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Article(id=1226136784468357386, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226136782408954119, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250672, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1756828800000, receivedDateStr=2025-09-03, revisedDate=null, revisedDateStr=null, acceptedDate=1760112000000, acceptedDateStr=2025-10-11, onlineDate=1770263389937, onlineDateStr=2026-02-05, pubDate=1770134400000, pubDateStr=2026-02-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1770263389937, onlineIssueDateStr=2026-02-05, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1770263389937, creator=13701087609, updateTime=1770263389937, updator=13701087609, issue=Issue{id=1226136782408954119, tenantId=1146029695717560320, journalId=1192105938417971205, year='2026', volume='66', issue='2', pageStart='481', pageEnd='955', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1770263389446, creator=13701087609, updateTime=1770268138976, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1226156703490683529, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226136782408954119, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1226156703490683530, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226136782408954119, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=801, endPage=814, ext={EN=ArticleExt(id=1226136784753570063, articleId=1226136784468357386, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Impacts of EvpP on the biological characteristics of Edwardsiella tarda and infection of macrophages, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

[Objective] The type VI secretion system (T6SS) is a novel virulence factor of Edwardsiella tarda. E. tarda virulence protein P (EvpP) is an effector of the T6SS. To date, the functional mechanisms of EvpP are still poorly understood. This study aimed to comprehensively investigate the biological functions of EvpP and elucidate the roles of T6SS in the pathogenicity of E. tarda. [Methods] We constructed the evpP-deleted mutant (ΔevpP) and complemented strain (ΔevpP-C) to study the effects of evpP deletion on the biological characteristics of E. tarda and infection in macrophages. [Results] No significant differences were observed in the growth curves or physiological and biochemical properties among the wild-type (WT), ΔevpP, and ΔevpP-C. However, compared with WT, ΔevpP exhibited significantly reduced motility, biofilm formation, adhesion rate to RAW264.7 macrophages, intracellular proliferation capacity, and ability to induce host cell autophagy, while triggering increased secretion of tumor necrosis factor-α (TNF-α) by macrophages. The complementation of evpP-C did not fully restore the intracellular proliferation capability, but completely rescued the other phenotypic defects. [Conclusion] EvpP does not affect the growth or physiological and biochemical properties of E. tarda. However, it could enhance the bacterial motility, biofilm formation, adhesion to macrophages, intracellular proliferation, and autophagy, while suppressing TNF-α secretion in E. tarda-infected macrophages. These findings confirm that EvpP plays a critical role in the pathogenicity of E. tarda.

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E-mail:
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VI型分泌系统(type VI secretion system, T6SS)是迟缓爱德华氏菌(Edwardsiella tarda)的一种新型毒力因子。迟缓爱德华氏菌毒力蛋白P (E. tarda virulence protein P, EvpP)是T6SS的效应蛋白，但目前对EvpP功能机制的研究仍十分有限。 【目的】 解析EvpP的生物学功能，深入探究T6SS在E. tarda致病过程中的作用。 【方法】 构建E. tarda的evpP基因缺失株(ΔevpP)和回补株(ΔevpP-C)，在此基础上开展evpP基因缺失对E. tarda生物学特性以及巨噬细胞感染影响的研究。 【结果】 E. tarda野生株、ΔevpP和ΔevpP-C在生长曲线和生理生化特性方面并无显著差异。相较于野生株，ΔevpP的运动性、生物被膜生成能力以及感染RAW264.7巨噬细胞的黏附率、胞内增殖率和触发细胞自噬的能力均显著下降，但能诱发巨噬细胞分泌更多的肿瘤坏死因子α (tumor necrosis factor-α, TNF-α)。ΔevpP-C的胞内增殖能力未完全恢复至野生株水平，但其他表型均得到完全恢复。 【结论】 EvpP不影响E. tarda的生长和生理生化特性，但能不同程度地增强该菌的运动性、生物膜形成能力以及对巨噬细胞的黏附、胞内增殖能力和自噬活性，且能抑制E. tarda感染的巨噬细胞分泌更多TNF-α。研究结果进一步证实EvpP参与E. tarda 的致病过程，并在其中发挥重要作用。

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作者贡献声明

杨子轩：实验操作、数据分析和论文撰写；康若茗：实验操作和数据收集；胡玉帅：数据分析和验证；高迎莉：提供菌株构建技术支持；王淑芳：提供Western blotting技术支持；秦蕾：提供经费支持、实验指导、论文修改。

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Autophagy participates in the Edwardsiella tarda-macrophage interaction and influences on the inflammatory response[J]. Microbiology China, 2025, 52(11): 5226-5239 (in Chinese)., articleTitle=null, refAbstract=null)], funds=[Fund(id=1226195564602241774, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, awardId=BK20221396, language=EN, fundingSource=the Natural Science Foundation of Jiangsu Province(BK20221396), fundOrder=null, country=null), Fund(id=1226195564816151283, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, awardId=BK20221396, language=CN, fundingSource=江苏省自然科学基金(BK20221396), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1226195555785814404, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, xref=null, ext=[AuthorCompanyExt(id=1226195555794203013, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, companyId=1226195555785814404, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=School of Marine Science and Fisheries, Jiangsu Ocean University, Lianyungang, Jiangsu, China), AuthorCompanyExt(id=1226195555802591623, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, companyId=1226195555785814404, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=江苏海洋大学 海洋科学与水产学院，江苏 连云港)])], figs=[ArticleFig(id=1226195560798007911, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=EN, label=Figure 1, caption=Detection of evpP deletion mutants (A) and complemented strains (B). M: DL2000 DNA marker; 1: ΔevpP; 2: Wild type; 3: ΔevpP-C; 4: ΔevpP; 5: E. coli β2163 (evpP-pBAD33cm-rp4)., figureFileSmall=ETOID5HlwvPe5UF+QAtxIA==, figureFileBig=ExdGjWmpU+2E/Lzv8NEVdQ==, tableContent=null), ArticleFig(id=1226195560902865521, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=CN, label=图1, caption=迟缓爱德华氏菌 evpP 缺失株(A)和回补株(B)的检测。M：DL2000 DNA marker；1：evpP缺失株；2：野生株；3：回补株；4：evpP缺失株；5：大肠杆菌β2163 (evpP-pBAD33cm-rp4)。, figureFileSmall=ETOID5HlwvPe5UF+QAtxIA==, figureFileBig=ExdGjWmpU+2E/Lzv8NEVdQ==, tableContent=null), ArticleFig(id=1226195561041277561, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=EN, label=Figure 2, caption=The growth curves of WT, ΔevpP and ΔevpP-C., figureFileSmall=BySgOOo0MXg2crTqVocQCQ==, figureFileBig=wHzdz3RXxW0AFwc+rhCzzw==, tableContent=null), ArticleFig(id=1226195561175495301, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=CN, label=图2, caption=野生株(WT)、缺失株(ΔevpP)和回补株(ΔevpP-C)的生长曲线, figureFileSmall=BySgOOo0MXg2crTqVocQCQ==, figureFileBig=wHzdz3RXxW0AFwc+rhCzzw==, tableContent=null), ArticleFig(id=1226195561271964298, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=EN, label=Figure 3, caption=Motility determination of WT, ΔevpP and ΔevpP-C strains. A: Swimming path of WT, ΔevpP and ΔevpP-C; B: Motility zone diameter of WT, ΔevpP and ΔevpP-C (***: P<0.001; ns: P>0.05)., figureFileSmall=Q75WSMHpCew/dI1aGXrnnA==, figureFileBig=pIKe3PpgFSEYyDkaRTb/1g==, tableContent=null), ArticleFig(id=1226195561364238993, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=CN, label=图3, caption=菌株WT、ΔevpP 和ΔevpP-C的运动性测定。A：三株菌株的运动轨迹；B：运动直径测定统计(***：P<0.001；ns：P>0.05)。, figureFileSmall=Q75WSMHpCew/dI1aGXrnnA==, figureFileBig=pIKe3PpgFSEYyDkaRTb/1g==, tableContent=null), ArticleFig(id=1226195561456513687, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=EN, label=Figure 4, caption=The biofilm forming of WT, ΔevpP和ΔevpP-C. A: Biofilm formation of the strains by crystal violet staining at 48 h; B: Absorbance measurement of biofilm formation at 595 nm (***: P<0.001; ns: P>0.05)., figureFileSmall=8gK9gJKBid6uc7H4rglVoQ==, figureFileBig=8bDSvlgegzu5ZuzP7pp3bw==, tableContent=null), ArticleFig(id=1226195561552982682, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=CN, label=图4, caption=菌株WT、ΔevpP 和ΔevpP-C的生物被膜形成情况。A：菌株培养48 h后结晶紫染色；B：生物被膜的吸光度(OD595)测定(***：P<0.001；ns：P>0.05)。, figureFileSmall=8gK9gJKBid6uc7H4rglVoQ==, figureFileBig=8bDSvlgegzu5ZuzP7pp3bw==, tableContent=null), ArticleFig(id=1226195561649451679, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=EN, label=Figure 5, caption=Loss of evpP results in reduced adherence of Edwardsiella tarda to RAW264.7 cells. A: Giemsa-stained RAW264.7 cells infected by WT, ΔevpP and ΔevpP-C strains, respectively; B: Adhesion rate of three strains expressed as cell-associated CFU/initial CFU (***: P<0.001; ns: P>0.05)., figureFileSmall=nx4YlCFgCs3RdxclPH2OKw==, figureFileBig=+cvqxcyUCC2orLQUQibsbg==, tableContent=null), ArticleFig(id=1226195561758503588, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=CN, label=图5, caption=evpP 的缺失导致 Edwardsiella tarda 对RAW264.7细胞的黏附性降低。A：WT、ΔevpP和ΔevpP-C菌株感染RAW264.7细胞的姬姆萨染色；B：菌株的黏附率统计(***：P<0.001；ns：P>0.05)。, figureFileSmall=nx4YlCFgCs3RdxclPH2OKw==, figureFileBig=+cvqxcyUCC2orLQUQibsbg==, tableContent=null), ArticleFig(id=1226195561875944107, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=EN, label=Figure 6, caption=Loss of evpP results in reduced intracellular proliferation of Edwardsiella tarda in RAW264.7 cells. A: Giemsa-stained RAW264.7 cells infected by WT, ΔevpP and ΔevpP-C strains, respectively; B: Intracellular proliferation rate of WT, ΔevpP and ΔevpP-C (*: P<0.05; ***: P<0.001)., figureFileSmall=/Y8os0Y8NHB9lURCIQ/png==, figureFileBig=LKo3i1NoZy+eMNyEVqeeDw==, tableContent=null), ArticleFig(id=1226195562022744751, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=CN, label=图6, caption=evpP 缺失削弱 Edwardsiella tarda 胞内增殖能力。A：WT、ΔevpP和ΔevpP-C菌株感染RAW264.7细胞的姬姆萨染色；B：各菌株的胞内增殖率(*：P<0.05；***：P<0.001)。, figureFileSmall=/Y8os0Y8NHB9lURCIQ/png==, figureFileBig=LKo3i1NoZy+eMNyEVqeeDw==, tableContent=null), ArticleFig(id=1226195562127602356, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=EN, label=Figure 7, caption=Analysis of TNF-α secreted by RAW264.7 cells after exposure to WT, ΔevpP and ΔevpP-C strains (***: P<0.001; ns: P>0.05)., figureFileSmall=PncY7AHA9tMJBCCz1HMAQw==, figureFileBig=TblxqRm4dRAmjeyYemmYUA==, tableContent=null), ArticleFig(id=1226195562286985918, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=CN, label=图7, caption=菌株感染RAW264.7巨噬细胞分泌TNF-α的检测分析, figureFileSmall=PncY7AHA9tMJBCCz1HMAQw==, figureFileBig=TblxqRm4dRAmjeyYemmYUA==, tableContent=null), ArticleFig(id=1226195563662717630, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=EN, label=Figure 8, caption=Autophagy induced by WT, ΔevpP and ΔevpP-C strains, respectively. A: Autophagy was confirmed by MDC staining (1: Control; 2: RAW264.7 cells infected with WT; 3: RAW264.7 cells infected with ΔevpP; 4: RAW264.7 cells infected with ΔevpP-C); B: LC3 was detected by Western blotting in cells infected with strains (+: Infected with this strain; -: Not infected with this strain; *: P<0.05; **: P<0.01; ***: P<0.001; ns: P>0.05)., figureFileSmall=+oVOxbvjZNZMlYEY8S4k9g==, figureFileBig=tnde26n9bQp+CctF1ZcItA==, tableContent=null), ArticleFig(id=1226195563822101190, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=CN, label=图8, caption=菌株感染RAW264.7巨噬细胞诱发的自噬。A：MDC染色观察(1：空白对照；2：WT感染巨噬细胞；3：ΔevpP感染巨噬细胞；4：ΔevpP-C感染巨噬细胞)；B：Western blotting检测菌株感染RAW264.7细胞的LC3表达水平(+：有；-：无；*：P<0.05；**：P<0.01；***：P<0.001；ns：P>0.05)。, figureFileSmall=+oVOxbvjZNZMlYEY8S4k9g==, figureFileBig=tnde26n9bQp+CctF1ZcItA==, tableContent=null), ArticleFig(id=1226195563947930319, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=EN, label=Table 1, caption=

Primers used in the construction of evpP deletion mutants and complemented strains

, figureFileSmall=null, figureFileBig=null, tableContent=

引物名称

Primer names

引物序列

Primer sequences (5′→3′)

产物大小

Product size (bp)

P1CATGACGTCTAGACGCTGACGCATCCATTATCGCAAC610
P2GATATGCCCGACCGTCAGGTTTGGTCCAACATCGGACCATTCCAGCTC
P3GAGCTGGAATGGTCCGATGTTGGACCAAACCTGACGGTCGGGCATATC587
P4TAGTCGTACTAGTTTGGCTGTCATCTCCGCTCAGGG
P5GACACAGTTGTAACTGGTCCA1 220
P6CAGGAACACTTAACGGCTGAC
P7CATGACGTCTAGAGCGTCGTGTCAGGAGGCGATGAA970/1 663
P8TAGTCGTACTAGTTTGCGCTCTTTCAGCTTGGGAAGC
P9CGAATTGGGTACCAGCGCTT273
P10TACCGTCGACGCCGGCCAGC
P11TGGGCTAGCGAATTCGAGCTAGGAGGAATTCACCATGTTGACTTTGAATTCCGAGCTG618
P12TGCCTGCAGGTCGACTCTAGCTATTTCAATATTGAAAATTGGTGGCTAC
P13CTAGAGTCGACCTGCAGGCA5 529
P14AGCTCGAATTCGCTAGCCCA
P15CCATAAGATTAGCGGATCCTACCT722
P16CTTCTCTCATCCGCCAAAACAG
), ArticleFig(id=1226195564082148054, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=CN, label=表1, caption=

evpP 缺失株和回补株构建所用引物

, figureFileSmall=null, figureFileBig=null, tableContent=

引物名称

Primer names

引物序列

Primer sequences (5′→3′)

产物大小

Product size (bp)

P1CATGACGTCTAGACGCTGACGCATCCATTATCGCAAC610
P2GATATGCCCGACCGTCAGGTTTGGTCCAACATCGGACCATTCCAGCTC
P3GAGCTGGAATGGTCCGATGTTGGACCAAACCTGACGGTCGGGCATATC587
P4TAGTCGTACTAGTTTGGCTGTCATCTCCGCTCAGGG
P5GACACAGTTGTAACTGGTCCA1 220
P6CAGGAACACTTAACGGCTGAC
P7CATGACGTCTAGAGCGTCGTGTCAGGAGGCGATGAA970/1 663
P8TAGTCGTACTAGTTTGCGCTCTTTCAGCTTGGGAAGC
P9CGAATTGGGTACCAGCGCTT273
P10TACCGTCGACGCCGGCCAGC
P11TGGGCTAGCGAATTCGAGCTAGGAGGAATTCACCATGTTGACTTTGAATTCCGAGCTG618
P12TGCCTGCAGGTCGACTCTAGCTATTTCAATATTGAAAATTGGTGGCTAC
P13CTAGAGTCGACCTGCAGGCA5 529
P14AGCTCGAATTCGCTAGCCCA
P15CCATAAGATTAGCGGATCCTACCT722
P16CTTCTCTCATCCGCCAAAACAG
), ArticleFig(id=1226195564212171483, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=EN, label=Table 2, caption=

Biochemical characteristics of WT, ΔevpP and ΔevpP-C

, figureFileSmall=null, figureFileBig=null, tableContent=
项目ItemsWTΔevpPΔevpP-C项目ItemsWTΔevpPΔevpP-C

V-P

Voges-Proskauer

---

棉子糖

Raffinose

---
山梨醇Sorbitol---麦芽糖Maltose+++
侧金盏花醇Adonitol---蜜二糖Melibiose---
甘露醇Mannitol---硫化氢Hydrogen sulfide+++
木糖醇Xylitol---苯丙氨酸Phenylalanine---

葡萄糖

Glucose

+++

赖氨酸脱羧酶

Lysine decarboxylas

+++

果糖

Fructose

+++

鸟氨酸脱羧酶

Ornithine decarboxylas

+++

蔗糖

Sucrose

---

精氨酸双水解酶

Arginase dihydrolase

+++
甘露糖Mannose+++脲酶Urease---
乳糖Lactose---吲哚Indole+++
木糖Xylose---氧化酶Oxidase---
), ArticleFig(id=1226195564333806307, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226136784468357386, language=CN, label=表2, caption=

菌株WT、ΔevpP 和ΔevpP-C的生理生化特性

, figureFileSmall=null, figureFileBig=null, tableContent=
项目ItemsWTΔevpPΔevpP-C项目ItemsWTΔevpPΔevpP-C

V-P

Voges-Proskauer

---

棉子糖

Raffinose

---
山梨醇Sorbitol---麦芽糖Maltose+++
侧金盏花醇Adonitol---蜜二糖Melibiose---
甘露醇Mannitol---硫化氢Hydrogen sulfide+++
木糖醇Xylitol---苯丙氨酸Phenylalanine---

葡萄糖

Glucose

+++

赖氨酸脱羧酶

Lysine decarboxylas

+++

果糖

Fructose

+++

鸟氨酸脱羧酶

Ornithine decarboxylas

+++

蔗糖

Sucrose

---

精氨酸双水解酶

Arginase dihydrolase

+++
甘露糖Mannose+++脲酶Urease---
乳糖Lactose---吲哚Indole+++
木糖Xylose---氧化酶Oxidase---
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EvpP对迟缓爱德华氏菌生物学特性及感染巨噬细胞的影响
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杨子轩 , 康若茗 , 胡玉帅 , 高迎莉 , 王淑芳 , 秦蕾
微生物学报 | 研究报告 2026,66(2): 801-814
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微生物学报 | 研究报告 2026, 66(2): 801-814
EvpP对迟缓爱德华氏菌生物学特性及感染巨噬细胞的影响
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杨子轩, 康若茗, 胡玉帅, 高迎莉, 王淑芳, 秦蕾
作者信息
  • 江苏海洋大学 海洋科学与水产学院，江苏 连云港
Impacts of EvpP on the biological characteristics of Edwardsiella tarda and infection of macrophages
Zixuan YANG, Ruoming KANG, Yushuai HU, Yingli GAO, Shufang WANG, Lei QIN
Affiliations
  • School of Marine Science and Fisheries, Jiangsu Ocean University, Lianyungang, Jiangsu, China
出版时间: 2026-02-04 doi: 10.13343/j.cnki.wsxb.20250672
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摘要
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VI型分泌系统(type VI secretion system, T6SS)是迟缓爱德华氏菌(Edwardsiella tarda)的一种新型毒力因子。迟缓爱德华氏菌毒力蛋白P (E. tarda virulence protein P, EvpP)是T6SS的效应蛋白，但目前对EvpP功能机制的研究仍十分有限。 【目的】 解析EvpP的生物学功能，深入探究T6SS在E. tarda致病过程中的作用。 【方法】 构建E. tarda的evpP基因缺失株(ΔevpP)和回补株(ΔevpP-C)，在此基础上开展evpP基因缺失对E. tarda生物学特性以及巨噬细胞感染影响的研究。 【结果】 E. tarda野生株、ΔevpP和ΔevpP-C在生长曲线和生理生化特性方面并无显著差异。相较于野生株，ΔevpP的运动性、生物被膜生成能力以及感染RAW264.7巨噬细胞的黏附率、胞内增殖率和触发细胞自噬的能力均显著下降，但能诱发巨噬细胞分泌更多的肿瘤坏死因子α (tumor necrosis factor-α, TNF-α)。ΔevpP-C的胞内增殖能力未完全恢复至野生株水平，但其他表型均得到完全恢复。 【结论】 EvpP不影响E. tarda的生长和生理生化特性，但能不同程度地增强该菌的运动性、生物膜形成能力以及对巨噬细胞的黏附、胞内增殖能力和自噬活性，且能抑制E. tarda感染的巨噬细胞分泌更多TNF-α。研究结果进一步证实EvpP参与E. tarda 的致病过程，并在其中发挥重要作用。

迟缓爱德华氏菌  /  EvpP  /  巨噬细胞  /  生物学特性

[Objective] The type VI secretion system (T6SS) is a novel virulence factor of Edwardsiella tarda. E. tarda virulence protein P (EvpP) is an effector of the T6SS. To date, the functional mechanisms of EvpP are still poorly understood. This study aimed to comprehensively investigate the biological functions of EvpP and elucidate the roles of T6SS in the pathogenicity of E. tarda. [Methods] We constructed the evpP-deleted mutant (ΔevpP) and complemented strain (ΔevpP-C) to study the effects of evpP deletion on the biological characteristics of E. tarda and infection in macrophages. [Results] No significant differences were observed in the growth curves or physiological and biochemical properties among the wild-type (WT), ΔevpP, and ΔevpP-C. However, compared with WT, ΔevpP exhibited significantly reduced motility, biofilm formation, adhesion rate to RAW264.7 macrophages, intracellular proliferation capacity, and ability to induce host cell autophagy, while triggering increased secretion of tumor necrosis factor-α (TNF-α) by macrophages. The complementation of evpP-C did not fully restore the intracellular proliferation capability, but completely rescued the other phenotypic defects. [Conclusion] EvpP does not affect the growth or physiological and biochemical properties of E. tarda. However, it could enhance the bacterial motility, biofilm formation, adhesion to macrophages, intracellular proliferation, and autophagy, while suppressing TNF-α secretion in E. tarda-infected macrophages. These findings confirm that EvpP plays a critical role in the pathogenicity of E. tarda.

Edwardsiella tarda  /  EvpP  /  macrophages  /  biological characteristics
杨子轩, 康若茗, 胡玉帅, 高迎莉, 王淑芳, 秦蕾. EvpP对迟缓爱德华氏菌生物学特性及感染巨噬细胞的影响. 微生物学报, 2026 , 66 (2) : 801 -814 . DOI: 10.13343/j.cnki.wsxb.20250672
Zixuan YANG, Ruoming KANG, Yushuai HU, Yingli GAO, Shufang WANG, Lei QIN. Impacts of EvpP on the biological characteristics of Edwardsiella tarda and infection of macrophages[J]. Acta Microbiologica Sinica, 2026 , 66 (2) : 801 -814 . DOI: 10.13343/j.cnki.wsxb.20250672
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迟缓爱德华氏菌(Edwardsiella tarda)是一种重要的人兽共患病原体，其宿主范围广泛，可引起鱼类、两栖类、爬行类、哺乳类甚至人类的感染[1]，尤其对水产养殖业危害巨大[2]。E. tarda属于胞内寄生菌，能够在宿主巨噬细胞内生存繁殖，进而促进自身感染[3-4]。目前，对E. tarda复杂的致病机理了解尚不充分，这在很大程度上限制了迟缓爱德华氏菌病的有效治疗与防控工作。
在众多毒力因子中，VI型分泌系统(type VI secretion system, T6SS)被认为是E. tarda发挥毒力所必需的，在E. tarda的致病过程中发挥着重要作用[5-6]。T6SS是新近确定的与细菌毒力和竞争生存密切相关的分泌系统，由结构蛋白、转位蛋白和分泌蛋白等组成[7-8]，其中分泌蛋白能够决定T6SS的杀伤功能[9]。作为T6SS的分泌蛋白，迟缓爱德华氏菌毒力蛋白P (E. tarda virulence protein P, EvpP)已被证实参与E. tarda的致病过程[10]。EvpP是E. tarda特有的分泌蛋白，其转录受到双组分系统EsrA-EsrB和铁摄取调节因子Fur的正向调控[11]；Zhang等[12]通过体内实验表明，evpP的转录可被调节因子H-NS抑制。阻断EvpP的表达会使E. tarda的毒性显著削弱，并影响菌株侵染上皮细胞、溶血活性、黏液穿透力以及抵抗血清杀伤和体内寄生的能力[6,11]。王鑫[13]研究发现，E. tarda入侵鼠巨噬细胞后EvpP的表达量显著上升，表明EvpP参与E. tarda与巨噬细胞的互作关系。进一步研究发现，EvpP能够抑制细胞质内Ca2+的增加，阻断炎性小体接头蛋白ASC的寡聚化，通过Ca2+依赖性MAPK-Jnk途径阻止NOD样受体热蛋白结构域相关蛋白3 (NOD-like receptor thermal protein domain associated protein 3, NLRP3)炎性小体的激活，从而促进E. tarda在巨噬细胞内的定殖[14]。动物实验表明EvpP能够抑制Jnk-caspy炎性小体信号通路，并显著抑制中性粒细胞向感染部位的募集[15]。作为E. tarda非常重要的毒力因子，目前尚不清楚T6SS如何通过其分泌蛋白EvpP影响E. tarda的生物学特性，以及在与巨噬细胞的互作过程中具体参与哪些环节。
本研究基于同源重组技术构建了迟缓爱德华氏菌的evpP基因缺失株和回补株。在此基础上，对E. tarda野生株、缺失株和回补株在生长曲线、生理生化特性、运动性、生物膜形成以及与巨噬细胞相互作用(包括黏附、胞内增殖、炎性因子分泌和细胞自噬等方面)进行比较分析，探究EvpP在E. tarda生物学特性及与巨噬细胞互作中所扮演的角色。研究结果综合评估了EvpP的功能，为证实EvpP参与E. tarda 致病过程并承担重要角色提供更多科学依据和参考，同时也为寻找防控迟缓爱德华氏菌病的有效措施提供新的思路。
1 材料与方法
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1.1 菌株、质粒与细胞系
E. tarda、E. coli β2163、E. coli DH5α λpir菌株以及pLP12、pBAD质粒均由本实验室保存；RAW264.7巨噬细胞系由中国科学院干细胞库提供。
1.2 主要试剂和仪器
TSB、TSA、LB培养基以及细菌微量生化鉴定管均购自杭州微生物试剂有限公司；DMEM培养基购自HyClone公司；胎牛血清购自Gibco公司；电泳预制胶购自南京金斯瑞生物科技有限公司；TNF-α检测试剂盒购自武汉伊莱瑞特生物科技股份有限公司；姬姆萨染色液购自北京索莱宝科技有限公司；MDC自噬检测试剂盒购自上海碧云天生物技术股份有限公司；兔抗鼠LC3多克隆抗体、兔抗鼠Tubulin内参抗体、羊抗兔二抗购自武汉三鹰生物技术有限公司；PrimeSTAR Max DNA Polymerase、限制性内切酶和DNA连接酶购自宝生物工程(大连)有限公司；ClonExpress® II One Step Cloning Kit购自南京诺唯赞生物科技股份有限公司；细菌基因组提取试剂盒、质粒提取试剂盒购自天根生化科技(北京)有限公司。
酶标仪，ThermoFisher Scientific公司；CO2细胞培养箱，RS Biotech公司；细菌浊度仪，杭州齐威仪器有限公司；荧光显微镜，Nikon公司；电泳仪，北京六一生物科技有限公司；半干转印仪，Cytiva公司。
1.3 E.tarda 的 evpP 缺失株(ΔevpP)和回补株(ΔevpP-C)的构建
1.3.1 引物的设计
根据GenBank中E. tardaevpP基因序列(Gene编号ID：8608773)，利用Primer Premier 5.0设计evpP缺失株和回补株构建所需引物(表1)，由上海捷瑞生物工程有限公司进行引物合成。
1.3.2 evpP 缺失株(ΔevpP)的构建
使用试剂盒提取E. tarda菌株基因组DNA并作为模板，分别扩增evpP基因的上游和下游同源臂。通过交叠PCR获得融合片段后克隆至pLP12自杀质粒构建重组质粒(pLP12-evpP)。将重组质粒转化大肠杆菌DH5α λpir并涂布LB平板(20 μg/mL Cm, 0.3% d-glucose)进行筛选。将重组质粒转化E. coli β2163，涂布LB平板(20 μg/mL Cm, 0.3 mmol/L DAP)筛选阳性克隆。利用TSA平板(0.3 mmol/L DAP, 0.3% d-glucose)开展接合试验，将重组质粒从供体菌E. coli β2163转移至E. tarda中，继续涂布平板(20 μg/mL Cm, 0.3% d-glucose)筛选插入突变株并进行PCR检测。取一个插入突变克隆经TSB (0.3% d-glucose)培养后涂布平板(0.4% l-arabinose)筛选阳性克隆，PCR检测并进行测序验证。
1.3.3 回补株(ΔevpP-C)的构建
参照1.3.2节缺失株构建的方法，将质粒pBAD33cm-rp4转化于E. coli β2163感受态细胞，涂布LB平板筛选阳性克隆。将ΔevpP与E. coli β2163 进行接合实验，确定质粒pBAD33cm-rp4能够在ΔevpP中进行复制。以E. tarda菌株基因组DNA为模板，扩增纯化evpP基因，并与扩增的载体片段进行无缝克隆连接，PCR检测筛选重组克隆，并进行测序验证。将阳性克隆质粒转化E. coli β2163，然后通过接合试验将evpP-pBAD33cm-rp4重组质粒转入ΔevpP。挑取若干重组克隆进行检测，阳性克隆纯化培养后测序验证。
1.4 菌株生长曲线的测定
分别挑取WT、ΔevpP和ΔevpP-C菌落接种至TSB培养基，28 ℃、120 r/min振荡培养至对数生长期(OD600为0.6)。使用细菌浊度仪将菌液调至统一浓度后，按1%接种量加入新鲜TSB培养基，28 ℃、120 r/min振荡培养24 h，在指定时间点取样测定OD600值并绘制生长曲线。试验重复3次。
1.5 菌株生理生化检测
分别将WT、ΔevpP和ΔevpP-C菌株接种于TSA培养基，28 ℃倒置培养12 h，挑取菌落用生理盐水制成菌悬液，采用细菌浊度仪统一调至0.5个麦氏比浊单位，分别吸取适量菌液加入细菌生理生化测定管中，28 ℃培养箱培养24 h，观察并根据鉴定管判读表记录结果。试验重复3次。
1.6 菌株运动性测定
将WT、ΔevpP和ΔevpP-C菌株分别使用TSB培养基28 ℃、120 r/min振荡培养至对数生长期(OD600为0.6)，采用细菌浊度仪将菌液浓度调至一致，分别吸取1 µL菌液垂直点在0.5%琼脂半固体培养基上，28 ℃培养12 h，拍照并测量细菌运动所得轨迹的直径，分析比较3组菌株的运动能力差异。试验重复3次。
1.7 菌株生物被膜形成能力测定
将WT、ΔevpP和ΔevpP-C菌株于28 ℃、120 r/min振荡培养至对数生长期(OD600为0.6)，使用新鲜TSB培养基调整各菌液OD600至0.1，各取200 μL菌液至96孔板，每个实验组设8个平行孔，TSB培养基作为空白对照。28 ℃静置培养48 h后弃除菌液，PBS洗涤3次，自然风干，加入200 μL纯甲醇固定10 min，吸净甲醇并晾干，1%结晶紫染色15 min，PBS洗涤3次后晾干。完全干燥后加入200 μL无水乙醇作用30 min，酶标仪测量光密度值(OD595)。试验重复3次。
1.8 巨噬细胞黏附试验
将状态良好的RAW264.7细胞平铺于24孔板，每孔1×105个细胞。37 ℃、5% CO2培养过夜后，按照感染复数MOI (细菌:巨噬细胞)=100:1的比例加入菌液进行感染实验。培养板800 r/min离心5 min，孵育1 h。吸弃培养基，无菌PBS轻柔润洗细胞5-6次，每孔加入1 mL 1% Triton X-100裂解细胞15 min，裂解液梯度稀释后涂布于TSA平板，28 ℃倒置培养24 h进行CFU计数并计算黏附率，计算如公式(1)所示。
黏附率=(黏附细胞菌数/孔内感染菌数)×100%
在24孔板中放入细胞爬片进行Giemsa染色观察。侵染步骤同上，细胞孵育完成后，将孔中培养液吸出，PBS洗3次后取出细胞爬片，甲醇固定10 min，最后用姬姆萨工作液染色10 min，冲洗脱色后镜检拍照。每个实验组设3个平行。试验重复3次。
1.9 胞内菌株增殖率检测
参照Subashchandrabose等[16]的方法并作适当修改。侵染步骤同1.8节黏附试验，孵育30 min后吸弃培养基，PBS轻柔洗涤3次，换含有100 µg/mL庆大霉素的完全培养基进行处理。实验分两组(T0和T1)，每个实验组设3个平行。T0组在庆大霉素加入15 min后采用PBS洗涤3次，1% Triton X-100裂解细胞，稀释涂布平板计数。T1组则在庆大霉素加入2 h后裂解并稀释涂布平板。细菌胞内增殖率计算如公式(2)所示。
增殖率=(T1-T0)/T0×100%
细胞侵染及姬姆萨染色步骤同1.8节，试验重复3次。
1.10 巨噬细胞分泌TNF-α的测定
采用ELISA试验检测菌株WT、ΔevpP和ΔevpP-C感染RAW264.7细胞后肿瘤坏死因子α (tumor necrosis factor-α, TNF-α)的分泌表达量。细胞侵染步骤同1.8节黏附试验，每株菌设3个平行，孵育1 h后用100 μg/mL庆大霉素培养基处理1 h，轻柔洗涤后换新鲜培养基继续培养。分别在感染后12 h和24 h取细胞上清，按照TNF-α ELISA Kit说明书进行TNF-α的检测。试验重复3次。
1.11 免疫荧光监测巨噬细胞自噬
细胞侵染步骤同1.8节黏附试验，庆大霉素处理1 h后更换新鲜培养基，继续培养6 h后按照MDC自噬检测试剂盒说明书进行细胞染色，倒置荧光显微镜下观察并拍照。
1.12 Western blotting检测LC3蛋白的表达量
采用免疫印迹法检测菌株WT、ΔevpP和ΔevpP-C感染巨噬细胞后微管相关蛋白1轻链3 (LC3)分子的表达量。感染实验同1.8节黏附试验，分别在感染后4 h和8 h提取细胞总蛋白，使用BCA蛋白定量试剂盒测定各组样品蛋白浓度。样品95 ℃变性15 min后进行SDS-PAGE分离LC3目的蛋白。电泳完成后使用14 V恒压半干转膜50 min。取出PVDF膜，甲醇浸泡1 min，室温下水平摇床封闭1 h。将膜分别置于兔抗鼠LC3一抗(1:2 500)、兔抗鼠β-tubulin一抗(1:7 500) 4 ℃孵育过夜，随后TBST清洗，羊抗兔二抗(1:7 500)孵育1 h，TBST漂洗。最后加入ECL发光液进行显色，成像系统曝光显影。使用ImageJ分析各条带灰度值。试验重复3次。
1.13 统计学分析
本研究数据使用GraphPad Prism 9进行统计学分析，且数据均以平均值±标准偏差(mean±SD)表示，两组之间差异的显著性使用独立的t检验进行评估，多组之间的比较则采用单因素方差分析。P<0.05被认为具有统计学意义。
2 结果与分析
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2.1 迟缓爱德华氏菌 evpP 缺失株(ΔevpP)和回补株(ΔevpP-C)的鉴定
通过自杀质粒介导的同源重组方法，以氯霉素抗性标记和阿拉伯糖诱导的致死基因vmt进行正、反向双重筛选。对筛选后的克隆进行检测，以野生型为对照，缺失突变的克隆PCR扩增产物片段大小为1 195 bp，而野生株的扩增片段大小为1 663 bp (图1A)，符合预期。克隆纯化后，再次扩增验证，PCR产物测序结果证实evpP缺失突变株构建成功，命名为ΔevpP。
扩增evpP基因与pBAD33载体序列，连接构建evpP-pBAD33cm-rp4重组质粒，转化ΔevpP。以ΔevpP和E. coli β2163 (evpP-pBAD33cm-rp4)为对照，PCR检测筛选重组克隆。如图1B所示，阳性克隆扩增片段为722 bp。纯化阳性克隆，并进行测序验证，测序结果表明回补菌株构建成功，命名为ΔevpP-C。
2.2 evpP 缺失不影响 E. tarda 的生长能力
对E. tarda野生株WT、ΔevpP和ΔevpP-C进行生长能力测定并绘制生长曲线(图2)。可见3株菌的生长特性并无明显差异，在相同培养条件下均于培养后8 h进入对数期，16 h进入稳定期。研究表明，evpP基因的缺失不会影响E. tarda的生长能力。
2.3 evpP 缺失不影响 E. tarda 的生理生化特性
取对数生长期的WT、ΔevpP和ΔevpP-C菌株进行生理生化特征测定，结果显示3株菌的生理生化特性完全一致，均能发酵葡萄糖、果糖、甘露糖、麦芽糖，具有赖氨酸脱羧酶活性、鸟氨酸脱羧酶活性、精氨酸双水解酶活性，能够生成吲哚和H2S (表2)。提示evpP缺失不影响E. tarda的生理生化特性。
2.4 EvpP影响 E. tarda 的运动能力
对菌株WT、ΔevpP和ΔevpP-C进行运动性检测，并根据菌株的运动直径评估各菌株运动能力的变化。结果显示各菌株均呈现明显的运动性，但ΔevpP的运动能力明显受损(P<0.01)，而ΔevpP-C恢复野生株运动能力(图3)，提示EvpP与E. tarda的运动性相关，能够增强菌株的运动能力。
2.5 EvpP参与 E. tarda 生物被膜的形成
采用结晶紫染色法评估菌株WT、ΔevpP和ΔevpP-C产生生物膜的能力。如图4所示，evpP缺失使得ΔevpP菌株生物膜形成能力显著下降(P<0.001)，而回补株ΔevpP-C生物被膜形成能力与野生株相比无显著差异(P>0.05)。提示EvpP参与E. tarda的生物被膜的形成过程，二者呈正相关。
2.6 EvpP促进 E. tarda 黏附巨噬细胞
通过测定菌株WT、ΔevpP和ΔevpP-C对RAW264.7巨噬细胞的黏附率并进行比较分析，探究EvpP对E. tarda侵染巨噬细胞的影响。研究结果显示：缺失株的细胞黏附率较野生株显著降低(P<0.001)，且回补株的黏附能力得到恢复(图5)。这表明EvpP与E. tarda黏附巨噬细胞的过程相关，能够显著增强细菌侵染细胞的能力。
2.7 EvpP有助于 E. tarda 在巨噬细胞内的增殖
分别对菌株WT、ΔevpP和ΔevpP-C感染细胞后15 min组(T0)和2 h组(T1)进行计数，并按照公式(2)计算增殖率。如图6所示，缺失株的胞内增殖率较野生株显著降低(P<0.001)，说明E. tarda的胞内增殖能力明显受到evpP缺失的影响。ΔevpP-C的胞内增殖能力未完全恢复至野生株水平，暗示EvpP可能通过与其他功能蛋白形成调控网络共同参与E. tarda的胞内寄生过程。
2.8 EvpP抑制 E. tarda 感染的巨噬细胞分泌TNF-α
采用ELISA检测菌株WT、ΔevpP和ΔevpP-C刺激RAW264.7细胞分泌产生TNF-α的水平，并进行分析比较。结果显示，感染ΔevpP的细胞在12 h及24 h时TNF-α分泌水与菌株WT相比平均显著上调(P<0.001)，而ΔevpP-C感染的细胞TNF-α分泌水平无明显差异(图7)。这表明EvpP能够抑制E. tarda感染的巨噬细胞分泌产生TNF-α。
2.9 EvpP诱发 E. tarda 感染的巨噬细胞自噬
采用免疫荧光法检测自噬小体以评估细胞自噬活性。MDC染色观察感染8 h后的RAW264.7细胞发现，ΔevpP感染细胞激发的荧光强度弱于WT和ΔevpP-C (图8A)。每1个荧光斑点相当于1个自噬小体，说明evpP的缺失导致E. tarda感染RAW264.7细胞的自噬水平下降。为进一步验证免疫荧光观察结果，以自噬关键标志性分子LC3表达水平为评价指标，采用免疫印迹法检测菌株WT、ΔevpP和ΔevpP-C在不同感染时间点诱发的RAW264.7细胞自噬活性。如图8B所示，在感染后4 h和8 h，ΔevpP感染细胞LC3-Ⅱ转化水平均显著下调(P<0.001)，且ΔevpP-C诱发细胞自噬活性恢复至野生株水平。上述结果表明，EvpP能够触发E. tarda感染的RAW264.7巨噬细胞发生自噬。
3 讨论与结论
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本研究基于同源重组技术成功构建了迟缓爱德华氏菌的evpP基因缺失株和回补株，为后续深入探究EvpP的功能及其在E. tarda致病过程中所扮演的角色奠定了基础，同时也为相关疫苗的开发提供了新的思路。将evpP基因缺失株与E. tarda野生株进行比较分析，证实evpP基因缺失对E. tarda的生长特性无影响，这表明EvpP并非E. tarda生长生存所必需，也为后续开展一系列相关实验研究提供了科学依据。已有研究发现，一些细菌的毒力与其生理生化特性相关，酶可通过不同代谢途径调控毒力因子的表达，间接或直接增强细菌的致病性[17-18]。本研究表明，evpP基因的缺失并不影响E. tarda的生理生化特性，提示EvpP可能并非通过直接调控细菌生理生化特性的方式对E. tarda的毒力产生影响。
细菌T6SS分泌系统的多种相关蛋白能通过调控细菌运动和生物膜形成来影响细菌的致病性[19-20]。本研究结果显示，evpP基因缺失后E. tarda的运动能力显著下降，表明EvpP与E. tarda的运动性密切相关。鞭毛运动是细菌最主要的运动方式，不仅有利于病原菌的传播，还与生物膜的形成和毒力因子的表达密切相关。研究发现T6SS相关基因hcp2的缺失能影响细菌鞭毛蛋白FliC的产生与分泌，并削弱细菌的运动能力[21-22]。由此推测，EvpP也可能通过影响E. tarda鞭毛相关蛋白的表达和输出来影响细菌的运动能力，但其具体调控机制还有待解析。本研究结果进一步证实，EvpP能够促进E. tarda生物膜的形成。生物膜被认为是致病菌毒力的一个重要因素，与细菌的运动性有关[23-24]。He等[25]的研究表明，鞭毛蛋白对于E. tarda生物膜的形成至关重要。Leung等[26]研究表明，E. tarda生物膜的形成受多个基因调控，因此EvpP是否通过介导E. tarda的运动性来影响生物膜的形成还是通过其他调控方式仍有待进一步明确。
巨噬细胞在宿主抵御E. tarda感染时扮演着非常重要的角色。E. tarda是胞内寄生菌，被巨噬细胞吞噬后能够“劫持”利用巨噬细胞，促进其在细胞内的生存、增殖和细胞间的扩散，E. tarda与巨噬细胞的互作结果将直接影响迟缓爱德华氏菌病的发展和转归[3-4]。鉴于巨噬细胞在宿主对E. tarda免疫反应中的重要性以及EvpP的致病性，推测EvpP可能直接或间接参与E. tarda与巨噬细胞的相互作用过程。细菌对细胞的黏附是其定殖过程中的重要环节，多种病原菌的T6SS相关基因已被证实参与细菌黏附的调控[20,27]。本研究表明，EvpP能够增强E. tarda对RAW264.7细胞的黏附，促进E. tarda与巨噬细胞的相互作用，为细菌T6SS参与宿主细胞黏附提供了新证据。Wu等[22]证实T6SS相关蛋白Hcp2能通过正向调控鞭毛系统介导溶藻弧菌的黏附性，但目前还不能确定EvpP促进E. tarda黏附的真正机制。EvpP是通过调控鞭毛蛋白的表达，还是通过其他方式，如调控黏附素分泌，还需进一步的试验来证实。E. tarda能够在巨噬细胞中存活和增殖是其致病的关键因素[3]。王鑫[13]研究发现EvpP的表达量在E. tarda感染的鼠巨噬细胞中显著上升，提示EvpP参与E. tarda与巨噬细胞的互作关系。本研究发现，与WT菌株相比，ΔevpP菌株在巨噬细胞内的增殖能力显著减弱，进一步证实了EvpP在E. tarda与巨噬细胞互作关系中的重要性。然而，ΔevpP-C菌株在本研究中并未完全恢复至野生型水平，暗示EvpP可能与其他毒力因子存在协同调控机制，共同形成某个调控网络对E. tarda的胞内命运产生影响。已有研究也发现，EvpP能通过Ca2+依赖性MAPK-Jnk途径促进E. tarda在巨噬细胞内的定殖[14]。是否还存在其他EvpP参与的E. tarda胞内生存调控的路径？这些调控路径所参与的蛋白是否受到evpP基因敲除的影响，使得ΔevpP-C只是部分恢复野生型的胞内增殖水平？这些问题都有待进一步探索与研究。巨噬细胞一旦识别外来细菌感染会被激活并分泌炎症细胞因子[28]。TNF-α是介导宿主炎症反应的关键细胞因子，是巨噬细胞控制外来入侵感染的重要动员信号[29-30]。TNF-α也被发现在针对鱼类迟缓爱德华氏菌病的免疫反应中发挥着重要作用[31]。本研究显示，ΔevpP菌株相较E. tarda野生株诱导巨噬细胞产生更多的TNF-α，提示E. tarda能够在巨噬细胞内生存、增殖与EvpP调控TNF-α的表达分泌相关，EvpP能够抑制巨噬细胞TNF-α的分泌进而达到E. tarda逃避免疫攻击的目的。
自噬是广泛存在于真核细胞内的一种溶酶体依赖性的降解途径。巨噬细胞作为极其重要的免疫细胞，其自噬过程在病原体感染中的地位日益受到关注[32]。自噬体的形成是自噬过程的关键步骤之一，其数量反映了细胞自噬的活跃程度。自噬发生时LC3-I会转化为LC3-II，附着在自噬体膜上。通过检测LC3蛋白的表达水平，可以评估细胞自噬的程度和活性[33]。在本研究中E. tarda缺失evpP后显著下调巨噬细胞自噬水平，表明EvpP能够诱发RAW264.7巨噬细胞发生自噬。类似报道也表明，副溶血弧菌T6SS的VgrG2蛋白能够诱发巨噬细胞自噬[34]。自噬在细菌感染中具有双重作用，既可帮助宿主细胞清除胞内细菌抵御感染，也可被一些病原菌利用加剧感染[35]。胡玉帅等[36]研究发现，E. tarda刚感染巨噬细胞时自噬能促进对细菌的清除，但很快E. tarda开始利用自噬促进其在胞内的增殖。推测E. tarda可能利用EvpP去触发巨噬细胞自噬，从而有助于自身的胞内生存。研究结果也为揭示E. tarda触发自噬的机制奠定了基础。
综上所述，本研究表明EvpP对E. tarda的生长和生理生化特性无显著影响，但参与E. tarda的运动和生物膜形成过程，并在E. tarda与巨噬细胞的互作环节，包括黏附、胞内增殖、炎症应答和自噬过程，都发挥了重要的作用。研究结果丰富了对EvpP的认知，为进一步研究E. tarda的感染机制及其对宿主免疫应答的影响奠定了基础。关于E. tarda与T6SS分泌系统致病性之间关系的解析，对开发研究新药和靶向治疗、制定迟缓爱德华氏菌有效防控措施具有重要意义。后续将进一步开展分子机制方面的研究，从分子层面深入探究EvpP对上述表型影响的具体机制，并对相关推测进行验证。
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  • 江苏省自然科学基金(BK20221396)
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doi: 10.13343/j.cnki.wsxb.20250672
  • 接收时间：2025-09-03
  • 首发时间：2026-02-05
  • 出版时间：2026-02-04
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  • 收稿日期：2025-09-03
  • 录用日期：2025-10-11
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the Natural Science Foundation of Jiangsu Province(BK20221396)
江苏省自然科学基金(BK20221396)
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    江苏海洋大学 海洋科学与水产学院，江苏 连云港
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鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
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