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Article(id=1259888476493889962, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1259888457367806489, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250898, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1764777600000, receivedDateStr=2025-12-04, revisedDate=null, revisedDateStr=null, acceptedDate=1768752000000, acceptedDateStr=2026-01-19, onlineDate=1778310420392, onlineDateStr=2026-05-09, pubDate=1777824000000, pubDateStr=2026-05-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1778310420392, onlineIssueDateStr=2026-05-09, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1778310420392, creator=13701087609, updateTime=1778310420392, updator=13701087609, issue=Issue{id=1259888457367806489, tenantId=1146029695717560320, journalId=1192105938417971205, year='2026', volume='66', issue='5', pageStart='2031', pageEnd='2556', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=0, articleOrder=1, issueType=-1, specialIssue=null, createTime=1778310415832, creator=13701087609, updateTime=1778320153326, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1259929299465921482, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1259888457367806489, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1259929299465921483, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1259888457367806489, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=2339, endPage=2351, ext={EN=ArticleExt(id=1259888480608502222, articleId=1259888476493889962, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Effects of outer membrane protein AhCirA on antibiotic resistance of Aeromonas hydrophila under quinolone and aminoglycoside stress, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

Objective The outer membrane protein CirA serves as a specific transporter for catecholate-type siderophores and is involved in the uptake of siderophores and other nutrients, playing a crucial role in bacterial physiology. However, its impact on bacterial antibiotic resistance remains unclear. This study aimed to investigate the role of ahcirA in the antibiotic resistance of Aeromonas hydrophila ATCC 7966 under antibiotic stress, thereby providing a theoretical basis for elucidating the molecular mechanism by which ahcirA regulates bacterial resistance. Methods With A. hydrophila ATCC 7966 as the model organism, an ahcirA knockout strain (ΔahcirA) was constructed, and its susceptibility to multiple quinolones and aminoglycosides was assessed. Quantitative proteomics was further employed to compare protein expression profiles of ΔahcirA with and without antibiotic stress. Bioinformatic approaches were adopted for the functional analysis of differentially expressed proteins. Results In the media containing enrofloxacin and norfloxacin, the growth of ΔahcirA was significantly impaired compared with that of the wild-type strain. In contrast, ΔahcirA exhibited enhanced growth in the media supplemented with kanamycin and streptomycin. Proteomic and bioinformatic analyses revealed that the deletion of ahcirA may alter bacterial antibiotic resistance by affecting the expression of proteins involved in multiple biological processes, such as small molecule metabolism, and by modulating the expression of antibiotic resistance genes. Conclusion CirA plays a significant role in the antibiotic resistance of A. hydrophila. Its absence influences bacterial susceptibility to different classes of antibiotics by regulating the expression of diverse functional proteins and antibiotic resistance genes.

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E-mail:
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These authors contributed equally to this work.

, authorsList=Xinrui ZHAO, Yuyue XIE, Yawen HUANG, Yankai LIU, Xiangmin LIN), CN=ArticleExt(id=1259888504893522700, articleId=1259888476493889962, tenantId=1146029695717560320, journalId=1192105938417971205, language=CN, title=喹诺酮类和氨基糖苷类抗生素胁迫下外膜蛋白AhCirA对嗜水气单胞菌耐药性的影响, columnId=1192149544164012138, journalTitle=微生物学报, columnName=研究报告, runingTitle=null, highlight=null, articleAbstract=

目的 外膜蛋白CirA作为特异性儿茶酚酸盐铁载体的转运通道，参与铁载体及其他营养物质的转运过程，在细菌生理功能中发挥着重要作用，但其对细菌耐药性的影响尚不明确。本研究探究ahcirA在抗生素胁迫下对嗜水气单胞菌ATCC 7966耐药机制的影响，为深入解析ahcirA调控细菌耐药性的分子机制提供理论基础。 方法 以嗜水气单胞菌ATCC 7966为研究对象，构建ahcirA基因缺失株(ΔahcirA)，并测定其对多种喹诺酮类和氨基糖苷类抗生素的敏感性。进一步采用定量蛋白质组学技术，比较ΔahcirA菌株在有无抗生素胁迫条件下的蛋白表达差异，并通过生物信息学方法对差异表达蛋白进行功能分析。 结果 在含恩诺沙星和诺氟沙星的培养基中ΔahcirA菌株的生长状况显著低于野生型菌株；而在含卡那霉素和链霉素的培养基中ΔahcirA菌株的生长状况优于野生型菌株。蛋白质组学与生物信息学分析表明，ahcirA缺失可能通过影响与小分子代谢过程等多种生物过程相关的蛋白及耐药基因的表达，从而改变细菌的耐药性。 结论 外膜蛋白CirA在嗜水气单胞菌耐药性中具有重要作用，其缺失可通过调控多种功能蛋白及耐药基因的表达影响细菌对不同类型抗生素的敏感性。

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作者贡献声明

赵心锐：实验设计与验证、数据整理、原稿撰写与修改；谢于悦：实验设计与验证、方法学、原稿撰写；黄雅雯：概念化、方法学、数据整理；刘彦楷：原稿撰写；林向民：概念化、监督、项目管理、稿件审阅与编辑。

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2.Key Provincial Key Laboratory of Agroecological Processing and Safety Monitoring, Fujian Agriculture and Forestry University, Fuzhou, Fujian, China
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2.福建农林大学，福建省农业生态过程与安全监控重点实验室，福建 福州
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2.Key Provincial Key Laboratory of Agroecological Processing and Safety Monitoring, Fujian Agriculture and Forestry University, Fuzhou, Fujian, China
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2.福建农林大学，福建省农业生态过程与安全监控重点实验室，福建 福州
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2.Key Provincial Key Laboratory of Agroecological Processing and Safety Monitoring, Fujian Agriculture and Forestry University, Fuzhou, Fujian, China
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A-D: Show the spot assay results of WT and ΔahcirA strains after 3 h of culture in media supplemented with the following antibiotics at the indicated final concentrations: 0.02 μg/mL ENR, 0.02 μg/mL NOR, 5 μg/mL SM, and 10 μg/mL KAN, respectively., figureFileSmall=iDMTYIoaOZCTNvSJg0GMqA==, figureFileBig=F8s6Dmv4t0mnb9WdUaRWMw==, tableContent=null), ArticleFig(id=1259928474517648125, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1259888476493889962, language=CN, label=图1, caption=野生菌和ΔahcirA 菌株的抗生素敏感性测定结果, figureFileSmall=iDMTYIoaOZCTNvSJg0GMqA==, figureFileBig=F8s6Dmv4t0mnb9WdUaRWMw==, tableContent=null), ArticleFig(id=1259928478665814793, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1259888476493889962, language=EN, label=Figure 2, caption=Growth curves of ΔahcirA under different concentrations of antibiotics stress. 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A: Principal component analysis (PCA) plot (Each color represents a distinct sample, with triplicate data points of the same color indicating three biological replicates); B: Density plot showing the distribution of protein expression ratios across different groups; C: Statistical summary of the number of DEPs in each group compared to the control; D: Distribution of differential proteins across groups., figureFileSmall=Vq/KDc4/Mlg0AuIAA2SVWQ==, figureFileBig=m9DlhvrmkO4YRC5OsNcgEw==, tableContent=null), ArticleFig(id=1259928484911133477, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1259888476493889962, language=CN, label=图3, caption=定量蛋白质组学数据分析, figureFileSmall=Vq/KDc4/Mlg0AuIAA2SVWQ==, figureFileBig=m9DlhvrmkO4YRC5OsNcgEw==, tableContent=null), ArticleFig(id=1259928487834563378, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1259888476493889962, language=EN, label=Figure 4, caption=GO enrichment analysis. A: Biological processes; B: Molecular functions. The circles represent the number of differentially expressed proteins involved in the relevant pathways, and the colored squares indicate the -lg P value., figureFileSmall=wiCThoxHKGJ6UAuDgxwduw==, figureFileBig=fvZoqCr5tIJA/fq0b1H8xg==, tableContent=null), ArticleFig(id=1259928490040767293, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1259888476493889962, language=CN, label=图4, caption=GO富集分析, figureFileSmall=wiCThoxHKGJ6UAuDgxwduw==, figureFileBig=fvZoqCr5tIJA/fq0b1H8xg==, tableContent=null), ArticleFig(id=1259928492716733255, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1259888476493889962, language=EN, label=Figure 5, caption=The upset plot illustrating the overlap of differentially expressed proteins among different groups. Bottom yellow bars: Total differential proteins per comparison; Top green bars: Group intersections (Numbers show overlap count). Central matrix: Single points denote group‑unique elements; Connecting lines indicate shared elements., figureFileSmall=/p5IyRWRNMqwe4A7J5+5JQ==, figureFileBig=mjt7U627weygTwH1zqyN3A==, tableContent=null), ArticleFig(id=1259928494251848532, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1259888476493889962, language=CN, label=图5, caption=Upset图表示不同组之间差异蛋白的重叠, figureFileSmall=/p5IyRWRNMqwe4A7J5+5JQ==, figureFileBig=mjt7U627weygTwH1zqyN3A==, tableContent=null), ArticleFig(id=1259928496843928414, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1259888476493889962, language=EN, label=Figure 6, caption=Screening of antibiotic resistance genes., figureFileSmall=2yCWb0ykmdv2uc6TDFgqCw==, figureFileBig=/WzkPvwCTzjzfmVGAFFWkA==, tableContent=null), ArticleFig(id=1259928497519211370, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1259888476493889962, language=CN, label=图6, caption=耐药基因筛选, figureFileSmall=2yCWb0ykmdv2uc6TDFgqCw==, figureFileBig=/WzkPvwCTzjzfmVGAFFWkA==, tableContent=null), ArticleFig(id=1259928498630701940, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1259888476493889962, language=EN, label=Figure 7, caption=qPCR verification results of differentially expressed genes of ΔahcirA strains under different antibiotic stresses. A-D: Show the up- and down-regulation of six randomly selected drug-resistance genes under treatment with streptomycin, kanamycin, enrofloxacin, and norfloxacin, respectively., figureFileSmall=gdkUuqBPnCaUf41YNqeOYQ==, figureFileBig=0VdeTA9Ef8wo2GtWpdtpTw==, tableContent=null), ArticleFig(id=1259928500975317892, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1259888476493889962, language=CN, label=图7, caption=qPCR验证ΔahcirA 菌株在不同抗生素胁迫下的基因表达情况, figureFileSmall=gdkUuqBPnCaUf41YNqeOYQ==, figureFileBig=0VdeTA9Ef8wo2GtWpdtpTw==, tableContent=null), ArticleFig(id=1259928502514627469, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1259888476493889962, language=EN, label=Table 1, caption=

The fluorescence quantitative primers of PCR

, figureFileSmall=null, figureFileBig=null, tableContent=
Gene namesSequences (5′→3′)
ppsA-FGTGGTGGGTTGTGGTGATGC
ppsA-RGGCGAAGTCGAACGCTCTGT
AHA_1804-FGGGGAAGAGTTCAGCAAGCG
AHA_1804-RCAGCCAGATGACGAGAGGCA
AHA_3092-FCTGGTTTCCGTTCGCAACAT
AHA_3092-RTGCACCCGGCCATACATCTC
AHA_2989-FCGCTGATCGCCCAGAACAAC
AHA_2989-RCGCCTCCTGCCTGAACATCT
AHA_1647-FGGCGTCCCCCGACTATTTGC
AHA_1647-RAAAGCGGTCCCCCTGCTCCT
AHA_4253-FCCGTTTGCTGATTATTTTCC
AHA_4253-RGAGTCGCAGTCGAGCTTGTT
cysA-FTCGGGTGGTGCTGATGAACG
cysA-RTGGGTAGCAGGCGGGTGATG
AHA_4258-FCGGCTGCTTTCTCTATGCTG
AHA_4258-RGACCACCGAGTAGGTTTCCC
AHA_0854-FCCCCACTTCTTGGACAAAAG
AHA_0854-RCTTCGTAGCCATCCATCTTC
), ArticleFig(id=1259928505106707352, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1259888476493889962, language=CN, label=表1, caption=

荧光定量PCR引物

, figureFileSmall=null, figureFileBig=null, tableContent=
Gene namesSequences (5′→3′)
ppsA-FGTGGTGGGTTGTGGTGATGC
ppsA-RGGCGAAGTCGAACGCTCTGT
AHA_1804-FGGGGAAGAGTTCAGCAAGCG
AHA_1804-RCAGCCAGATGACGAGAGGCA
AHA_3092-FCTGGTTTCCGTTCGCAACAT
AHA_3092-RTGCACCCGGCCATACATCTC
AHA_2989-FCGCTGATCGCCCAGAACAAC
AHA_2989-RCGCCTCCTGCCTGAACATCT
AHA_1647-FGGCGTCCCCCGACTATTTGC
AHA_1647-RAAAGCGGTCCCCCTGCTCCT
AHA_4253-FCCGTTTGCTGATTATTTTCC
AHA_4253-RGAGTCGCAGTCGAGCTTGTT
cysA-FTCGGGTGGTGCTGATGAACG
cysA-RTGGGTAGCAGGCGGGTGATG
AHA_4258-FCGGCTGCTTTCTCTATGCTG
AHA_4258-RGACCACCGAGTAGGTTTCCC
AHA_0854-FCCCCACTTCTTGGACAAAAG
AHA_0854-RCTTCGTAGCCATCCATCTTC
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喹诺酮类和氨基糖苷类抗生素胁迫下外膜蛋白AhCirA对嗜水气单胞菌耐药性的影响
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赵心锐 1, 2, 3 , 谢于悦 1, 2, 3 , 黄雅雯 1, 2, 3 , 刘彦楷 1, 2, 3 , 林向民 1, 2, 3
微生物学报 | 研究报告 2026,66(5): 2339-2351
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微生物学报 | 研究报告 2026, 66(5): 2339-2351
喹诺酮类和氨基糖苷类抗生素胁迫下外膜蛋白AhCirA对嗜水气单胞菌耐药性的影响
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赵心锐1, 2, 3, 谢于悦1, 2, 3, 黄雅雯1, 2, 3, 刘彦楷1, 2, 3, 林向民1, 2, 3
作者信息
  • 1.福建农林大学 生命科学学院，福建 福州
  • 2.福建农林大学，福建省农业生态过程与安全监控重点实验室，福建 福州
  • 3.福建农林大学，作物生态与分子生理重点实验室，福建 福州
Effects of outer membrane protein AhCirA on antibiotic resistance of Aeromonas hydrophila under quinolone and aminoglycoside stress
Xinrui ZHAO1, 2, 3, Yuyue XIE1, 2, 3, Yawen HUANG1, 2, 3, Yankai LIU1, 2, 3, Xiangmin LIN1, 2, 3
Affiliations
  • 1.College of Life Sciences, Fujian Agriculture and Forestry University, Fuzhou, Fujian, China
  • 2.Key Provincial Key Laboratory of Agroecological Processing and Safety Monitoring, Fujian Agriculture and Forestry University, Fuzhou, Fujian, China
  • 3.Key Laboratory of Crop Ecology and Molecular Physiology, Fujian Agriculture and Forestry University, Fuzhou, Fujian, China
出版时间: 2026-05-04 doi: 10.13343/j.cnki.wsxb.20250898
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摘要
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目的 外膜蛋白CirA作为特异性儿茶酚酸盐铁载体的转运通道，参与铁载体及其他营养物质的转运过程，在细菌生理功能中发挥着重要作用，但其对细菌耐药性的影响尚不明确。本研究探究ahcirA在抗生素胁迫下对嗜水气单胞菌ATCC 7966耐药机制的影响，为深入解析ahcirA调控细菌耐药性的分子机制提供理论基础。 方法 以嗜水气单胞菌ATCC 7966为研究对象，构建ahcirA基因缺失株(ΔahcirA)，并测定其对多种喹诺酮类和氨基糖苷类抗生素的敏感性。进一步采用定量蛋白质组学技术，比较ΔahcirA菌株在有无抗生素胁迫条件下的蛋白表达差异，并通过生物信息学方法对差异表达蛋白进行功能分析。 结果 在含恩诺沙星和诺氟沙星的培养基中ΔahcirA菌株的生长状况显著低于野生型菌株；而在含卡那霉素和链霉素的培养基中ΔahcirA菌株的生长状况优于野生型菌株。蛋白质组学与生物信息学分析表明，ahcirA缺失可能通过影响与小分子代谢过程等多种生物过程相关的蛋白及耐药基因的表达，从而改变细菌的耐药性。 结论 外膜蛋白CirA在嗜水气单胞菌耐药性中具有重要作用，其缺失可通过调控多种功能蛋白及耐药基因的表达影响细菌对不同类型抗生素的敏感性。

嗜水气单胞菌  /  外膜蛋白AhCirA  /  喹诺酮类  /  氨基糖苷类  /  耐药性

Objective The outer membrane protein CirA serves as a specific transporter for catecholate-type siderophores and is involved in the uptake of siderophores and other nutrients, playing a crucial role in bacterial physiology. However, its impact on bacterial antibiotic resistance remains unclear. This study aimed to investigate the role of ahcirA in the antibiotic resistance of Aeromonas hydrophila ATCC 7966 under antibiotic stress, thereby providing a theoretical basis for elucidating the molecular mechanism by which ahcirA regulates bacterial resistance. Methods With A. hydrophila ATCC 7966 as the model organism, an ahcirA knockout strain (ΔahcirA) was constructed, and its susceptibility to multiple quinolones and aminoglycosides was assessed. Quantitative proteomics was further employed to compare protein expression profiles of ΔahcirA with and without antibiotic stress. Bioinformatic approaches were adopted for the functional analysis of differentially expressed proteins. Results In the media containing enrofloxacin and norfloxacin, the growth of ΔahcirA was significantly impaired compared with that of the wild-type strain. In contrast, ΔahcirA exhibited enhanced growth in the media supplemented with kanamycin and streptomycin. Proteomic and bioinformatic analyses revealed that the deletion of ahcirA may alter bacterial antibiotic resistance by affecting the expression of proteins involved in multiple biological processes, such as small molecule metabolism, and by modulating the expression of antibiotic resistance genes. Conclusion CirA plays a significant role in the antibiotic resistance of A. hydrophila. Its absence influences bacterial susceptibility to different classes of antibiotics by regulating the expression of diverse functional proteins and antibiotic resistance genes.

Aeromonas hydrophila  /  outer membrane protein AhCirA  /  quinolones  /  aminoglycosides  /  antibiotic resistance
赵心锐, 谢于悦, 黄雅雯, 刘彦楷, 林向民. 喹诺酮类和氨基糖苷类抗生素胁迫下外膜蛋白AhCirA对嗜水气单胞菌耐药性的影响. 微生物学报, 2026 , 66 (5) : 2339 -2351 . DOI: 10.13343/j.cnki.wsxb.20250898
Xinrui ZHAO, Yuyue XIE, Yawen HUANG, Yankai LIU, Xiangmin LIN. Effects of outer membrane protein AhCirA on antibiotic resistance of Aeromonas hydrophila under quinolone and aminoglycoside stress[J]. Acta Microbiologica Sinica, 2026 , 66 (5) : 2339 -2351 . DOI: 10.13343/j.cnki.wsxb.20250898
正文
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嗜水气单胞菌(Aeromonas hydrophila)是一种广泛分布于多种水体的革兰氏阴性菌，属于气单胞菌属[1-2]。该菌是自然水域中常见的条件致病菌，当水体环境或宿主体内环境发生剧烈变化时可引发鱼类患出血性败血症，还能同时感染人类、鱼类及两栖动物等多种宿主[3-4]。细菌外膜在维持细胞生存与致病过程中起着关键作用，它既能提供有效保护，又允许生命活动所必需的物质进行交换[5]。目前，抗生素的作用机制大多依赖于进入细胞内干扰生物过程，因此外膜蛋白直接影响细菌对抗生素的敏感性[6-7]。
Colicin I receptor (CirA)是嗜水气单胞菌中的一种外膜蛋白，同时也是一种重要的铁载体转运蛋白[8]。它能够在外膜形成特异性通道，负责将儿茶酚酸盐类铁载体从胞外转运至周质空间，该过程依赖TonB系统提供能量；随后，铁载体进一步通过内膜上的ABC转运蛋白进入胞内，从而参与维持细胞内的铁稳态[9]。除铁载体转运功能外，CirA在细菌毒力、生物膜形成以及抗生素耐药性等方面也发挥着重要作用。例如，肠道沙门氏菌中cirA基因的缺失会导致生物膜形成能力显著下降，并削弱抗生素耐药性[10]。多项研究还表明，在抗生素胁迫、铁限制、氧化应激及胆盐应激等外界压力条件下，多种细菌中CirA同源物的表达会显著上调[11]。
尽管已有研究显示嗜水气单胞菌的耐药性问题日趋严峻，且对其部分耐药机制有所阐明，然而其内在分子机制仍存在诸多未解之处，有待进一步深入探索[12-13]。值得注意的是，喹诺酮类和氨基糖苷类抗生素可通过降低膜通透性影响细菌耐药性，是两类常用的抗菌药物[14]。AhCirA作为嗜水气单胞菌中关键的铁载体转运蛋白和主要外膜通道蛋白，不仅影响生物膜形成和细菌运动能力，还可能参与调控多种抗生素耐受性。
本研究旨在探究AhCirA蛋白在嗜水气单胞菌耐药性形成中的作用机制。通过定量蛋白质组学方法分析ΔahcirA在多种抗生素胁迫下的蛋白表达谱，可系统比较该缺失菌株与野生型菌株的响应差异。进一步结合生物信息学分析，有助于揭示差异表达蛋白所涉及的生物过程与分子功能，从而为阐明AhCirA是否通过调控外膜通透性等机制影响细菌耐药性提供线索。该研究可为深入理解嗜水气单胞菌的耐药机制提供新视角，并为后续防控策略及新型抗菌药物的研发奠定理论基础。
1 材料与方法
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1.1 材料
本研究所用菌株包括嗜水气单胞菌野生型ATCC 7966、课题组先前研究中构建的ahcirA基因缺失菌株(ΔahcirA)[15]和ΔahcirA基因回补菌株[16]。前期已对ΔahcirA突变株和回补株的基础生理表型进行了系统性验证，实验结果显示ΔahcirA菌株回补后其生理功能也得到恢复[16]。
1.2 抗生素抗性测定
本研究选用2种喹诺酮类抗生素(恩诺沙星、诺氟沙星)和2种氨基糖苷类抗生素(卡那霉素、链霉素)进行ahcirA基因的抗生素敏感性检测。配制含有不同终浓度抗生素的LB液体培养基，将4种待测菌株接种后，于30 ℃、200 r/min条件下培养至OD600为1.0。将菌液进行10倍梯度稀释(共8个梯度)，使用多通道移液器将各菌株不同稀释度的样品点至普通LB平板上，30 ℃倒置培养12 h后拍照记录。
1.3 最低抑菌浓度和生长曲线测定
采用二倍稀释法，用LB培养基对抗生素进行梯度稀释。将活化过夜的菌液稀释后，并分别加入含有抗生素的LB培养基中，以不含抗生素的LB培养基作为阴性对照。然后，于37 ℃静置培养16 h，记录最低抑菌浓度(minimum inhibitory concentration, MIC)值。根据测得的ΔahcirA菌株对卡那霉素(kanamycin, KAN)、链霉素(streptomycin, SM)、恩诺沙星(enrofloxacin, ENR)和诺氟沙星(norfloxacin, NOR)的MIC值，用LB培养基对上述抗生素进行梯度稀释，浓度梯度为1/8、1/4、1/2 MIC。将培养至稳定状态的ΔahcirA菌株以1:100的比例分别转入添加/不添加抗生素的LB培养基中，再转移至生长曲线板中，每个样品重复3次，最后使用全自动生长曲线分析仪于30 ℃下连续监测16 h，每小时记录1次OD600值。
1.4 蛋白质样品制备
将过夜菌以1:100比例转接到50 mL的LB液体培养基中，30 ℃、200 r/min培养至OD600=1.0时，4 ℃、12 000 r/min离心2 min收集所有菌体，用预冷的PBS溶液洗涤2次，再于4 ℃、8 000 r/min离心10 min。接着加入1 mL的Lysis buffer [6 mol/L尿素和2 mol/L硫脲溶解于0.1 mol/L的Tris-HCl (pH 7.6)溶液，并加入蛋白酶抑制剂]混匀，置于冰上，在超声破碎仪(30%功率)中超声破碎15 min至菌液呈现透明，4 ℃、8 000 r/min离心15 min，然后将上清液转移到新的离心管中测定蛋白浓度。每组约50 μg蛋白质样品，在56 ℃下用10 mmol/L二硫苏糖醇(dithiothreitol, DTT)还原40 min，然后在室温黑暗环境中使用50 mmol/L碘乙酰胺(iodoacetamide, IAA)烷基化30 min。接着加入胰蛋白酶(1:50)，在37 ℃下过夜酶解以获得多肽样品。然后用C18柱对得到的肽进行脱盐，并进行质谱鉴定。
1.5 质谱鉴定
脱盐后的肽样品使用EASY-nano-LC系统，以600 nL/min的流速进行分离。分离梯度设置为：0-12 min，10%-14%的流动相B (乙腈含0.1%的FA)；12-57 min，14%-26%的流动相B，随后在10 min内将流动相B的比例从26%上升至42%。利用Q Exactive HF质谱仪(ThermoFisher Scientific公司)进行数据非依赖性采集(data-independent aquisition, DIA)数据库构建与分析。全扫描设置为350-1 500 m/z范围内，分辨率60 000；DIA扫描分辨率设置为30 000；NCE：28%；AGC目标：3×106，最大注入时间为auto。为DIA采集设置了45个可变DIA窗口。将通过质谱收集的原始DIA数据与使用Maxquant软件版本v.1.6.3.4的数据库进行比较，该数据库包含嗜水气单胞菌ATCC 7966的蛋白质组序列。在肽和蛋白质水平上的错误发现率(false discovery rate, FDR)设置为1%。
1.6 生物信息学分析
生物信息学分析使用David (https://david.ncifcrf.gov/)在线数据库对差异蛋白(differentially expressed proteins, DEPs)进行基因本体论(gene ontology, GO)富集分析。此外，本研究还使用TBtools软件[17]进行相关性分析及UpSet分析，综合抗生素耐药数据库CARD在线网站(https://card.mcmaster.ca/)预测耐药基因，然后使用Cytoscape 3.9.1进行可视化。
1.7 RNA提取
将过夜菌转接培养至OD600=1.0，预冷离心机后，以5 000×g离心10 min收集菌体，加入1 mL TRIzol，充分吹打混匀，静置2-3 min。氯仿抽提：加入200 µL氯仿(所有试剂全部提前预冷)，涡旋振荡1 min充分反应至淡粉色、微黏稠，室温静置5 min，4 ℃、10 000 r/min离心15 min，取上清并移至新的RNA free 1.5 mL EP管中(约为600 µL)，加入体积分数为60%预冷的异丙酮(约为480 µL)，涡旋振荡1 min充分反应，室温静置5-10 min；洗涤：4 ℃、10 000 r/min离心15 min，弃上清，留沉淀(尽量吸干)加入75%乙醇1 mL (无水乙醇750 mL+DEPC水250 µL，现用现配)，轻轻吹打混匀，4 ℃、7 500 r/min离心15 min，重复1次，4 ℃、7 500 r/min离心15 min，弃上清，留沉淀(将残留液体吸净)，无菌台吹3-5 min至无乙醇味；加入30 µL的DEPC水，溶解沉淀，置于冰上测定其浓度。按照反转录试剂盒说明书(TaKaRa公司)进行反转录获得cDNA。
1.8 实时荧光定量PCR
采用TRIzol-氯仿法提取细菌总RNA，并逆转录成cDNA作为实时荧光定量PCR (quantitative real-time PCR, qPCR)模板。使用SYBR Premix Ex Taq II (TaKaRa公司)试剂，在CFX96 Touch Deep Well实时荧光定量PCR系统(Bio-Rad公司)上检测目标基因(表1)的相对表达水平。以16S rRNA基因作为内参，反应体系(25 μL)：2×Phanta Max Mix (p515) 12.5 µL，上、下游引物(10 µmol/L)各1 µL，DNA模板0.5 µL，ddH2O 10 µL。PCR反应条件：95 ℃预变性5 min；95 ℃变性30 s，60 ℃退火30 s，72 ℃延伸2 min，共35个循环；72 ℃终延伸5 min。反应结束后，对扩增曲线与溶解曲线进行质量评估。基因相对表达量采用2-ΔΔCt法计算，并使用TBtools (v2.109)软件进行数据分析与可视化。
2 结果与分析
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2.1 ahcirA 缺失菌株对嗜水气单胞菌抗生素耐药性的影响
将含有不同终浓度的恩诺沙星、诺氟沙星、卡那霉素和链霉素的液体LB培养基，按1%的比例接种野生型菌株(wild-type strain, WT)和ΔahcirA菌株，于30 ℃培养3 h，利用稀释点板法分别将上述培养好的WT和ΔahcirA菌株进行10倍稀释，并接种到普通LB平板上。结果如图1所示，在正常液体LB培养基中培养3 h后，WT和ΔahcirA菌株在平板上的生长状况无差异；在诺氟沙星培养基中培养3 h后，ΔahcirA菌株的生长状况明显低于野生菌株，恩诺沙星抗性测定中也出现同样情况(图1A、1B)，而在卡那霉素和链霉素培养基中培养3 h并稀释点板的结果则相反，缺失菌株的生长状况优于野生菌株，如图1C、1D所示。
2.2 ΔahcirA 菌株对SM、KAN、ENR和NOR的MIC值和生长曲线
根据ΔahcirA菌株在不同浓度的SM、KAN、ENR和NOR下的生长情况，测得其对SM、KAN、ENR和NOR的MIC值分别为20、3.12、0.031 2、0.05 μg/mL。依据测定的MIC值，利用生长曲线仪测定ΔahcirA菌株分别在1/8、1/4、1/2 MIC的SM、KAN、ENR和NOR条件下的生长曲线。如图2A-2D所示，当SM、KAN、ENR和NOR的浓度分别为5、0.781 25、0.007 8、0.012 5 μg/mL时，与未加抗生素处理的对照相比，加抗生素处理的ΔahcirA菌株的生长受到显著抑制。因此，本研究分别选择5 μg/mL SM、0.781 25 μg/mL KAN、0.007 8 μg/mL ENR和0.012 5 μg/mL NOR作为处理浓度进行后续研究。
2.3 ΔahcirA 在抗生素胁迫下的蛋白表达差异
为全面了解抗生素对ΔahcirA菌株生理功能的影响，本研究分别用SM、KAN、ENR和NOR处理ΔahcirA菌株后进行蛋白质组学分析。通过PCA主成分分析(图3A)结果显示，同一处理组内的生物学重复紧密聚集，表明实验具有良好的重复性。然而，当将抗生素处理的缺失样本与ΔahcirA进行比较时组间表现出更大的分离趋势，且同一类抗生素的分散性相近，表明同类型抗生素存在一定程度上的系统性差异。此外，密度图显示抗生素处理的缺失样本与ΔahcirA菌株之间的蛋白质比率分布无显著差异(图3B)。随后，采用P值<0.05，且变化倍数>1.5倍的标准来鉴定差异蛋白。与ΔahcirA相比，ΔahcirA在SM处理下共鉴定出522个差异蛋白，其中209个蛋白表达上调，313个蛋白表达下调；同样，ΔahcirA在KAN处理下共鉴定出678个差异表达蛋白，包括251个上调蛋白和427个下调蛋白；ΔahcirA在ENR处理下共鉴定出480个差异蛋白，其中125个蛋白上调，355个蛋白下调；ΔahcirA在NOR处理下有614个差异蛋白，包括235个上调蛋白和379个下调蛋白(图3C)。此外，通过火山图(图3D)可以直观了解这些差异蛋白的分布情况。
2.4 差异表达蛋白的代谢途径分析
本研究通过GO富集分析了解抗生素胁迫下差异表达蛋白对代谢途径的富集情况。结果显示，在被显著富集的前10个生物过程中，卡那霉素和链霉素胁迫组中有8个是完全相同的生物过程，分别是α-氨基酸代谢过程、小分子代谢过程、蛋白质源性氨基酸代谢过程、氨基酸代谢过程、L-氨基酸代谢过程、氧酸代谢过程、有机酸代谢过程和羧酸代谢过程；恩诺沙星和诺氟沙星胁迫组中有6个是完全相同的，分别是氨基酸代谢过程、小分子代谢过程、细胞分解代谢过程、有机酸分解代谢过程、芳香氨基酸家族的分解代谢过程和小分子分解代谢过程。可见在相同类型抗生素胁迫下的差异表达蛋白所富集到的生物过程高度重合，所富集到的分子功能也较为类似。此外，在这些抗生素处理组中DEPs的生物过程富集程度最高的途径是小分子代谢过程，其富集程度显著，P值较低。同样，在分子功能中小分子结合代谢途径也显示出大量富集的DEPs (图4)。总的来说，抗生素胁迫下影响了ΔahcirA菌株的生物功能和各种代谢途径。
2.5 差异表达蛋白分析
根据蛋白质组学结果比较处理组和对照组之间的差异蛋白。图5中的重叠图表明，KAN和NOR处理下的差异蛋白数量最多。此外，在不同抗生素胁迫下，各组共同差异表达上调和下调的蛋白分别有12个和126个。这12个共同差异表达上调的蛋白分别是SecD-2、RpoH、TrmA、AHA_0044、AHA_3652、AHA_4271、AHA_0458、TrpS-1、AHA_3005、AHA_0035、DeoD-2、Tpx，这表明抗生素胁迫下的调控模块具有一定的相似模式。
2.6 耐药基因
本研究还将差异蛋白的氨基酸序列与CARD数据库进行比对，以确定本研究中已知的抗性基因(antibiotic resistance gene, ARGs)蛋白。如图6所示，共鉴定出93个ARGs，其中30个耐药基因为卡那霉素组和链霉素组所共有，19个耐药基因为恩诺沙星组和诺氟沙星组所共有，而6个耐药基因为卡那霉素组、链霉素组、恩诺沙星组和诺氟沙星组所共有，这6个耐药基因分别是ppsA (phosphoenolpyruvate synthase)、AHA_1804 (major facilitator family transporter)、AHA_3092 (histidine kinase)、AHA_1684 (histidine ABC transporter, ATP-binding protein)、AHA_3291 (DNA-binding response regulator)和AHA_3230 (glycerate dehydrogenase)。
2.7 qPCR验证相关基因
为验证抗生素胁迫下ΔahcirA菌株差异表达蛋白结果的准确性，本研究在4种抗生素胁迫下的差异表达蛋白中随机各挑选6个耐药基因，利用qPCR技术验证其表达量。结果如图7所示，在卡那霉素或链霉素胁迫下，AHA_1804、AHA_3092、ppsA均表达下调，AHA_2989、AHA_1647、AHA_4253均表达上调(图7A、7B)；在恩诺沙星或诺氟沙星胁迫下，AHA_1804、AHA_3092、ppsA均表达下调，cysA、AHA_4258、AHA_0854均表达上调(图7C、7D)，转录结果与定量蛋白质组学数据基本一致。
3 讨论
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AhCirA蛋白是嗜水气单胞菌中重要的铁载体转运蛋白，同时也是大肠杆菌素I的受体[18-19]。先前研究发现作为外膜重要的通道蛋白，AhCirA还是多功能蛋白，对嗜水气单胞菌的铁摄取、生长繁殖、生物膜形成以及营养物质的跨外膜运输均具有显著影响[16]。然而，AhCirA是否直接参与嗜水气单胞菌的耐药机制，目前尚未有系统研究。本研究通过系统评估ΔahcirA缺失株在喹诺酮类(恩诺沙星、诺氟沙星)和氨基糖苷类(卡那霉素、链霉素)抗生素胁迫下的表型变化及蛋白质组响应，初步揭示AhCirA在调控嗜水气单胞菌耐药性中的潜在作用。
研究结果显示，ΔahcirA菌株对喹诺酮类抗生素的敏感性增强，而对氨基糖苷类抗生素的耐受性反而提高，表明AhCirA对不同类型抗生素的耐药性具有差异性调控作用。这一现象可能与AhCirA作为外膜蛋白对不同抗生素的渗透性调控有关。李永慧等[20]在大肠杆菌中的研究也发现，cirA基因缺失会导致菌株对多种抗生素的敏感性发生变化，提示CirA同源物在细菌耐药性中具有保守功能。
为进一步探究其分子机制，本研究采用定量蛋白质组学技术，并分析ΔahcirA菌株在4种抗生素胁迫下的蛋白表达谱。PCA分析显示，同一类型抗生素处理下蛋白表达模式高度相似，说明AhCirA可能通过调控一组共同的代谢通路来影响细菌的耐药性。GO富集分析进一步表明，差异表达蛋白主要富集于小分子代谢过程、氨基酸代谢过程及相关分解代谢途径，提示AhCirA缺失可能通过干扰细胞的基础代谢状态，间接影响细菌对抗生素的应激反应。此外，通过CARD数据库比对，本研究共鉴定出93个与抗生素耐药性相关的基因，其中多个基因在氨基糖苷类和喹诺酮类抗生素处理下共同表达。例如，ppsA、AHA_1804、AHA_3092等6个基因在4组处理中均出现差异表达，值得注意的是，AHA_1804 (主要易化子超家族转运蛋白)作为一种多药外排泵，在结核分枝杆菌等临床病原体的耐药性形成中起关键作用[21]。上述结果表明，这些共同响应的基因可能构成AhCirA调控嗜水气单胞菌耐药性的核心靶标群。qPCR验证结果进一步证实了蛋白质组数据的可靠性，表明AhCirA缺失确实影响了这些耐药相关基因的转录水平。
综上所述，AhCirA可能通过调控外膜通透性、代谢重编程以及耐药基因表达等多重机制影响嗜水气单胞菌对喹诺酮类和氨基糖苷类抗生素的敏感性。本研究为深入解析AhCirA在细菌耐药性中的功能提供了新的实验依据和理论支持。同时需要指出，本研究在抗生素种类覆盖上存在一定局限性，AhCirA可能同样参与其他类别抗生素的耐药机制，这将是未来研究的重要方向。
4 结论
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本研究以嗜水气单胞菌ATCC 7966为研究对象，系统分析ahcirA在喹诺酮类和氨基糖苷类抗生素胁迫下的功能。研究发现ahcirA缺失显著增强菌株对恩诺沙星和诺氟沙星的敏感性，但提高对卡那霉素和链霉素的耐受性，说明ahcirA对不同类型抗生素的耐药性具有特异性调控作用。蛋白质组学分析显示，ΔahcirA菌株在抗生素胁迫下出现大量差异表达蛋白，主要富集于小分子代谢、氨基酸代谢等生物过程，提示AhCirA可能通过影响细胞代谢状态参与耐药调控。此外，通过CARD数据库筛选及qPCR验证，鉴定出多个共同响应不同类型抗生素的耐药基因，如ppsA、AHA_1804、AHA_3092等，这些基因可能构成ahcirA调控网络的核心组成部分。本研究从蛋白质组水平揭示了AhCirA在嗜水气单胞菌抗生素耐药性中的多重调控功能，为理解外膜蛋白在细菌耐药机制中的作用提供了新视角，也为后续开发针对嗜水气单胞菌的新型抗菌策略奠定了理论基础。
基金
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  • 国家级大学生创新创业训练计划(202510389041)
  • 国家级大学生创新创业训练计划(202510389036)
文献
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2026年第66卷第5期
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doi: 10.13343/j.cnki.wsxb.20250898
  • 接收时间：2025-12-04
  • 首发时间：2026-05-09
  • 出版时间：2026-05-04
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出版历史
  • 收稿日期：2025-12-04
  • 录用日期：2026-01-19
基金
The National College Students Innovation Training Program(202510389041)
国家级大学生创新创业训练计划(202510389041)
The National College Students Innovation Training Program(202510389036)
国家级大学生创新创业训练计划(202510389036)
作者信息
    1.福建农林大学 生命科学学院，福建 福州
    2.福建农林大学，福建省农业生态过程与安全监控重点实验室，福建 福州
    3.福建农林大学，作物生态与分子生理重点实验室，福建 福州
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https://castjournals.cast.org.cn/joweb/wswxb/CN/10.13343/j.cnki.wsxb.20250898
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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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