有差异[27]。通常分离自植物根际的假单胞菌可产生环脂肽，且P. protegens主要分泌orfamide A类环脂肽[28]。本研究通过质谱检测证实，分离自油菜根际的P. protegens FD6在PA培养基中也能产生orfamide A。
P. protegens FD6粗提物具有广谱的病原真菌抑制作用。0.6 mg/mL粗提物能显著抑制B. cinerea分生孢子的萌发和芽管伸长，而2.0 mg/mL粗提物则可完全抑制B. cinerea分生孢子的萌发。Geudens等[26]报道指出，细胞膜的扰动，尤其是膜孔的形成造成细胞内容物外泄，是抗菌物质的主要抑菌机制。本研究发现，FD6粗提物处理B. cinerea分生孢子可促使其膜内物质泄漏，导致细胞外电导率显著增强。通过PI染色后在荧光显微镜下观察，发现FD6粗提物处理B. cinerea分生孢子可破坏其细胞膜，且2.0 mg/mL粗提物处理可导致接近50%的分生孢子细胞膜受到破坏。此外，粗提物处理菌丝后，使菌丝原生质体皱缩、菌丝顶端出现膨大以及无规则分叉。Ma等[28]报道，环脂肽orfamide A-C这3种化合物均可抑制R. solani菌丝生长，导致菌丝产生更多的分枝。假单胞菌产生的环脂肽粗提物也可以抑制稻瘟病菌(Magnaporthe oryzae)分生孢子萌发及附着胞的形成[27]。因此，P. protegens FD6可能通过合成环脂肽类物质抑制植物病原真菌的孢子萌发并破坏细胞膜。
抗菌肽是微生物产生的具有广谱抑菌作用的天然产物，可以影响微生物的细胞膜结构，具有快速、高效和持久的抑菌活性且不易产生耐药性等优点，在病害防治方面具有潜在的研究价值[29]。已有相关抗菌肽的报道，例如抗菌肽O3TR和C12O3TR在高温和强酸强碱条件下仍能保持对甜菜根腐病菌(F. culmorum)的抑菌活性[30]；抗菌肽GNU6和GNU7在胰蛋白酶、胰凝乳蛋白酶和金属蛋白酶处理后仍有抑菌活性[31]。本研究发现，包含环脂肽的P. protegens FD6粗提物具有热稳定性和耐酸碱性，并且对蛋白酶K和胰蛋白酶不敏感，说明粗提物中的环脂肽具有较好的稳定性。后续试验将进一步优化P. protegens FD6发酵条件，以提高环脂肽类抗生素的产量，从而提高P. protegens FD6的防病效果，为生防菌剂的开发奠定基础。
作者利益冲突公开声明
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作者声明不存在任何可能会影响本文所报告工作的已知经济利益或个人关系。
基金
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  • 国家自然科学基金(32072471)
  • 国家自然科学基金(32472628)
  • 扬州市重点研发项目(YZ2023056)
  • 扬州大学自制实验仪器设备项目(zzyq2024dy03)
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2025年第65卷第7期
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doi: 10.13343/j.cnki.wsxb.20240818
  • 接收时间：2024-12-17
  • 首发时间：2026-02-06
  • 出版时间：2025-07-04
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  • 收稿日期：2024-12-17
  • 录用日期：2025-02-15
基金
National Natural Science Foundation of China(32072471)
国家自然科学基金(32072471)
National Natural Science Foundation of China(32472628)
国家自然科学基金(32472628)
Yangzhou Key Research and Development Program(YZ2023056)
扬州市重点研发项目(YZ2023056)
Self-made Experimental Instruments and Equipment Project of Yangzhou University(zzyq2024dy03)
扬州大学自制实验仪器设备项目(zzyq2024dy03)
作者信息
    扬州大学 植物保护学院，江苏 扬州
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https://castjournals.cast.org.cn/joweb/wswxb/CN/10.13343/j.cnki.wsxb.20240818
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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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