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Article(id=1226460582279299613, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226460576751206672, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20240715, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1731340800000, receivedDateStr=2024-11-12, revisedDate=null, revisedDateStr=null, acceptedDate=1745942400000, acceptedDateStr=2025-04-30, onlineDate=1770340589350, onlineDateStr=2026-02-06, pubDate=1754236800000, pubDateStr=2025-08-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1770340589350, onlineIssueDateStr=2026-02-06, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1770340589350, creator=13701087609, updateTime=1770340589350, updator=13701087609, issue=Issue{id=1226460576751206672, tenantId=1146029695717560320, journalId=1192105938417971205, year='2025', volume='65', issue='8', pageStart='1', pageEnd='3812', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1770340588033, creator=13701087609, updateTime=1770363610188, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1226557138735117113, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226460576751206672, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1226557138735117114, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226460576751206672, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=3540, endPage=3551, ext={EN=ArticleExt(id=1226460582694535752, articleId=1226460582279299613, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Mechanisms of bacterial infection mediated by the lipoprotein PplA of Listeria monocytogenes, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

[Objective] We employed the wild-type strain of Listeria monocytogenes, the lipoprotein gene pplA- deleted strain, and the complementary strain to investigate the role of PplA in the infection of L. monocytogenes. [Methods] We compared the hemolytic capacity, cell adhesion and invasion, intracellular proliferation, cell-to-cell migration, mouse organ colonization, transcription levels of virulence factors in mouse organs, and transcription levels of quorum sensing-related genes among wild-type, pplA-deleted, and complementary strains to explore the role of PplA in the infection of L. monocytogenes. [Results] After the deletion of pplA, L. monocytogenes showed no significant change in intracellular proliferation or cell-to-cell migration. However, its hemolytic capacity, cell adhesion and invasion, mouse organ colonization, and transcription levels of virulence factors such as plcB, hly, and prfA in mouse organs were significantly reduced. Moreover, the transcription levels of quorum sensing-related genes agrA, agrB, agrC, and luxS were altered in the pplA-deleted strain. [Conclusion] Thelipoprotein PplA is involved in the virulence regulation and affects the pathogenicity of L. monocytogenes.

, correspAuthors=Changyong CHENG, Mianmian CHEN, authorNote=null, correspAuthorsNote=
*E-mail: CHEN Mianmian,
CHENG changyong,
, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, authorCompany=null, fund=null, authors=

These authors contributed equally to this work.

, authorsList=Yucong LIU, Lifeng YANG, Gengjia REN, Qinjia SANG, Houhui SONG, Jing SUN, Lingli JIANG, Changyong CHENG, Mianmian CHEN), CN=ArticleExt(id=1226460585055929114, articleId=1226460582279299613, tenantId=1146029695717560320, journalId=1192105938417971205, language=CN, title=单核增生李斯特氏菌信息素脂蛋白PplA介导的细菌感染机制, columnId=1192149544164012138, journalTitle=微生物学报, columnName=研究报告, runingTitle=null, highlight=null, articleAbstract=

【目的】 利用单核增生李斯特氏菌野生株、脂蛋白基因pplA缺失株和回补株探究PplA在单核增生李斯特氏菌感染功能中的作用。 【方法】 比较野生株、缺失株和回补株在细胞溶血能力、细胞黏附和侵袭能力、胞内增殖能力、胞间迁移能力、小鼠脏器定殖能力、小鼠脏器内细菌毒力因子转录水平和群体感应相关基因转录水平等方面的差异，从而解析PplA在单核增生李斯特氏菌感染宿主中的作用。 【结果】 缺失pplA基因后，单核增生李斯特氏菌的胞内增殖能力和胞间迁移能力无显著差异，但溶血能力、细胞黏附和侵袭能力、小鼠脏器定殖能力均显著降低，且小鼠脏器内细菌毒力因子plcB、hly和prfA的转录水平显著下降。此外，群体感应相关基因agrA、agrB、agrC和luxS的转录水平也发生了显著变化。 【结论】 研究表明单核增生李斯特氏菌脂蛋白PplA参与细菌毒力调控过程，显著影响细菌的致病力。

, correspAuthors=程昌勇, 陈绵绵, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=x6sNuDlyoHYWEPSy8nE3kQ==, magXml=FcWhe3z2fA7KV6w0K3aBag==, pdfUrl=null, pdf=9EwHoEReQ0BxRsmjII4Mqw==, pdfFileSize=6181518, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=zeJaI1BqhPz/XkKCvILJzw==, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=s5rJ3AhJsfI3aqvZJXm2YQ==, mapNumber=null, authorCompany=null, fund=null, authors=

作者贡献声明

刘郁匆：实验操作、数据分析和论文撰写；杨立锋：数据分析、论文撰写和获取基金；任耿佳：实验操作和数据收集；桑琴嘉：数据分析和验证；宋厚辉：获取基金，数据监管；孙静：获取基金，提供技术支持；江玲丽：提出概念，提供技术支持；程昌勇：研究构思和设计，获取基金，论文修改；陈绵绵：研究构思和设计，数据收集和处理，论文撰写与修改。

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Infection and Immunity, 2003, 71(8): 4463-4471., articleTitle=Identification of the agr locus of Listeria monocytogenes: role in bacterial virulence, refAbstract=null), Reference(id=1226596310212588042, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460582279299613, doi=null, pmid=null, pmcid=null, year=2006, volume=72, issue=8, pageStart=5653, pageEnd=5658, url=null, language=null, rfNumber=[25], rfOrder=24, authorNames=SELA S, FRANK S, BELAUSOV E, PINTO R, journalName=Applied and Environmental Microbiology, refType=null, unstructuredReference=SELA S, FRANK S, BELAUSOV E, PINTO R. A mutation in the luxS gene influences Listeria monocytogenes biofilm formation[J]. 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Primer sequences used in this study

, figureFileSmall=null, figureFileBig=null, tableContent=
Primers namePrimer sequences (5′→3′)
plcA-fCGAGCAAAACAGCAACGATAG
plcA-rCGTGTCAGTTCTGGGAGTAGTGTAA
plcB-fATCATACCCTCCAGGCTACCA
plcB-rCGCCCTTTTCGCATTTTC
hly-fTCACATCGTCCATCTATTTGCC
hly-rATTACCGTTCTCCACCATTCC
inlA-fTAGCGATGGCGGTAGTTACACAGA
inlA-rTTAAGTGGCTGCGTCACGGTTC
inlB-fGCAGTCAACTTAACCCGCTATGTCA
inlB-rGTGCGAGGCTAGAGTTCCTTGTT
actA-fCAGCAGATGAGTCTTCACCACA
actA-rCCATTTCCCCGCATCTTTTA
prfA-fACAGCTGAGCTATGTGCGAT
prfA-rAATGATTTTTCGATTAACGGGAAGCT
agrA-fGTTTGAGTTAGTATCACGAATGCC
agrA-rCAAACTTCCGAATTTCCTGAGC
agrB-fGAAAGAATGGCGGATGTGTTG
agrB-rCCTGTTACTAAGGCGATACCG
agrC-fACAAAAGGAGAAGGTCGTGG
agrC-rCGGTCACTTTCGTATCTAAGGC
agrD-fACAATCCATGAAAGTTGCGGA
agrD-rTCACAAATGGACTTTTTGGTTCGT
luxS-fCAATGGGATGTCAAACAGGC
luxS-rCAACCGCATTGGACTTCATTG
rpoB-fCTACACTTAGGTATGGCTGCTCG
rpoB-rGGCTTCTTCCACTGTGCTCC
), ArticleFig(id=1226596304965513507, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460582279299613, language=CN, label=表1, caption=

本研究所用引物

, figureFileSmall=null, figureFileBig=null, tableContent=
Primers namePrimer sequences (5′→3′)
plcA-fCGAGCAAAACAGCAACGATAG
plcA-rCGTGTCAGTTCTGGGAGTAGTGTAA
plcB-fATCATACCCTCCAGGCTACCA
plcB-rCGCCCTTTTCGCATTTTC
hly-fTCACATCGTCCATCTATTTGCC
hly-rATTACCGTTCTCCACCATTCC
inlA-fTAGCGATGGCGGTAGTTACACAGA
inlA-rTTAAGTGGCTGCGTCACGGTTC
inlB-fGCAGTCAACTTAACCCGCTATGTCA
inlB-rGTGCGAGGCTAGAGTTCCTTGTT
actA-fCAGCAGATGAGTCTTCACCACA
actA-rCCATTTCCCCGCATCTTTTA
prfA-fACAGCTGAGCTATGTGCGAT
prfA-rAATGATTTTTCGATTAACGGGAAGCT
agrA-fGTTTGAGTTAGTATCACGAATGCC
agrA-rCAAACTTCCGAATTTCCTGAGC
agrB-fGAAAGAATGGCGGATGTGTTG
agrB-rCCTGTTACTAAGGCGATACCG
agrC-fACAAAAGGAGAAGGTCGTGG
agrC-rCGGTCACTTTCGTATCTAAGGC
agrD-fACAATCCATGAAAGTTGCGGA
agrD-rTCACAAATGGACTTTTTGGTTCGT
luxS-fCAATGGGATGTCAAACAGGC
luxS-rCAACCGCATTGGACTTCATTG
rpoB-fCTACACTTAGGTATGGCTGCTCG
rpoB-rGGCTTCTTCCACTGTGCTCC
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单核增生李斯特氏菌信息素脂蛋白PplA介导的细菌感染机制
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刘郁匆 1 , 杨立锋 2 , 任耿佳 1 , 桑琴嘉 1 , 宋厚辉 1 , 孙静 1 , 江玲丽 2 , 程昌勇 1, * , 陈绵绵 1, *
微生物学报 | 研究报告 2025,65(8): 3540-3551
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微生物学报 | 研究报告 2025, 65(8): 3540-3551
单核增生李斯特氏菌信息素脂蛋白PplA介导的细菌感染机制
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刘郁匆1, 杨立锋2, 任耿佳1, 桑琴嘉1, 宋厚辉1, 孙静1, 江玲丽2, 程昌勇1, * , 陈绵绵1, *
作者信息
  • 1.浙江农林大学 动物医学院，浙江省畜禽绿色生态健康养殖应用技术研究重点实验室，动物健康互联网检测技术浙江省工程研究中心，浙江省动物医学与健康管理国际科技合作基地，同一健康和食品安全“一带一路”国际联合实验室，中澳动物健康大数据分析联合实验室，浙江 杭州
  • 2.宁波卫生职业技术学院，浙江 宁波
Mechanisms of bacterial infection mediated by the lipoprotein PplA of Listeria monocytogenes
Yucong LIU1, Lifeng YANG2, Gengjia REN1, Qinjia SANG1, Houhui SONG1, Jing SUN1, Lingli JIANG2, Changyong CHENG1, * , Mianmian CHEN1, *
Affiliations
  • 1.Key Laboratory of Applied Biotechnology on Animal Science & Veterinary Medicine of Zhejiang Province, Zhejiang Engineering Research Center for Veterinary Diagnostics & Advanced Technology, Zhejiang International Science and Technology Cooperation Base for Veterinary Medicine and Health Management, the Belt and Road International Joint Laboratory for One Health and Food Safety, China-Australia Joint Laboratory for Animal Health Big Data Analytics, College of Veterinary Medicine, Zhejiang A&F University, Hangzhou, Zhejiang, China
  • 2.Ningbo College of Health Sciences, Ningbo, Zhejiang, China
出版时间: 2025-08-04 doi: 10.13343/j.cnki.wsxb.20240715
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摘要
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【目的】 利用单核增生李斯特氏菌野生株、脂蛋白基因pplA缺失株和回补株探究PplA在单核增生李斯特氏菌感染功能中的作用。 【方法】 比较野生株、缺失株和回补株在细胞溶血能力、细胞黏附和侵袭能力、胞内增殖能力、胞间迁移能力、小鼠脏器定殖能力、小鼠脏器内细菌毒力因子转录水平和群体感应相关基因转录水平等方面的差异，从而解析PplA在单核增生李斯特氏菌感染宿主中的作用。 【结果】 缺失pplA基因后，单核增生李斯特氏菌的胞内增殖能力和胞间迁移能力无显著差异，但溶血能力、细胞黏附和侵袭能力、小鼠脏器定殖能力均显著降低，且小鼠脏器内细菌毒力因子plcB、hly和prfA的转录水平显著下降。此外，群体感应相关基因agrA、agrB、agrC和luxS的转录水平也发生了显著变化。 【结论】 研究表明单核增生李斯特氏菌脂蛋白PplA参与细菌毒力调控过程，显著影响细菌的致病力。

单核增生李斯特氏菌  /  脂蛋白PplA  /  细胞感染  /  毒力调控

[Objective] We employed the wild-type strain of Listeria monocytogenes, the lipoprotein gene pplA- deleted strain, and the complementary strain to investigate the role of PplA in the infection of L. monocytogenes. [Methods] We compared the hemolytic capacity, cell adhesion and invasion, intracellular proliferation, cell-to-cell migration, mouse organ colonization, transcription levels of virulence factors in mouse organs, and transcription levels of quorum sensing-related genes among wild-type, pplA-deleted, and complementary strains to explore the role of PplA in the infection of L. monocytogenes. [Results] After the deletion of pplA, L. monocytogenes showed no significant change in intracellular proliferation or cell-to-cell migration. However, its hemolytic capacity, cell adhesion and invasion, mouse organ colonization, and transcription levels of virulence factors such as plcB, hly, and prfA in mouse organs were significantly reduced. Moreover, the transcription levels of quorum sensing-related genes agrA, agrB, agrC, and luxS were altered in the pplA-deleted strain. [Conclusion] Thelipoprotein PplA is involved in the virulence regulation and affects the pathogenicity of L. monocytogenes.

Listeria monocytogenes  /  lipoprotein PplA  /  cell infection  /  virulence regulation
刘郁匆, 杨立锋, 任耿佳, 桑琴嘉, 宋厚辉, 孙静, 江玲丽, 程昌勇, 陈绵绵. 单核增生李斯特氏菌信息素脂蛋白PplA介导的细菌感染机制. 微生物学报, 2025 , 65 (8) : 3540 -3551 . DOI: 10.13343/j.cnki.wsxb.20240715
Yucong LIU, Lifeng YANG, Gengjia REN, Qinjia SANG, Houhui SONG, Jing SUN, Lingli JIANG, Changyong CHENG, Mianmian CHEN. Mechanisms of bacterial infection mediated by the lipoprotein PplA of Listeria monocytogenes[J]. Acta Microbiologica Sinica, 2025 , 65 (8) : 3540 -3551 . DOI: 10.13343/j.cnki.wsxb.20240715
正文
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单核增生李斯特氏菌(Listeria monocytogenes)，是一种食源性人畜共患病原菌[1]。作为一种胞内寄生菌[2]，单核增生李斯特氏菌生存能力极强，不仅可以从水和土壤中分离得到[3]，还特别容易适应人体胃肠道环境，并耐受酸性、胆汁、渗透压和抗菌肽等变化[4]。该菌常在食物的储存和运输环节污染食品[5]，抵抗力较弱的人群感染后很容易患上李斯特菌病[6]，严重者还会并发败血症、胎儿流产和脑膜炎等疾病，因此被称为第三大最有害的人畜共患病原体[7-8]。目前，单核增生李斯特氏菌在欧洲的感染人数逐年增加，且具有较高的发病率和致死率[9]。
单核增生李斯特氏菌主要通过4个阶段主动入侵宿主细胞并在细胞内定殖[10]：第1阶段是黏附与侵袭，单核增生李斯特氏菌首先黏附并内化至细胞内；第2阶段是在其分泌的毒力因子帮助下进入细胞质；第3阶段是胞内增殖，单核增生李斯特氏菌在宿主细胞胞质中大量增殖；第4阶段是胞间迁移，单核增生李斯特氏菌通过形成彗星状尾巴迁移至邻近细胞[11]。单核增生李斯特氏菌在细胞内的生长和转移主要依赖于转录激活因子positive regulatory factor A (PrfA)的调节，在细菌进入宿主细胞后，PrfA能够激活和诱导单核增生李斯特氏菌的毒力基因表达，而peptide pheromone-encoding lipoprotein A (PplA)是一种能够激活PrfA的小分子辅助因子[12]。PplA是一种信息素脂蛋白，其N端存在信号肽，可在蛋白酶Eep和信号肽酶SPII的切割作用下产生多肽信息素peptide pheromone encoded by peptide pheromone-encoding lipoprotein A (pPplA)，从而刺激有助于液泡逃逸的相关基因表达，帮助单核增生李斯特氏菌从宿主吞噬泡中逃逸，并促进PrfA的激活[13]。在革兰阳性菌中，多肽信息素能够直接参与群体感应，改变微生物相互作用的细胞间通讯模式，调节生物膜的形成，影响细胞外酶、代谢物和细胞外多糖等公共物质的分泌以及抗菌物质的合成，从而直接或间接影响微生物群落对不断变化环境的适应性[14]。然而，目前关于PplA如何调控单核增生李斯特氏菌感染机制的研究尚不明确。
本研究利用单核增生李斯特氏菌野生株EGD-e、缺失株ΔpplA及其回补株，探究PplA对单核增生李斯特氏菌溶血能力、细胞黏附和侵袭能力、胞内增殖和胞间迁移能力、小鼠脏器定殖能力以及小鼠脏器内细菌毒力因子转录水平的影响，旨在揭示PplA在单核增生李斯特氏菌感染功能中的作用。
1 材料与方法
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1.1 材料
1.1.1 菌株、质粒及培养条件
本研究所用菌株与细胞均来自本实验室。菌株均由本实验室前期构建，包括单核增生李斯特氏菌野生型菌株EGD-e、脂蛋白PplA缺失株(利用pKSV7重组质粒构建，缺失pplA基因900 bp总长的第13-882 bp)和回补株[携带有pplA基因及其启动子片段的pIMK2重组整合型质粒(卡那霉素抗性)整合至pplA缺失株基因组]；细胞包括Caco-2、RAW264.7和L929细胞。EGD-e和pplA基因缺失株接种于无抗性脑心浸液(brain heart infusion, BHI)培养基中，pplA基因回补株接种于含有卡那霉素抗性的BHI培养基中，37 ℃、250 r/min振荡培养，人源结肠癌上皮细胞Caco-2使用RPMI 1640培养基(含15% FBS)培养，鼠源巨噬细胞RAW264.7和鼠源成纤维细胞系L929使用DMEM培养基(含10% FBS)培养[15]，以上细胞均于5% CO2的细胞培养箱中37 ℃恒温静置培养。
1.1.2 主要试剂和仪器
BHI培养基购自Oxoid公司；无菌脱纤绵羊血购自南京森贝伽生物科技公司；DMEM细胞培养基购自Sigma-Aldrich公司；RPMI 1640细胞培养基和胎牛血清(fetal bovine serum, FBS)均购自赛默飞世尔科技公司；二硫苏糖醇(dithiothreitol, DTT)和聚乙二醇辛基苯基醚(Triton X-100)均购自北京索莱宝科技有限公司；低熔点琼脂糖购自西格玛奥德里奇(上海)贸易有限公司；反转录试剂盒购自南京诺唯赞生物科技股份有限公司。
荧光定量PCR仪购自Bio-Rad Laboratories公司；酶标仪购自伯腾(BioTek)仪器有限公司；扫描仪购自爱普生(Epson)有限公司。
1.2 细胞的复苏与培养
将冻存的细胞于37 ℃恒温水浴锅中加热至融化，700 r/min离心3 min后弃上清，用5 mL细胞培养基重悬细胞并将其转移至T25细胞培养瓶中，放置于含5% CO2的细胞培养箱中37 ℃恒温静置培养。细胞贴壁生长至铺满细胞瓶后，弃掉培养基，用PBS清洗1次，再加入2 mL EDTA放置于培养箱中静置，待细胞全部脱落后加入3 mL细胞培养基，700 r/min离心3 min弃上清，加入适量细胞培养基重悬并计数，根据细胞数量转移至T75细胞培养瓶继续培养。
1.3 血细胞溶血实验
将平板划线培养的EGD-e、ΔpplA和CΔpplA的单菌落分别接种于相应抗性的BHI液体培养基中，37 ℃、200 r/min振荡培养过夜，然后取适量菌液于4 ℃、4 800 r/min离心10 min后用无菌PBS清洗2次并将OD600调至0.6，按照1:100接种于BHI中37 ℃、200 r/min培养至OD600为0.6时，将菌液12 000 r/min离心2 min，吸取上清液按照1:1的比例加入PBS-DTT形成混合液并在96孔板中梯度稀释。在阴性对照孔中加入100 μL PBS-DTT混合液；阳性对照孔在阴性对照孔的基础上每孔添加2 μL Triton X-100。取出1 mL新鲜绵羊血置于2 mL离心管中，4 ℃、1 000 r/min离心10 min，将上清和白细胞层弃掉；加入生理盐水重悬红细胞，再次离心，弃上清，重复洗涤步骤1次，将红细胞与生理盐水以1:20混匀，制成5%绵羊血红细胞-生理盐水悬液。每孔加入100 μL上述5%红细胞悬液，37 ℃静置培养30 min以裂解红细胞。96孔板4 ℃、1 000 r/min离心10 min后使用扫描仪扫描并检测OD550，绘制溶解曲线并使用GraphPad软件对数据进行分析，统计结果用one-way ANOVA比较差异(***：P<0.001；ns：P>0.05)。
1.4 人源上皮细胞Caco-2的黏附与侵袭实验
使用RPMI 1640 (含15% FBS)培养Caco-2细胞并计数，按照每孔约3×105个细胞转移至12孔板中，放入37 ℃、5% CO2细胞培养箱静置培养；将平板划线培养的EGD-e、ΔpplA和CΔpplA单菌落分别接种于相应抗性的BHI培养基中，37 ℃、200 r/min振荡培养过夜，然后使用10 mmol/L PBS将菌液清洗2遍后调整OD600值至0.6；将菌液以感染复数(multiplicity of infection, MOI)为10的比例感染细胞，30 min后弃去原有培养基，使用10 mmol/L PBS将细胞清洗2遍，加入1 mL细胞裂解液并进行稀释点板，统计数目并根据公式(1)计算黏附率。
黏附率=黏附菌量/感染菌量×100%
另取一块细胞培养板在感染60 min后，用PBS洗涤2次后加入1 mL预热的RPMI 1640培养基(添加庆大霉素，终浓度50 μg/mL)，放入培养箱中继续培养1 h，弃去培养孔中废液，使用PBS反复洗涤细胞后加入1 mL细胞裂解液进行稀释点板，统计数目并根据公式(2)计算侵袭率，使用GraphPad进行绘图并使用t检验分析差异(*：P<0.05；**：P<0.01)。
侵袭率=侵袭菌量/感染菌量×100%
1.5 鼠源巨噬细胞RAW264.7胞内细菌增殖试验
使用DMEM培养基(含10% FBS)培养并重悬细胞，按照每孔2×105 cell/mL的密度铺板于12孔板。将EGD-e、ΔpplA及CΔpplA分别接种于BHI液体培养基中，37 ℃、200 r/min振荡培养过夜，并将菌液OD600调整至0.6。将菌液按1:1 000比例加入DMEM培养基中，感染1 h后弃去上清，并加入DMEM培养基(含庆大霉素，终浓度50 μg/mL)；分别于感染后2、5、8 h取出一块细胞培养板，清洗后加入1 mL 4 ℃细胞裂解液使细胞破裂，随后进行稀释点板计数。使用GraphPad软件分析数据，并通过t检验比较差异(ns：P>0.05)。
1.6 鼠源成纤维细胞L929内细菌胞间迁移实验
使用DMEM培养基(含10% FBS)培养并重悬细胞，按照每孔2×105 cell/mL的密度铺板于12孔板。将EGD-e、ΔpplA及CΔpplA分别接种于BHI液体培养基中，37 ℃、200 r/min振荡培养过夜，并将菌液OD600调整至0.6。将菌液按1:1 000比例加入DMEM培养基中，感染1 h后弃去上清，并加入DMEM培养基(含庆大霉素，终浓度50 μg/mL)。感染2 h后，弃去细胞培养液并用PBS洗涤2次，每孔加入3.6 mL琼脂覆盖液，冷却至琼脂凝固后，置于在37 ℃细胞培养箱中倒置培养，约48-60 h后可见空斑出现。每孔加入700 μL的40%甲醛溶液，在37 ℃培养箱中静置70 min以固定空斑。将琼脂凝块挑出，向每孔中加入0.5%的结晶紫溶液染色30 min，使用ddH2O冲洗未结合的结晶紫溶液，拍摄空斑图像并统计空斑直径。使用GraphPad软件分析数据，并通过t检验比较差异(ns：P>0.05)。
1.7 细菌的小鼠脏器定殖实验
选用4周龄SPF级雌性ICR小鼠为模型，分成3组，每组7只，饲养3-4 d。将EGD-e、ΔpplA及CΔpplA分别接种于BHI液体培养基中，37 ℃、200 r/min振荡培养过夜，并将菌液OD600调整至0.6。每只小鼠腹腔注射100 μL菌液，48 h后处死小鼠，取全部脾脏和部分肝脏称量并记录质量。将肝脏和脾脏研磨后点板计数，使用GraphPad软件分析数据，并通过t检验比较差异(*：P<0.05；***：P<0.001)。
本研究动物实验均已通过浙江农林大学实验动物伦理委员会审查，批准号：ZAFUAC202485。
1.8 小鼠体内单核增生李斯特氏菌毒力基因转录水平检测
将4周龄SPF级雌性ICR小鼠分为2组，每组5只。将EGD-e和ΔpplA分别接种于BHI液体培养基中，37 ℃、200 r/min振荡培养过夜，并将菌液OD600调整至0.6。腹腔注射菌液感染小鼠，48 h后取肝脏进行研磨。4 ℃、4 800 r/min离心10 min获取沉淀，将胰酶与ddH2O按照1:2的比例均匀加入匀浆沉淀静置30 min以裂解真核细胞；4 ℃、4 800 r/min离心10 min收集沉淀，加入DNA酶(5 mg/mL)和RNA酶(5 mg/mL)各100 μL，振荡混匀后于37 ℃培养箱静置30 min以降解真核细胞的DNA和RNA。4 ℃、4 800 r/min离心10 min收集菌体，加入溶菌酶裂解菌体，并使用TRIzol法提取细菌总RNA进行反转录。对表1所列的单核增生李斯特氏菌重要毒力基因(prfA、inlA、inlB、plcA、plcB、actA、hly)进行实时荧光定量PCR检测，以rpoB作为内参基因。PCR反应体系：2×Taq Pro Universal SYBR qPCR Master 10 μL，上、下游引物(1 µmol/L)各4 μL，cDNA 2 μL。PCR反应程序：95 ℃ 30 s；95 ℃ 10 s，60 ℃ 30 s，共40个循环。采用2-ΔΔCt算法分析数据并用GraphPad软件绘图，当log2 fold change (FC)>1时说明基因转录水平显著上调，log2FC < -1时说明基因转录水平显著下调，否则为基因转录水平无显著变化。
1.9 单核增生李斯特氏菌群体感应相关基因转录水平检测
将过夜培养的EGD-e和ΔpplA菌液分别转接入BHI培养基，置于37 ℃、200 r/min摇床振荡培养。当菌液的OD600值在0.4-0.5时，取1 mL菌液4 ℃、4 800 r/min离心10 min收集菌体。提取细菌总RNA进行反转录。采用荧光定量PCR方法研究PplA对单核增生李斯特氏菌Agr系统和LuxS系统群体感应相关基因(agrA、agrB、agrC、agrD和luxS，见表1)的转录调控。采用1.8节的反应体系、反应程序和数据分析算法进行研究。
2 结果与分析
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2.1 pplA 缺失后细菌溶血能力显著降低
通过比较EGD-e、ΔpplA和CΔpplA培养上清对绵羊血细胞的破坏能力，探究PplA对单核增生李斯特氏菌溶血能力的影响。如图1A所示，缺失株ΔpplA的溶血能力较野生株EGD-e显著降低，而回补株的溶血能力与野生株无显著差异。比较各菌株及阴性/阳性对照溶血后溶液的OD550值，绘制溶血曲线(图1B)并进行显著性分析，结果显示pplA缺失后的溶血能力显著低于野生株(P<0.001)，表明PplA能显著提高单核增生李斯特氏菌的溶血能力。
2.2 pplA 缺失后细菌对细胞的黏附侵袭能力显著降低
通过EGD-e、ΔpplA和CΔpplA感染Caco-2细胞，进行稀释点板计数，探究PplA在单核增生李斯特氏菌黏附侵袭Caco-2细胞中的作用。如图2A所示，ΔpplA感染后的细胞表面黏附细菌数较野生株显著减少(P<0.05)，而回补株与野生株差异不显著，表明PplA显著提高了单核增生李斯特氏菌对Caco-2细胞的黏附能力。如图2B所示，ΔpplA感染后侵袭至细胞内的细菌数量显著低于野生株和回补株(P<0.01)，即PplA能显著促进单核增生李斯特氏菌侵袭肠上皮细胞。
2.3 pplA 基因缺失后细菌的胞内增殖能力不变
使用EGD-e、ΔpplA和CΔpplA感染鼠源巨噬细胞RAW264.7后进行稀释点板计数，探究PplA在单核增生李斯特氏菌胞内增殖中的作用。如图3所示，在感染后2、5及8 h细胞中各菌株在细胞内的数量差异均不显著(P>0.05)。这表明PplA对单核增生李斯特氏菌在巨噬细胞RAW264.7中的增殖能力无显著影响。
2.4 pplA 缺失后细菌在细胞间的迁移能力不变
使用EGD-e、ΔpplA和CΔpplA感染L929细胞后，通过检测其形成的空斑大小来探究pplA对细菌胞间迁移能力的影响。如图4所示，各菌株形成的空斑大小和数量差异均不显著，表明EGD-e、ΔpplA和CΔpplA的胞间迁移能力差异不显著，说明PplA对单核增生李斯特氏菌的胞间迁移能力无显著影响。
2.5 pplA 缺失后细菌在小鼠脏器中的定殖能力显著降低
为了探究pplA对细菌毒力的影响，通过使用EGD-e、ΔpplA和CΔpplA感染小鼠，并对脾脏和肝脏细菌定殖数量进行统计。如图5所示，在感染48 h后，ΔpplA感染小鼠脾脏与肝脏中分离的细菌数量显著少于EGD-e和CΔpplA感染小鼠，表明pplA缺失后单核增生李斯特氏菌在小鼠脾脏与肝脏中的定殖能力显著减弱，即PplA对单核增生李斯特氏菌在宿主器官中的定殖起促进作用。
2.6 pplA 缺失后单核增生李斯特氏菌毒力因子 inlB 、 plcA 、 plcB 、 hly 和 prfA 转录水平降低
通过感染小鼠后单核增生李斯特氏菌毒力因子的转录水平测定，探究pplA对细菌在宿主细胞内的毒力因子转录水平的影响。如图6所示，在pplA缺失后，plcB、hly以及调控以上基因的单核增生李斯特氏菌重要毒力因子prfA的转录水平均显著下调，其中，prfA的变化最为显著，而inlA、actA、inlB和plcA则无显著变化。这表明在单核增生李斯特氏菌感染宿主时，PplA可以通过改变细菌毒力基因的转录水平进而促进单核增生李斯特氏菌的毒力。
2.7 pplA 缺失影响细菌群体感应相关基因 agrA 、 agrB 、 agrC 和 luxS 的转录
单核增生李斯特氏菌群体感应系统包括Agr系统和LuxS系统[16]。通过对pplA缺失的单核增生李斯特氏菌群体感应相关基因的转录水平研究，探究pplA对细菌群体感应系统的调控。结果显示pplA缺失后，Agr系统群体感应相关基因agrD转录水平未发生显著变化，而agrA和agrB显著上调，agrC显著下调；LuxS系统群体感应相关基因luxS显著下调(图7)。这表明单核增生李斯特氏菌中，PplA可以调控Agr系统和LuxS系统群体感应相关基因的转录水平。
3 讨论与结论
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本研究主要通过单核增生李斯特氏菌野生株EGD-e、ΔpplA和CΔpplA进行体内和体外感染实验，以探究pplA对单核增生李斯特氏菌致病力的影响。在体外研究中，使用绵羊血细胞、肠上皮细胞Caco-2、巨噬细胞RAW264.7以及成纤维细胞L929作为感染模型，以探究pplA对单核增生李斯特氏菌溶血活性、细胞黏附侵袭能力、胞内增殖和胞间迁移能力的影响。结果表明，单核增生李斯特氏菌缺失pplA后，其溶血能力显著降低，表明PplA在细菌的膜裂解能力中起到了促进作用。膜裂解能力是单核增生李斯特氏菌逃逸宿主吞噬泡的重要机制之一，PplA可能通过调控溶血素LLO的表达或活性来增强细菌的膜裂解能力。此外，pplA缺失株在细胞黏附与侵袭能力方面也显著减弱，说明PplA在感染初级阶段细菌与宿主细胞的相互作用中起到了关键作用，可能通过影响细菌表面蛋白的表达或修饰，增强细菌与宿主细胞的黏附能力。在胞内增殖和胞间迁移能力方面，pplA缺失株与野生株相比无显著性差异，表明PplA主要参与细菌的早期感染阶段。在体内试验中，以小鼠作为感染模型，结果显示pplA缺失后，单核增生李斯特氏菌在小鼠脏器中的定殖能力显著下降。为了进一步探究pplA是否影响单核增生李斯特氏菌感染宿主动物时毒力因子的表达水平，通过RT-qPCR检测了感染小鼠体内单核增生李斯特氏菌主要毒力因子的基因转录水平。结果表明，在缺失pplA后，其毒力因子plcB、hly和prfA基因转录水平显著下调，这与感染结果一致，说明PplA不仅在感染的初级阶段影响单核增生李斯特氏菌，还通过调控其毒力因子的表达来增强细菌在宿主体内的生存和增殖能力。这些结果进一步说明了PplA在调控单核增生李斯特氏菌毒力中的重要作用。
群体感应机制对细菌的致病力具有调控作用，当细菌密度达到一定水平时，群体感应使细菌接收信号分子，进而触发一系列级联反应和基因表达，导致相关毒力因子的产生和分泌，从而影响细菌的毒力[14]。群体感应系统最初在海洋细菌中被报道，当细胞群密度足以产生自诱导剂的积累时会触发生物发光[17]。革兰氏阳性菌通常利用分泌的寡肽和双组分系统进行群体感应，这些系统由膜结合的传感器激酶受体和能够调控基因表达的细胞质转录因子组成[18]。群体感应在革兰氏阳性菌中的生物学作用包括协调群体行为，如生物膜形成、毒力因子产生和抗生素抗性等[18]。例如，金黄色葡萄球菌中存在一种典型的双组分系统，即所谓的辅助基因调节因子(auxiliary gene regulation, Agr)系统，该系统以反应调节因子AgrA和调节RNAIII作为主要的细胞内效应子，能够感应细菌密度，调整细菌毒力因子表达的时机以响应应激以及代谢和环境方面的动态变化[19]。芽孢杆菌属也存在群体感应系统，可通过感受细菌密度调节相关基因表达，其调控网络的一个重要参与者是信息素ComX，它在整个过程中起诱导剂的作用，除此以外，枯草芽孢杆菌还存在ComQXPA这一四蛋白群体感应系统[20]。肺炎链球菌具有TprA/PhrA肽信号系统，能够介导群体感应，帮助其在鼻咽处定殖，除此以外该系统还能控制抗微生物肽的基因簇表达，从而影响肺炎链球菌在鼻咽定殖期间的竞争性适应性[21]。革兰氏阳性菌的群体感应通常利用小的翻译后修饰的肽信号分子，这些肽信号与组氨酸激酶双组分信号转导系统的传感器元件相互作用，能够调节枯草芽孢杆菌和肺炎链球菌中细菌能力的发育、粪肠球菌中的结合以及金黄色葡萄球菌中的毒力[22]。单核增生李斯特氏菌中也存在群体感应系统，其中Agr群体感应系统是其重要组成部分[23-24]。相关研究表明，Agr系统中agrD基因的缺失会导致细菌对宿主的侵袭能力降低、在小鼠体内毒力衰减以及prfA依赖性基因表达水平的降低[23]，而agrA基因的缺失则会使细菌在宿主内的脏器定殖能力降低[24]。LuxS/AI-2群体感应系统中的luxS基因，其缺失可减弱细菌的生物被膜形成能力，但对细菌的溶血活性和磷脂酶活性并无影响[25]。单核增生李斯特氏菌信息素脂蛋白PplA产生的多肽信息素pPplA可以介导细菌之间的信号传递，有利于感染中吞噬体膜的破坏和细菌的逃逸[13]。在本研究中，首先探究了PplA是否调节Agr和LuxS群体感应系统。结果表明，缺失PplA并不影响与细菌毒力相关的Agr系统中agrD基因的转录水平，但会使agrA基因的转录水平上调。由于agrA基因能够促进单核增生李斯特氏菌的毒力，且PplA本身也具有促进单核增生李斯特氏菌毒力的功能。由此可以推断，PplA发挥其促进细菌毒力的功能并非通过调控agrA基因来实现的。相比之下，LuxS群体感应系统在单核增生李斯特氏菌感染中的作用尚不明确，因此PplA调控LuxS群体感应系统介导的致病机制也仍待探究。此外，研究发现PplA可以提高感染小鼠体内单核增生李斯特氏菌重要毒力因子prfA、hly、plcA和plcB等的基因转录水平。因此，推测PplA的功能发挥可能类似于枯草芽孢杆菌和肺炎链球菌，即PplA产生的多肽信息素pPplA与双组分信号转导系统的传感器元件或转录调节因子等调控元件相互作用，提高转录激活因子PrfA的表达，PrfA进而提高毒力因子LLO、PlcA和PlcB等的表达，从而提高单核增生李斯特氏菌在宿主内的存活能力(图8)。然而，该过程的具体机制仍需进一步探究。
本研究通过比较单核增生李斯特氏菌野生株、脂蛋白PplA缺失株和回补株在细胞溶血能力、细胞黏附和侵袭能力、胞内增殖能力、胞间迁移能力、小鼠脏器定殖能力、小鼠脏器内细菌毒力因子转录水平以及群体感应相关基因转录水平等方面的差异，揭示了PplA在单核增生李斯特氏菌感染宿主过程中的作用，为理解单核增生李斯特氏菌的感染机制提供了新的分子基础，也为抗感染治疗提供了潜在靶点。
基金
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  • 国家重点研发计划(2023YFD1801800)
  • 宁波市自然科学基金(2023J241)
  • 浙江省自然科学基金(LY23C180002)
文献
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参考文献 引证文献
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2025年第65卷第8期
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doi: 10.13343/j.cnki.wsxb.20240715
  • 接收时间：2024-11-12
  • 首发时间：2026-02-06
  • 出版时间：2025-08-04
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  • 收稿日期：2024-11-12
  • 录用日期：2025-04-30
基金
National Key Research and Development Program of China(2023YFD1801800)
国家重点研发计划(2023YFD1801800)
Ningbo Natural Science Foundation(2023J241)
宁波市自然科学基金(2023J241)
Natural Science Foundation of Zhejiang Province(LY23C180002)
浙江省自然科学基金(LY23C180002)
作者信息
    1.浙江农林大学 动物医学院，浙江省畜禽绿色生态健康养殖应用技术研究重点实验室，动物健康互联网检测技术浙江省工程研究中心，浙江省动物医学与健康管理国际科技合作基地，同一健康和食品安全“一带一路”国际联合实验室，中澳动物健康大数据分析联合实验室，浙江 杭州
    2.宁波卫生职业技术学院，浙江 宁波

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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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