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Article(id=1226956558710521893, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226956547847275311, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250180, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1741190400000, receivedDateStr=2025-03-06, revisedDate=null, revisedDateStr=null, acceptedDate=1744905600000, acceptedDateStr=2025-04-18, onlineDate=1770458839347, onlineDateStr=2026-02-07, pubDate=1756915200000, pubDateStr=2025-09-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1770458839347, onlineIssueDateStr=2026-02-07, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1770458839347, creator=13701087609, updateTime=1770458839347, updator=13701087609, issue=Issue{id=1226956547847275311, tenantId=1146029695717560320, journalId=1192105938417971205, year='2025', volume='65', issue='9', pageStart='3821', pageEnd='4232', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1770458836757, creator=13701087609, updateTime=1770459153781, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1226957877613605816, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226956547847275311, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1226957877613605817, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226956547847275311, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=4162, endPage=4173, ext={EN=ArticleExt(id=1226956559587131480, articleId=1226956558710521893, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Biological characteristics and pathogenicity of a uvrY-deleted mutant of Vibrio parahaemolyticus, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

UvrY is a key response regulator of the BarA/UvrY two-component system (TCS) and plays an important role in regulating bacterial virulence and environmental adaptability. [Objective] To investigate the regulatory role of UvrY in the biological characteristics and pathogenicity of Vibrio parahaemolyticus SH112. [Methods] The uvrY-deleted mutant (ΔuvrY) and its complementary strain (CΔuvrY) were constructed by homologous recombination. Phenotypes were systematically compared among the wild type, mutant, and complementary strains by growth curve plotting, motility (swimming and swarming) assays, biofilm formation assay, bacterial competition assay, HeLa cell adhesion and cytotoxicity assays, as well as a mouse infection model (analysis of bacterial loads in tissues and lethality). [Results] Compared with the wild type strain, ΔuvrY exhibited significant growth defects during the late exponential phase and weakened motility, with swimming and swarming reduced by 33% and 70%, respectively, while the biofilm formation of the mutant remained unaffected. Additionally, ΔuvrY showed weakened competitive inhibition against Escherichia coli, a 36.7% reduction in HeLa cell adhesion, and a 15.8% decrease in cytotoxicity. Mouse infection experiments further demonstrated that ΔuvrY had significantly reduced tissue colonization capacity and the attenuation of 75% in pathogenicity. [Conclusion] This study reveals that UvrY plays a crucial role in the pathogenicity of V. parahaemolyticus by regulating the growth, motility, competitive ability, and interaction with the host, giving insights into the regulatory network of the BarA/UvrY two-component system.

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*E-mail: JIANG Wei,
HAN Xiangan,
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#These authors contributed equally to this work.

, authorsList=Shuqi LU, Chuang RUI, Rong GUO, Peijie WU, Suoping QIU, Weihuan FANG, Tingting LI, Xiangan HAN, Wei JIANG), CN=ArticleExt(id=1226956562929992025, articleId=1226956558710521893, tenantId=1146029695717560320, journalId=1192105938417971205, language=CN, title=副溶血弧菌 uvrY 基因缺失株的生物学特性及致病性分析, columnId=1192149544164012138, journalTitle=微生物学报, columnName=研究报告, runingTitle=null, highlight=null, articleAbstract=

UvrY是BarA/UvrY双组分系统的关键反应调节因子，在调控细菌毒力与环境适应性中起重要作用。 【目的】 探究UvrY对副溶血弧菌SH112株的生物学特性及致病性的调控作用。 【方法】 通过同源重组技术构建uvrY基因缺失株(ΔuvrY)及其互补株(CΔuvrY)，并采用生长曲线测定、运动能力(泳动和群集运动)、生物被膜形成、细菌竞争、HeLa细胞黏附能力与毒性实验，以及小鼠感染模型(组织载菌量和致死率分析)等多方面实验，系统比较野生株、缺失株和互补株的表型差异。 【结果】 与野生株相比，ΔuvrY菌株在指数生长后期表现出显著的生长缺陷；运动能力明显下降，其中泳动和群集运动分别降低了33%和70%，但生物被膜形成能力未受到显著影响。此外，ΔuvrY菌株对大肠杆菌的竞争抑制能力减弱，对HeLa细胞的黏附率降低了36.7%，细胞毒性下降了15.8%。小鼠感染实验进一步表明，ΔuvrY菌株的组织定殖能力显著降低，致病性减弱了75%。 【结论】 本研究揭示了UvrY通过调控副溶血弧菌的生长、运动性、竞争能力和宿主互作等过程在其致病机制中发挥关键作用，为深入理解BarA/UvrY双组分系统的调控网络提供了重要依据。

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作者贡献声明

卢淑淇：实验操作、数据分析和撰写文章；芮闯：数据收集和监督管理；郭容：实验操作、数据收集与分析；吴佩洁、邱索平、方维焕、李婷婷：软件程序、监督管理；韩先干：研究构思和设计；蒋蔚：获取基金、项目管理、论文审阅和修改。

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M: DL2000 DNA marker; Lanes 1-3: Target fragments amplified with WT, ΔuvrY and CΔuvrY as templates using uvrY E/F primers; Lanes 4-6: Target fragments amplified with WT, ΔuvrY and CΔuvrY as templates, and the target fragment amplified withuvrY-pMMB-F/R primers; Lane 7: The target fragment of ΔuvrY amplified with sacB-F/R primers., figureFileSmall=0tpUvxZufz3Pjc+/YX3GJg==, figureFileBig=MkDS3GrUohFR03Br+x7iPw==, tableContent=null), ArticleFig(id=1226964059686355937, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=CN, label=图1, caption=uvrY 基因缺失株及其互补株的PCR鉴定结果。M：DL2000 DNA marker；泳道1-3：以WT、ΔuvrY和CΔuvrY为模板，用uvrY-E/F引物扩增的目的片段；泳道4-6：以WT、ΔuvrY和CΔuvrY为模板，用uvrY-pMMB-F/R引物扩增的目的片段；泳道7：以ΔuvrY为模板，用sacB-F/R引物扩增的目的片段。, figureFileSmall=0tpUvxZufz3Pjc+/YX3GJg==, figureFileBig=MkDS3GrUohFR03Br+x7iPw==, tableContent=null), ArticleFig(id=1226964059828962285, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=EN, label=Figure 2, caption=The growth curves of the WT, ΔuvrY and CΔuvrY. *: P<0.05., figureFileSmall=im6kC5xe+ZNS9EIsyN0ZZA==, figureFileBig=aPDcYsqnR34Q42iBUp6uqA==, tableContent=null), ArticleFig(id=1226964059929625588, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=CN, label=图2, caption=WT、ΔuvrY 和CΔuvrY 的生长曲线, figureFileSmall=im6kC5xe+ZNS9EIsyN0ZZA==, figureFileBig=aPDcYsqnR34Q42iBUp6uqA==, tableContent=null), ArticleFig(id=1226964060034483194, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=EN, label=Figure 3, caption=Motility analysis. A, B: The swimming pattern of the strains; C, D: The swarming pattern of the strains; E: Growth of each strain during swimming motility. ****: P<0.000 1., figureFileSmall=AsvjD5bMe+SVFZTRjX0OHg==, figureFileBig=GkKPRtek+7a91ZRWOSyuQw==, tableContent=null), ArticleFig(id=1226964060172895230, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=CN, label=图3, caption=运动性分析。A、B：菌株的泳动运动；C、D：菌株的群集运动；E：各菌株在泳动过程中的生长情况。, figureFileSmall=AsvjD5bMe+SVFZTRjX0OHg==, figureFileBig=GkKPRtek+7a91ZRWOSyuQw==, tableContent=null), ArticleFig(id=1226964060323889155, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=EN, label=Figure 4, caption=Biofilm forming capacity. Quantitative profiling of biofilm formation ability in bacterial strains using crystal violet assay: phenotypic morphology (A) and spectrophotometric quantification (B) following 48 h static incubation., figureFileSmall=QAQz+wwAPBUHv87Vyz5L9w==, figureFileBig=kHPKKZGOs0sik3+3Yoo0NA==, tableContent=null), ArticleFig(id=1226964060458106885, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=CN, label=图4, caption=生物被膜形成能力。通过结晶紫染色法观察菌株在48 h内形成生物被膜的能力(A)和测量数据(B)。, figureFileSmall=QAQz+wwAPBUHv87Vyz5L9w==, figureFileBig=kHPKKZGOs0sik3+3Yoo0NA==, tableContent=null), ArticleFig(id=1226964060554575889, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=EN, label=Figure 5, caption=Antibacterial efficacy of V. parahaemolyticus against E. coli in co-culture systems. A, B: Comparative quantification of E. coli and V. parahaemolyticus populations at pre-interaction (0 h) vs. post-competitive (4 h) phases under 30 °C; C, D: Comparative quantification of E. coli and V. parahaemolyticus populations at pre-interaction (0 h) vs. post-competitive (4 h) phases under 37 °C. ***: P<0.001; ****: P<0.000 1., figureFileSmall=mWxjMrWmJLQvyMHK+hnAJQ==, figureFileBig=BNnACedDGY1B8MbwFfkxzA==, tableContent=null), ArticleFig(id=1226964060688793623, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=CN, label=图5, caption=副溶血弧菌菌株与大肠杆菌共培养时的抗菌作用。A、B：30 ℃下竞争前(0 h)和竞争后(4 h)大肠杆菌和副溶血弧菌的计数；C、D：37 ℃下竞争前(0 h)和竞争后(4 h) 大肠杆菌和副溶血弧菌的计数。***： P<0.001；****：P<0.000 1。, figureFileSmall=mWxjMrWmJLQvyMHK+hnAJQ==, figureFileBig=BNnACedDGY1B8MbwFfkxzA==, tableContent=null), ArticleFig(id=1226964060764291103, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=EN, label=Figure 6, caption=Adhesion of each strain to HeLa cells. A: Adhesion to HeLa cell monolayers of the strains; B: Growth of each strain 1 h after infection of HeLa cells. ***: P<0.001., figureFileSmall=9wBmX7JCudCav75gLkRU5w==, figureFileBig=RQtVVJ0fHmLfzv3xIhbJbA==, tableContent=null), ArticleFig(id=1226964060843982886, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=CN, label=图6, caption=各菌株对HeLa细胞的黏附作用。A：菌株对HeLa细胞的黏附作用；B：感染HeLa细胞1 h后各菌株的生长情况。, figureFileSmall=9wBmX7JCudCav75gLkRU5w==, figureFileBig=RQtVVJ0fHmLfzv3xIhbJbA==, tableContent=null), ArticleFig(id=1226964060982394925, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=EN, label=Figure 7, caption=Cytotoxic effects of the WT, ΔuvrY, and CΔuvrY on HeLa cells. **: P<0.01., figureFileSmall=oCNBuWO5vGHFZKwC31qAcQ==, figureFileBig=TDa37tcsBHTxmP5tPcyw8A==, tableContent=null), ArticleFig(id=1226964061116612662, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=CN, label=图7, caption=WT、ΔuvrY 和CΔuvrY 对HeLa细胞的毒性影响, figureFileSmall=oCNBuWO5vGHFZKwC31qAcQ==, figureFileBig=TDa37tcsBHTxmP5tPcyw8A==, tableContent=null), ArticleFig(id=1226964061255024699, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=EN, label=Figure 8, caption=The survival rate of ICR mice infected with different strains., figureFileSmall=qZApMV1ICjuD4zWIIGQabA==, figureFileBig=Aha8lHT0J5wHTcnBQMHnJA==, tableContent=null), ArticleFig(id=1226964061372465221, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=CN, label=图8, caption=各菌株感染ICR小鼠后的存活率, figureFileSmall=qZApMV1ICjuD4zWIIGQabA==, figureFileBig=Aha8lHT0J5wHTcnBQMHnJA==, tableContent=null), ArticleFig(id=1226964061515071563, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=EN, label=Figure 9, caption=The colonization ability of each strain. Bacterial loads in heart (A), liver (B), spleen (C), kidney (D) of infected mice. **: P<0.01; ****: P<0.000 1., figureFileSmall=ccjdT8PqS9+mhEz5neg3hg==, figureFileBig=OnpxThw4zvC8dERogcAjsA==, tableContent=null), ArticleFig(id=1226964061632512079, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=CN, label=图9, caption=各菌株的定殖能力。各菌株在小鼠心(A)、肝(B)、脾(C)、肾(D)组织的细菌载量。, figureFileSmall=ccjdT8PqS9+mhEz5neg3hg==, figureFileBig=OnpxThw4zvC8dERogcAjsA==, tableContent=null), ArticleFig(id=1226964061733175382, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=EN, label=Table 1, caption=

Primers used in this experiment

, figureFileSmall=null, figureFileBig=null, tableContent=
Primers namePrimer sequences (5′→3′)Restriction enzymeProduct size (bp)
uvrY-AACCGGATCCTTGGCGAGTAAGTAGGAAACBamH I548
uvrY-BACTTGTATCTCCACACTTTTTATTGGC
uvrY-CTGTGGAGATACAAGTTGTGAATCCTCCTTTC606
uvrY-DACCCTGCAGCGTCTTATCGCTTGTATCTGPst I
uvrY-ETTCTGGATACGATGAAAGCC

Wide type: 2 461

Mutant: 1 816

uvrY-FTACCTCTTGTTGGCTCGTGT
uvrY-pMMB-FCGCGGCCTGCAG TTGATTAATGTTTTCCTTGTPst I663
uvrY-pMMB-RAATGAGCTCCTAAAGGGTCTCGGTGTCCASac I
sacB-FACGGCACTGTCGCAAACTATA600
sacB-RTTCCGTCACCGTCAAAGAT
), ArticleFig(id=1226964061905141856, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956558710521893, language=CN, label=表1, caption=

本研究所用引物

, figureFileSmall=null, figureFileBig=null, tableContent=
Primers namePrimer sequences (5′→3′)Restriction enzymeProduct size (bp)
uvrY-AACCGGATCCTTGGCGAGTAAGTAGGAAACBamH I548
uvrY-BACTTGTATCTCCACACTTTTTATTGGC
uvrY-CTGTGGAGATACAAGTTGTGAATCCTCCTTTC606
uvrY-DACCCTGCAGCGTCTTATCGCTTGTATCTGPst I
uvrY-ETTCTGGATACGATGAAAGCC

Wide type: 2 461

Mutant: 1 816

uvrY-FTACCTCTTGTTGGCTCGTGT
uvrY-pMMB-FCGCGGCCTGCAG TTGATTAATGTTTTCCTTGTPst I663
uvrY-pMMB-RAATGAGCTCCTAAAGGGTCTCGGTGTCCASac I
sacB-FACGGCACTGTCGCAAACTATA600
sacB-RTTCCGTCACCGTCAAAGAT
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副溶血弧菌 uvrY 基因缺失株的生物学特性及致病性分析
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卢淑淇 1 , 芮闯 2 , 郭容 1 , 吴佩洁 3 , 邱索平 3 , 方维焕 4 , 李婷婷 2 , 韩先干 1 , 蒋蔚 1
微生物学报 | 研究报告 2025,65(9): 4162-4173
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微生物学报 | 研究报告 2025, 65(9): 4162-4173
副溶血弧菌 uvrY 基因缺失株的生物学特性及致病性分析
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卢淑淇1, 芮闯2, 郭容1, 吴佩洁3, 邱索平3, 方维焕4, 李婷婷2, 韩先干1 , 蒋蔚1
作者信息
  • 1 中国农业科学院上海兽医研究所，上海
  • 2 上海健康医学院，上海
  • 3 从化海关综合技术服务中心，广东 广州
  • 4 浙江农林大学 动物科技学院，浙江 临安
Biological characteristics and pathogenicity of a uvrY-deleted mutant of Vibrio parahaemolyticus
Shuqi LU1, Chuang RUI2, Rong GUO1, Peijie WU3, Suoping QIU3, Weihuan FANG4, Tingting LI2, Xiangan HAN1 , Wei JIANG1
Affiliations
  • 1 Shanghai Veterinary Research Institute, Chinese Academy of Agricultural Sciences, Shanghai, China
  • 2 Shanghai University of Medicine & Health Sciences, Shanghai, China
  • 3 Conghua Customs Comprehensive Technical Service Center, Guangzhou, Guangdong, China
  • 4 College of Animal Science and Technology, Zhejiang A&F University, Lin’an, Zhejiang, China
出版时间: 2025-09-04 doi: 10.13343/j.cnki.wsxb.20250180
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摘要
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UvrY是BarA/UvrY双组分系统的关键反应调节因子，在调控细菌毒力与环境适应性中起重要作用。 【目的】 探究UvrY对副溶血弧菌SH112株的生物学特性及致病性的调控作用。 【方法】 通过同源重组技术构建uvrY基因缺失株(ΔuvrY)及其互补株(CΔuvrY)，并采用生长曲线测定、运动能力(泳动和群集运动)、生物被膜形成、细菌竞争、HeLa细胞黏附能力与毒性实验，以及小鼠感染模型(组织载菌量和致死率分析)等多方面实验，系统比较野生株、缺失株和互补株的表型差异。 【结果】 与野生株相比，ΔuvrY菌株在指数生长后期表现出显著的生长缺陷；运动能力明显下降，其中泳动和群集运动分别降低了33%和70%，但生物被膜形成能力未受到显著影响。此外，ΔuvrY菌株对大肠杆菌的竞争抑制能力减弱，对HeLa细胞的黏附率降低了36.7%，细胞毒性下降了15.8%。小鼠感染实验进一步表明，ΔuvrY菌株的组织定殖能力显著降低，致病性减弱了75%。 【结论】 本研究揭示了UvrY通过调控副溶血弧菌的生长、运动性、竞争能力和宿主互作等过程在其致病机制中发挥关键作用，为深入理解BarA/UvrY双组分系统的调控网络提供了重要依据。

副溶血弧菌  /  UvrY  /  双组分系统  /  致病性  /  毒力调控

UvrY is a key response regulator of the BarA/UvrY two-component system (TCS) and plays an important role in regulating bacterial virulence and environmental adaptability. [Objective] To investigate the regulatory role of UvrY in the biological characteristics and pathogenicity of Vibrio parahaemolyticus SH112. [Methods] The uvrY-deleted mutant (ΔuvrY) and its complementary strain (CΔuvrY) were constructed by homologous recombination. Phenotypes were systematically compared among the wild type, mutant, and complementary strains by growth curve plotting, motility (swimming and swarming) assays, biofilm formation assay, bacterial competition assay, HeLa cell adhesion and cytotoxicity assays, as well as a mouse infection model (analysis of bacterial loads in tissues and lethality). [Results] Compared with the wild type strain, ΔuvrY exhibited significant growth defects during the late exponential phase and weakened motility, with swimming and swarming reduced by 33% and 70%, respectively, while the biofilm formation of the mutant remained unaffected. Additionally, ΔuvrY showed weakened competitive inhibition against Escherichia coli, a 36.7% reduction in HeLa cell adhesion, and a 15.8% decrease in cytotoxicity. Mouse infection experiments further demonstrated that ΔuvrY had significantly reduced tissue colonization capacity and the attenuation of 75% in pathogenicity. [Conclusion] This study reveals that UvrY plays a crucial role in the pathogenicity of V. parahaemolyticus by regulating the growth, motility, competitive ability, and interaction with the host, giving insights into the regulatory network of the BarA/UvrY two-component system.

Vibrio parahaemolyticus  /  UvrY  /  two-component system  /  pathogenicity  /  virulence regulation
卢淑淇, 芮闯, 郭容, 吴佩洁, 邱索平, 方维焕, 李婷婷, 韩先干, 蒋蔚. 副溶血弧菌 uvrY 基因缺失株的生物学特性及致病性分析. 微生物学报, 2025 , 65 (9) : 4162 -4173 . DOI: 10.13343/j.cnki.wsxb.20250180
Shuqi LU, Chuang RUI, Rong GUO, Peijie WU, Suoping QIU, Weihuan FANG, Tingting LI, Xiangan HAN, Wei JIANG. Biological characteristics and pathogenicity of a uvrY-deleted mutant of Vibrio parahaemolyticus[J]. Acta Microbiologica Sinica, 2025 , 65 (9) : 4162 -4173 . DOI: 10.13343/j.cnki.wsxb.20250180
正文
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副溶血弧菌(Vibrio parahaemolyticus)是一种革兰氏阴性菌，广泛分布在气候条件温暖的海洋和河口环境中[1-2]。自1950年首次分离鉴定以来，副溶血弧菌已逐渐成为全球食源性海产品中毒事件的首要致病因子，并引起虾等水产动物的急性坏死性肝胰腺炎，对水产养殖业造成经济损失，严重威胁公共卫生安全[3]。截至目前，其主要毒力因子包括黏附素、溶血素、III型分泌系统(type III secretion system, T3SS)以及VI型分泌系统(type VI secretion system, T6SS)[4-6]。然而副溶血弧菌的致病机制仍未被完全阐明。因此深入研究副溶血弧菌的致病机理，尤其是关键信号转导系统的功能，将为开发精准防控策略提供理论依据。
在细菌适应宿主环境的过程中，信号转导系统通过感知外界刺激并协调毒力因子表达调控致病作用。其中，双组分系统在原核生物中广泛存在，调控多种细胞过程[7-8]。UvrY作为BarA/UvrY双组分系统的反应调节因子，在细菌中发挥转录因子的作用[9]。BarA/UvrY双组分系统是信号转导的关键，由跨膜感应激酶蛋白BarA及其协同调控的胞内反应调节蛋白UvrY构成[10]。BarA通过感知环境信号发生自磷酸化，随后将磷酸基团转移至UvrY，调控非编码RNA控制下游基因表达[11]。该调控模块在大肠杆菌、副溶血弧菌、沙门氏菌等革兰氏阴性病原菌中的毒力调控具有高度保守性[12-14]。BarA/UvrY双组分系统精密调控细菌的运动性、生物被膜形成、黏附、定殖及毒力因子表达等关键生理过程[15-17]。例如，UvrY可正向调控大肠杆菌I型菌毛的转录，增强其肠道定殖能力[18]。在气单胞菌中，UvrY通过调控一系列毒力因子来影响病原体的毒力[19]。此外，研究表明BarA/UvrY是副溶血弧菌的重要毒力因子，能够调控III型和VI型分泌系统等毒力因子，在致病过程中发挥关键作用[13]。BarA/UvrY系统在许多细菌的毒力调控中发挥了核心作用。然而副溶血弧菌的BarA/UvrY双组分系统与毒力之间的密切关系目前尚不完全清楚。
本研究成功构建了副溶血弧菌SH112株uvrY基因缺失株及互补株，并对各菌株的生物学特性进行了深入分析，以期为揭示BarA/UvrY双组分系统与副溶血弧菌生物学特性和毒力调控之间关联性的研究提供重要依据。
1 材料与方法
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1.1 菌种和质粒
本研究所用菌株及质粒均由本实验室保存：副溶血弧菌SH112 (tdh+，血清型O3:K6)、携带pRK2013质粒的大肠杆菌HB101、大肠杆菌CC118λpir、含pBAD33-Gm质粒的大肠杆菌DH5α、自杀质粒pYAK1、互补质粒pMMB207、HeLa细胞。
1.2 主要试剂和仪器
细菌基因组提取试剂盒、质粒提取试剂盒，天根生化科技(北京)有限公司；CytoTox 96®非放射性细胞毒性检测试剂盒，Promega公司；BamH I、Pst I、Sac I内切酶与T4 DNA连接酶，TaKaRa公司；Triton X-100，国药集团化学试剂有限公司；DMEM培养基、胎牛血清，Gibco公司。
全波长酶标仪、PCR仪，Eppendorf公司。
1.3 实验动物
Institute of Cancer Research (ICR)小鼠购自上海杰思捷实验动物有限公司。本研究所有动物实验经中国农业科学院上海兽医研究所动物伦理委员会批准，编号为SYXK (沪) 2020-0027。
1.4 引物设计与合成
根据GenBank中副溶血弧菌RIMD2210633标准株的uvrY基因序列，采用Primer Premier 5.0软件设计特异性引物(表1)。其中，sacB-F/R引物特异性靶向质粒pYAK1多克隆位点两侧的sacB标记基因，通过PCR特异性检测同源重组后质粒在宿主菌中的丢失情况。该筛选策略利用sacB基因编码的果聚糖蔗糖酶特性，当质粒成功切除时sacB基因随载体同步丢失，从而实现重组菌株的精准筛选。
1.5 uvrY 基因缺失株与互补株的构建
参考文献[20]构建uvrY基因缺失株及互补株。以副溶血弧菌SH112株为野生株(wild type strain, WT)，并提取全基因组作为扩增模板，所用引物见表1。分别使用引物uvrY-A/B和uvrY-C/D扩增同源臂AB、CD片段，通过重叠PCR获得缺失uvrY基因的融合片段uvrY-AD。将该片段经BamH I/Pst I酶切后连接至自杀质粒pYAK1，获得重组质粒pYAK1-ΔuvrY。通过接合转移(供体菌：CC118λpir/pYAK1-ΔuvrY；受体菌：SH112；辅助菌：pRK2013-HB101)将重组质粒导入SH112。用含10 μg/mL氯霉素的TCBS平板和含20%蔗糖的LB平板筛选结合子。将筛选到的疑似缺失株用相应引物进行鉴定，PCR产物送至生工生物工程(上海)股份有限公司进行测序分析，经鉴定正确的缺失株命名为ΔuvrY。经过20代传代后，用引物uvrY-E/F和sacB-F/R进行鉴定，确定自杀质粒已丢失，保存菌株并用于后续实验。
参考构建缺失株的方法获得互补质粒pMMB207-uvrY，并通过结合转移的方法导入副溶血弧菌SH112，分别使用uvrY-E/F和uvrY-pMMB-F/R特异性引物进行PCR验证，经电泳检测将阳性菌株命名为互补株CΔuvrY。
1.6 生长曲线的测定
将过夜培养的WT、ΔuvrY及CΔuvrY菌株分别接种至含3% NaCl的LB液体培养基中，在37 ℃、180 r/min振荡培养至对数生长期(OD600为0.20±0.02)。采用无菌96孔板进行动态监测，每组设置3个平行重复，按100 μL/孔定量接种菌悬液，使用酶标仪测定OD600 (间隔1 h)，持续监测12 h。最终通过生长曲线比较分析各菌株间的增殖差异。
1.7 运动性测定
根据Li等[21]的方法，分别检测菌株的泳动和群集运动能力。泳动运动(swimming)：将WT、ΔuvrY及CΔuvrY培养至对数生长期，各取1 µL的菌液垂直接种于泳动平板(含0.3%琼脂、3% NaCl的LB培养基)上，37 ℃恒温培养4 h后观察扩散圈；群集运动(swarming)：用含1.5%琼脂的HIB培养基制备运动平板，接种1 µL对数期菌液后置于30 ℃培养箱中持续孵育16 h。拍照记录泳动和群集运动情况，并测量直径。
1.8 生物被膜的测定
采用结晶紫染色法评估细菌生物被膜形成能力[15]。将各菌株培养至对数生长期，取200 μL菌液加入96孔板中，分别在30 ℃和37 ℃静置孵育48 h。用PBS洗涤3次，甲醇固定15 min，结晶紫染色15 min，PBS洗涤并晾干后，加入95%乙醇溶解，测量OD595值。
1.9 细菌竞争试验
参照文献[22]进行细菌竞争试验。将WT、ΔuvrY及CΔuvrY与大肠杆菌DH5α (含pBAD33质粒)按4:1的比例混合(副溶血弧菌为攻击者，大肠杆菌为猎物)。取其中1份倍比稀释后分别涂布于TCBS培养基和含有庆大霉素的LB平板上，以确定共培养0 h时攻击者和猎物的菌体浓度(CFU/mL)；另外2份混合菌液分别在30 ℃和37 ℃共培养4 h后稀释至10-4-10-6，涂布于相应平板上，获得共培养4 h时各副溶血弧菌和大肠杆菌的菌体浓度(CFU/mL)。
1.10 细胞黏附试验
将HeLa细胞接种于24孔板，于37 ℃、5% CO2条件下培养至约80%汇合度。分别以MOI=10:1的WT、ΔuvrY及CΔuvrY感染细胞1 h，加入含0.5% Triton X-100的裂解液处理20 min。裂解液经PBS梯度稀释后涂布于含3% NaCl的LB平板，记录菌落数量，计算相对黏附效率。
1.11 细胞毒性检测
将HeLa细胞接种于96孔细胞板中，培养至约80%汇合度。将各菌株培养至对数生长期，以MOI=10:1感染细胞，37 ℃、5% CO2条件下孵育1.5 h。按照CytoTox96®试剂盒说明书操作，反应终止后通过酶标仪测定OD490，并计算各菌株的细胞毒性率。
1.12 小鼠致病性分析
参照Bai等[22]的方法，将3-4周龄雌性ICR小鼠随机分为4组(WT组、ΔuvrY组、CΔuvrY组和空白组，n=8)，分别通过腹腔注射接种终浓度为3×108 CFU/mL的菌悬液或等体积无菌生理盐水。感染后连续监测7 d，每小时记录小鼠的存活状态。
1.13 组织载量测定
参考文献[23]将5周龄的小鼠分成4组，每组10只，分别通过腹腔注射接种终浓度为5×107 CFU/mL的对数生长期菌悬液和无菌生理盐水。感染10 h后无菌解剖取样，用PBS倍比稀释，涂布于TCBS琼脂平板上，以评估细菌定植情况。
1.14 数据分析
连续变量以均值±标准差(mean±SD)形式表达；采用GraphPad Prism v8.0软件进行单因素方差分析(one-way ANOVA)，以P<0.05作为差异显著性阈值。
2 结果与分析
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2.1 uvrY 基因缺失株和互补株的鉴定
采用菌株的基因组DNA作为扩增模板进行PCR验证(图1)。以WT基因组为模板，引物uvrY-E/F扩增出大小约为2 500 bp的特异性条带(图1，泳道1)，而以ΔuvrY及CΔuvrY基因组为模板，均呈现约1 800 bp的扩增产物(图1，泳道2、3)。PCR产物经比对正确，WT和ΔuvrY扩增出的目的片段大小分别是2 461 bp和1 816 bp。用引物uvrY-pMMB-F/R扩增WT和CΔuvrY基因组，获得目的片段(图1，泳道4、6)，而ΔuvrY扩增不出目的条带(图1，泳道5)，经测序比对表明目的片段为663 bp，表明互补株构建正确。多次传代后，采用sacB-F/R引物对ΔuvrY进行自杀质粒的丢失情况进行筛查，电泳未检测到sacB基因特异性条带(图1，泳道7)，表明ΔuvrY与CΔuvrY构建成功且传代稳定，可用于后续实验。
2.2 uvrY 基因参与调控副溶血弧菌生长能力
如图2所示，在相同的培养条件下(37 ℃、含3% NaCl的LB培养基)培养6 h内uvrY基因的缺失对副溶血弧菌的生长无显著影响。从7 h开始，缺失株ΔuvrY的生长速度低于WT和CΔuvrY (*：P<0.05)。这表明uvrY基因参与调控副溶血弧菌的生长。
2.3 UvrY参与副溶血弧菌的泳动及群集运动
各菌株在泳动平板上培养4 h时后，均形成特征性半圆形扩散环；定量分析显示，ΔuvrY突变株的泳动直径较WT下降了33% (****：P<0.000 1)，CΔuvrY未恢复至WT水平(****：P<0.000 1) (图3A、3B)。各菌株在群集平板上均表现出明显的运动性；ΔuvrY的群集运动能力较WT下降了70% (****：P<0.000 1)，而CΔuvrY的群集运动能力恢复至WT水平(图3C、3D)。上述结果证实UvrY对副溶血弧菌的泳动及群集运动具有调控作用。为排除ΔuvrY生长缺陷对运动表型的干扰，同步检测了泳动平板菌体的增殖情况(6 h培养)。经菌落计数验证(图3E)，WT、ΔuvrY和CΔuvrY在运动检测期间的生长能力无统计学差异(P>0.05)，确证运动表型变化是uvrY缺失的直接效应。上述结果表明，作为双组分系统的调控枢纽，UvrY对副溶血弧菌的泳动和群集运动具有差异性调控作用，且该功能独立于其生长调控途径。
2.4 UvrY不参与副溶血弧菌生物被膜的形成
采用结晶紫染色法定量评估WT、ΔuvrY及CΔuvrY的生物被膜形成能力。经48 h静态培养后，染色分析显示(图4A、4B)在30 ℃和37 ℃的培养条件下，ΔuvrY菌株的生物被膜吸光度值与WT组无统计学差异(P>0.05)，提示uvrY基因不参与副溶血弧菌生物被膜的形成。
2.5 UvrY参与副溶血弧菌细菌竞争过程
将各组副溶血弧菌与大肠杆菌进行混合孵育，分别在30 ℃和37 ℃作用4 h后进行计数，考察uvrY基因缺失后对副溶血弧菌杀伤大肠杆菌能力的影响。如图5A所示，在30 ℃时，与大肠杆菌共孵育4 h后ΔuvrY组的大肠杆菌存活数显著高于WT组(***：P<0.001)，表明ΔuvrY缺失株杀伤大肠杆菌的能力显著低于野生株。同时，对共孵育4 h后的各组副溶血弧菌进行计数。结果显示，各组活菌量均无显著差异，排除了由于uvrY基因缺失导致副溶血弧菌生长变慢而引起的杀菌作用减弱的可能性(图5B)。在37 ℃时，各组与大肠杆菌共孵育4 h后大肠杆菌和副溶血弧菌的浓度差异均不显著(图5C、5D)。然而，在30 ℃和37 ℃共孵育4 h后，CΔuvrY组的大肠杆菌存活数未能恢复至WT水平(****：P<0.000 1)。上述结果表明，在30 ℃时uvrY基因有助于副溶血弧菌对大肠杆菌的抗菌作用。
2.6 UvrY参与副溶血弧菌对细胞的黏附过程
以MOI为10的条件感染HeLa细胞1 h，细胞黏附实验结果显示(图6A)：ΔuvrY的黏附水平较WT显著下降36.7% (***：P<0.001)，而CΔuvrY则恢复至WT的细胞黏附水平。该结果证实uvrY基因正向调控副溶血弧菌对宿主细胞的黏附过程。为明确黏附缺陷是否源于细菌增殖差异，实验采用平行对照设计：各菌株以MOI=10:1感染HeLa细胞1 h后裂解细胞，接着连续10倍比稀释后涂布至LB平板上，观察并记录菌落的生长情况。如图6B所示，感染时各菌株活菌浓度无统计学差异(P>0.05)，排除了生长速率对黏附表型的潜在影响。UvrY可能通过调控细菌表面黏附因子的表达，而非影响基础生长特性，来介导副溶血弧菌与宿主细胞的黏附作用。
2.7 uvrY 基因缺失影响副溶血弧菌对细胞的毒性作用
为了研究UvrY是否影响副溶血弧菌的毒力，本研究对感染WT、ΔuvrY和CΔuvrY菌株的HeLa细胞(MOI=10:1，感染1.5 h)的乳酸脱氢酶释放水平进行了定量分析。与WT相比，ΔuvrY诱导的细胞毒性下降了15.8% (**：P<0.01)(图7)，表明UvrY作为关键调控因子能够促进副溶血弧菌对宿主细胞的毒性作用。
2.8 UvrY调控副溶血弧菌对小鼠的致病性作用
本研究利用小鼠腹腔感染模型系统评估了UvrY在哺乳动物感染模型中的毒力调控作用。WT组小鼠在感染2 h后即出现精神萎靡、呼吸急促、眼球充血、粪便稀薄等临床症状，并在10 h内全部死亡(图8)。与WT组相比，ΔuvrY组小鼠的临床症状显著减轻，且死亡起始时间延迟，最终存活率达75%；CΔuvrY组小鼠的症状与WT组相似，存活率为25%；空白对照组小鼠未出现上述临床症状，存活率为100%。该结果不仅证实uvrY基因缺失可显著降低副溶血弧菌对小鼠的致病性，还为BarA/UvrY双组分系统的宿主体内功能研究提供了依据。
2.9 UvrY参与副溶血弧菌在小鼠组织的定殖过程
为了解UvrY对副溶血弧菌在宿主体内定殖的影响，本研究进行了小鼠组织载菌量测定。结果显示(图9)，接种10 h后ΔuvrY组菌株在小鼠心脏、肝脏、脾脏和肾脏的载菌量分别下降为WT组的1/27、1/273、1/54和1/47 (**：P<0.01；****：P<0.000 1)，表明uvrY基因参与副溶血弧菌在小鼠组织中的定殖过程。然而，CΔuvrY在各个组织中的定殖能力未能恢复到WT水平(****：P<0.000 1)。
3 讨论与结论
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UvrY作为BarA/UvrY双组分系统的核心反应调节因子，在多种致病菌的碳代谢、生物被膜形成、运动性和毒力过程中发挥调控作用[13,24-25]。本研究通过构建ΔuvrY突变株系统解析了该调控因子对细菌生长、细菌竞争、运动性、对宿主的黏附和毒性以及致病性的影响，揭示了其在副溶血弧菌感染过程中的关键作用。
在生长特性方面，ΔuvrY表现出独特的表型：在标准培养条件下，短周期培养(6 h内)其生长能力与WT相当，但在后期出现生长迟滞。这一现象提示UvrY可能通过双相调控机制影响代谢通路，缺失uvrY可能阻碍了BarA/UvrY系统对碳代谢的调控，进而限制能量供应。早期通过非碳代谢依赖途径维持基础生长，而在对数生长期后期则通过激活碳代谢相关基因来优化能量分配，导致ΔuvrY在中后期出现生长缺陷。这种时序性调控模式拓展了对BarA/UvrY系统功能的认识。本研究还通过实验证实，在短周期培养下泳动检测(图3E)及细胞黏附实验(图6B)中生长未受影响。
UvrY及其同源蛋白还影响大肠杆菌、沙门氏菌、霍乱弧菌、铜绿假单胞菌和黏质沙雷氏菌等的鞭毛基因表达和/或运动性[24,26-28]。副溶血弧菌通过游动运动在液体中移动，并利用群集运动在表面上移动，这使得细胞能够到达有利的环境。UvrY在副溶血弧菌的环境适应功能中扮演关键角色，uvrY基因的缺失显著降低了副溶血弧菌的泳动和群集运动能力。此外，细菌竞争试验表明UvrY可增强副溶血弧菌在30 ℃条件下的环境生存优势，这是揭示BarA/UvrY双组分系统对抗菌活性具有调控作用。运动能力作为细菌毒力的重要决定因素，结合已有研究[29]，我们推测UvrY可能通过调控鞭毛基因的表达，从而协调运动性与抗菌活性以及致病性的关系，这一发现为理解病原体环境适应机制提供了新视角。
通过比较分析，本研究证实UvrY促进了副溶血弧菌对宿主细胞的黏附和毒性作用。黏附于宿主组织是微生物建立感染的基础，有助于避免被宿主的天然免疫机制清除[30]。BarA/UvrY双组分系统是禽致病性大肠杆菌黏附和毒力的决定因素[31]。Zhang等[13]发现UvrY通过调控T3SS2和T6SS效应蛋白的分泌来促进副溶血弧菌的毒力。本研究表明，uvrY基因的缺失显著削弱了副溶血弧菌对HeLa细胞的黏附能力及细胞毒性作用。值得注意的是，病原菌通过释放毒性因子介导的宿主细胞损伤过程是其突破宿主防御屏障实现体内定殖的重要病理机制。在肠出血性大肠杆菌中，UvrY能够增加T3SS基因的表达，促进其在小鼠组织中的定殖能力[12]。本研究证实UvrY通过多途径调控副溶血弧菌的黏附-定殖过程。小鼠感染实验进一步证实了UvrY在系统性毒力调控中的核心地位。uvrY基因缺失导致临床症状延迟出现以及致死率显著降低，同时，ΔuvrY表现出的黏附率下降与组织载菌量降低存在显著相关性。这些结果提示，UvrY为副溶血弧菌提供了生存优势，导致突变菌株的毒力、黏附和定殖能力下降，使病原菌更易被宿主清除，而WT在组织内持续存在，导致宿主死亡。这表明UvrY可能通过调控副溶血弧菌的系统性毒力因子显著影响其在哺乳动物宿主中的致病进程。
上述结果提示，UvrY是副溶血弧菌重要的毒力因子，参与协调细菌生长、抗菌活性、运动性、毒力、黏附和定殖能力，进而影响副溶血弧菌的生物学特性和对宿主的致病性。这些发现不仅显著深化了对副溶血弧菌致病分子机制的理解，同时也为相关感染的防治开辟了新的思路，使其成为一个极具潜力的抗菌靶标。基于当前研究成果，未来我们将继续深入解析UvrY的精确调控网络，从而为开发针对副溶血弧菌的新型防控策略打下坚实的基础。
基金
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  • 国家自然科学基金(32473039)
  • 上海市自然科学基金(21ZR1477000)
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doi: 10.13343/j.cnki.wsxb.20250180
  • 接收时间：2025-03-06
  • 首发时间：2026-02-07
  • 出版时间：2025-09-04
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  • 收稿日期：2025-03-06
  • 录用日期：2025-04-18
基金
National Natural Science Foundation of China(32473039)
国家自然科学基金(32473039)
Shanghai Natural Science Foundation of China(21ZR1477000)
上海市自然科学基金(21ZR1477000)
作者信息
    1 中国农业科学院上海兽医研究所，上海
    2 上海健康医学院，上海
    3 从化海关综合技术服务中心，广东 广州
    4 浙江农林大学 动物科技学院，浙江 临安
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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
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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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