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Article(id=1228017375233110351, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1228017371202388759, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20240645, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1729008000000, receivedDateStr=2024-10-16, revisedDate=null, revisedDateStr=null, acceptedDate=1732032000000, acceptedDateStr=2024-11-20, onlineDate=1770711757715, onlineDateStr=2026-02-10, pubDate=1741017600000, pubDateStr=2025-03-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1770711757715, onlineIssueDateStr=2026-02-10, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1770711757715, creator=13701087609, updateTime=1770711757715, updator=13701087609, issue=Issue{id=1228017371202388759, tenantId=1146029695717560320, journalId=1192105938417971205, year='2025', volume='65', issue='3', pageStart='871', pageEnd='1336', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1770711756754, creator=13701087609, updateTime=1770719134572, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1228048316089434941, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1228017371202388759, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1228048316093629246, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1228017371202388759, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=1137, endPage=1147, ext={EN=ArticleExt(id=1228017375719649629, articleId=1228017375233110351, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Construction and biological characterization of Brucella S2Δbp26, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

[Objective] To develop a safe, effective, and identifiable new vaccine. [Methods] The gene deletion strain S2Δbp26 was constructed by homologous recombination with S2 as the parental strain. The stability, safety, and efficacy of S2Δbp26 were then evaluated. [Results] The phenotype and genotype of S2Δbp26 did not change after 20 successive passages invitro. The experiments in mice and guinea pigs showed that there was no difference in biosafety between S2Δbp26 and S2. The bacterial load was less than 2×105 CFU per gram of spleen in guinea pigs inoculated with S2Δbp26 at different passages and S2, which suggested that the virulence of the mutant strain was attenuated. The mice inoculated with S2ΔBP26 had a shorter 50% recovery time than those inoculated with S2. Meanwhile, the mice immunized with S2Δbp26 could successfully resist the challenge with M28 wild-type strain at a dose of 2×105 CFU/mouse. [Conclusion] S2Δbp26 was successfully constructed, with excellent safety and immunoprotective capacity, which provided technical reserves for the development of identifiable vaccines for brucellosis.

, correspAuthors=Xiaowei PENG, Junping LI, authorNote=null, correspAuthorsNote=
*E-mail: LI Junping,
PENG Xiaowei,
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#These authors contributed equally to this work.

, authorsList=Shibo LIU, Xinyue TANG, Xiaoqian ZHANG, Yinghui ZHANG, Jihui JIN, Weifeng SUN, Xiaowei PENG, Junping LI), CN=ArticleExt(id=1228017379540660708, articleId=1228017375233110351, tenantId=1146029695717560320, journalId=1192105938417971205, language=CN, title=布鲁氏菌S2Δbp26缺失株的构建与生物学特性, columnId=1192149544164012138, journalTitle=微生物学报, columnName=研究报告, runingTitle=null, highlight=null, articleAbstract=

【目的】 研发一种安全、有效且可鉴别的新型疫苗。 【方法】 以S2作为亲本株，利用同源重组技术构建S2Δbp26基因缺失株，并对缺失菌株的稳定性、安全性和有效性进行全面评价。 【结果】 构建了S2Δbp26缺失株，该缺失株在体外连续传代20代后其表型和基因型均未发生改变。小鼠和豚鼠的安全性试验结果表明，缺失株在生物安全性方面与S2疫苗株无显著差异。不同代次的S2Δbp26缺失株和S2亲本株接种豚鼠后，每克脾脏分菌数量均小于2×105个，表明该缺失株毒力较低。残余毒力试验数据表明，接种缺失株的小鼠半数痊愈时间比S2亲本株更短。同时，免疫S2Δbp26菌株的小鼠可以成功抵御2×105 CFU/只剂量的M28野毒株攻毒。 【结论】 本研究构建的S2Δbp26基因缺失株展现出优异的安全性和免疫保护力，为布鲁氏菌病可鉴别诊断疫苗的研发提供了技术储备。

, correspAuthors=彭小薇, 李俊平, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=4Y5Dv6bJaCk/StFznw2yZw==, magXml=46pnRphliIsddt1REMxfVg==, pdfUrl=null, pdf=UuJlXDKO6ZRcpeZGhE9oAA==, pdfFileSize=2473057, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=pTu4EChPIu4iyfu4jO9ZFQ==, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=EPoZUVEEgd/riS7xrAhC9g==, mapNumber=null, authorCompany=null, fund=null, authors=

作者贡献声明

刘世博：文章撰写、数据分析、安全性试验；唐新月：缺失株构建、缺失株表型鉴定、安全性试验、残余毒力试验；张晓茜：安全性试验、免疫保护力试验；张莹辉：安全性试验、残余毒力试验；靳继惠：免疫保护力试验；孙伟峰：残余毒力试验；彭小薇：试验指导、文章修改；李俊平：试验设计、试验指导、文章修改。

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M5 (M5-90Δ26)株和S2株布氏杆菌疫苗对羊诱导的抗体消长动态规律及免疫效果研究[J]. 北方牧业, 2024(15): 13., articleTitle=M5 (M5-90Δ26)株和S2株布氏杆菌疫苗对羊诱导的抗体消长动态规律及免疫效果研究, refAbstract=null), Reference(id=1228088887831097923, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017375233110351, doi=null, pmid=null, pmcid=null, year=null, volume=2024, issue=15, pageStart=13, pageEnd=null, url=null, language=null, rfNumber=[24], rfOrder=39, authorNames=WEI JP, WANG HY, ZHANG HJ, SUN TR, LIU B, LIU YT, YUAN F, MA HW, HAN QA, journalName=North China Pastoral Industry Journal, refType=null, unstructuredReference=WEI JP, WANG HY, ZHANG HJ, SUN TR, LIU B, LIU YT, YUAN F, MA HW, HAN QA. Study on the dynamic law of antibody growth and decline induced by M5 (M5-90Δ26) strain and S2 strain Brucella vaccine in sheep and its immune effect[J]. 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A: The splenic bacteria load of mice on days 15; B: The splenic bacteria load of mice on days 45. **: P<0.01., figureFileSmall=cHu/CAPd7xTuv7iqL6j9RA==, figureFileBig=6/cXF95nJgOgGtqgvXauwg==, tableContent=null), ArticleFig(id=1228088877148205268, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017375233110351, language=CN, label=图8, caption=攻毒后小鼠脾脏载菌量, figureFileSmall=cHu/CAPd7xTuv7iqL6j9RA==, figureFileBig=6/cXF95nJgOgGtqgvXauwg==, tableContent=null), ArticleFig(id=1228088877261451488, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017375233110351, language=EN, label=Table 1, caption=

Primers related to the construction of bp26 gene deletion strain

, figureFileSmall=null, figureFileBig=null, tableContent=
Primers namePrimer sequences (5′→3′)Restriction sitesLenth of products
D-S2-Bp26F-FGGGGTACCTCAACGAGGCAGATTATGGKpn I523
D-S2-Bp26F-RAAGCGGGTTATGTCAGG-
D-S2-Bp26R-FAACCCGCTTGCCCTTTGCCACCTGAT-506
D-S2-Bp26R-RCCGGATCCGCTCAAATCGTTCGTCGTTBamH I
YZ1-Bp26-FGCAGCCTCATTTTCCACAA-441
YZ1-Bp26-RACCGCCCTGATTAACACC-
18Ap-FCTGCAAGGCGATTAAGTTGGGTAAC-1 193
18Ap-RTATGTTGTGTGGAATTGTGAGCGGA-
), ArticleFig(id=1228088877345337571, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017375233110351, language=CN, label=表1, caption=

构建 bp26 基因缺失株相关引物

, figureFileSmall=null, figureFileBig=null, tableContent=
Primers namePrimer sequences (5′→3′)Restriction sitesLenth of products
D-S2-Bp26F-FGGGGTACCTCAACGAGGCAGATTATGGKpn I523
D-S2-Bp26F-RAAGCGGGTTATGTCAGG-
D-S2-Bp26R-FAACCCGCTTGCCCTTTGCCACCTGAT-506
D-S2-Bp26R-RCCGGATCCGCTCAAATCGTTCGTCGTTBamH I
YZ1-Bp26-FGCAGCCTCATTTTCCACAA-441
YZ1-Bp26-RACCGCCCTGATTAACACC-
18Ap-FCTGCAAGGCGATTAAGTTGGGTAAC-1 193
18Ap-RTATGTTGTGTGGAATTGTGAGCGGA-
), ArticleFig(id=1228088877483749614, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017375233110351, language=EN, label=Table 2, caption=

The spleen bacteria load

, figureFileSmall=null, figureFileBig=null, tableContent=
StrainsBacterial load of spleen (CFU/g)
S23.56×104
S2Δbp26-F17.95×104
S2Δbp26-F52.19×104
S2Δbp26-F101.30×104
), ArticleFig(id=1228088877592801531, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017375233110351, language=CN, label=表2, caption=

脾脏载菌量

, figureFileSmall=null, figureFileBig=null, tableContent=
StrainsBacterial load of spleen (CFU/g)
S23.56×104
S2Δbp26-F17.95×104
S2Δbp26-F52.19×104
S2Δbp26-F101.30×104
), ArticleFig(id=1228088877693464835, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017375233110351, language=EN, label=Table 3, caption=

The recovery rate of mice and the calculation of 50% recovery time

, figureFileSmall=null, figureFileBig=null, tableContent=
StrainsInoculation dose (CFU/mice)Recovery rate (%)RT50 (weeks)
3 weeks5 weeks7 weeks9 weeks
S21×108268998994.08
S2Δbp26-F11×108569598992.62
), ArticleFig(id=1228088877823488266, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017375233110351, language=CN, label=表3, caption=

小鼠痊愈率及半数痊愈时间计算

, figureFileSmall=null, figureFileBig=null, tableContent=
StrainsInoculation dose (CFU/mice)Recovery rate (%)RT50 (weeks)
3 weeks5 weeks7 weeks9 weeks
S21×108268998994.08
S2Δbp26-F11×108569598992.62
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布鲁氏菌S2Δbp26缺失株的构建与生物学特性
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刘世博 1, 2 , 唐新月 1, 3 , 张晓茜 1 , 张莹辉 1 , 靳继惠 1 , 孙伟峰 1 , 彭小薇 1, * , 李俊平 1, *
微生物学报 | 研究报告 2025,65(3): 1137-1147
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微生物学报 | 研究报告 2025, 65(3): 1137-1147
布鲁氏菌S2Δbp26缺失株的构建与生物学特性
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刘世博1, 2, 唐新月1, 3, 张晓茜1, 张莹辉1, 靳继惠1, 孙伟峰1, 彭小薇1, * , 李俊平1, *
作者信息
  • 1 中国兽医药品监察所，国家动物布鲁氏菌病参考实验室，北京
  • 2 吉林农业大学 动物医学院，吉林 长春
  • 3 北京三元集团畜牧兽医总站，北京
Construction and biological characterization of Brucella S2Δbp26
Shibo LIU1, 2, Xinyue TANG1, 3, Xiaoqian ZHANG1, Yinghui ZHANG1, Jihui JIN1, Weifeng SUN1, Xiaowei PENG1, * , Junping LI1, *
Affiliations
  • 1 National Reference Laboratory for Animal Brucellosis, China Institute of Veterinary Drug Control, Beijing, China
  • 2 College of Animal Medicine, Jilin Agricultural University, Changchun, Jilin, China
  • 3 Animal Husbandry & Veterinary Station of Beijing SanYuan Group, Beijing, China
出版时间: 2025-03-04 doi: 10.13343/j.cnki.wsxb.20240645
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摘要
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【目的】 研发一种安全、有效且可鉴别的新型疫苗。 【方法】 以S2作为亲本株，利用同源重组技术构建S2Δbp26基因缺失株，并对缺失菌株的稳定性、安全性和有效性进行全面评价。 【结果】 构建了S2Δbp26缺失株，该缺失株在体外连续传代20代后其表型和基因型均未发生改变。小鼠和豚鼠的安全性试验结果表明，缺失株在生物安全性方面与S2疫苗株无显著差异。不同代次的S2Δbp26缺失株和S2亲本株接种豚鼠后，每克脾脏分菌数量均小于2×105个，表明该缺失株毒力较低。残余毒力试验数据表明，接种缺失株的小鼠半数痊愈时间比S2亲本株更短。同时，免疫S2Δbp26菌株的小鼠可以成功抵御2×105 CFU/只剂量的M28野毒株攻毒。 【结论】 本研究构建的S2Δbp26基因缺失株展现出优异的安全性和免疫保护力，为布鲁氏菌病可鉴别诊断疫苗的研发提供了技术储备。

布鲁氏菌  /  疫苗  /  S2Δbp26  /  生物学特性

[Objective] To develop a safe, effective, and identifiable new vaccine. [Methods] The gene deletion strain S2Δbp26 was constructed by homologous recombination with S2 as the parental strain. The stability, safety, and efficacy of S2Δbp26 were then evaluated. [Results] The phenotype and genotype of S2Δbp26 did not change after 20 successive passages invitro. The experiments in mice and guinea pigs showed that there was no difference in biosafety between S2Δbp26 and S2. The bacterial load was less than 2×105 CFU per gram of spleen in guinea pigs inoculated with S2Δbp26 at different passages and S2, which suggested that the virulence of the mutant strain was attenuated. The mice inoculated with S2ΔBP26 had a shorter 50% recovery time than those inoculated with S2. Meanwhile, the mice immunized with S2Δbp26 could successfully resist the challenge with M28 wild-type strain at a dose of 2×105 CFU/mouse. [Conclusion] S2Δbp26 was successfully constructed, with excellent safety and immunoprotective capacity, which provided technical reserves for the development of identifiable vaccines for brucellosis.

Brucella  /  vaccine  /  S2Δbp26  /  biological characterization
刘世博, 唐新月, 张晓茜, 张莹辉, 靳继惠, 孙伟峰, 彭小薇, 李俊平. 布鲁氏菌S2Δbp26缺失株的构建与生物学特性. 微生物学报, 2025 , 65 (3) : 1137 -1147 . DOI: 10.13343/j.cnki.wsxb.20240645
Shibo LIU, Xinyue TANG, Xiaoqian ZHANG, Yinghui ZHANG, Jihui JIN, Weifeng SUN, Xiaowei PENG, Junping LI. Construction and biological characterization of Brucella S2Δbp26[J]. Acta Microbiologica Sinica, 2025 , 65 (3) : 1137 -1147 . DOI: 10.13343/j.cnki.wsxb.20240645
正文
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布鲁氏菌病(brucellosis)是由胞内寄生的布鲁氏菌(Brucella spp.)感染引起的一种具有严重危害的人畜共患传染病[1]。该病主要引起动物流产、睾丸炎、关节炎等症状[2]；人感染布鲁氏菌后主要会出现波浪热、多汗、肌肉疼痛、乏力、全身不适等临床表现。该病在全球范围内170多个国家均有流行[3]。进入21世纪以来，由于我国畜牧养殖业的快速发展和活畜调运的便捷，布鲁氏菌病在我国迅速传播，严重影响了畜牧业的健康发展和人民群众的公共卫生安全[4]。
疫苗免疫是防控家畜布鲁氏菌病的有效措施之一，全球已有多个国家通过疫苗免疫实现了该病的有效控制。目前我国已批准使用的布鲁氏菌病疫苗共有7种，包括S2株、M5/M5-90株、M5-90Δ26株、A19株、A19ΔVirB12株、Rev.1株和BA0711株。这些均为弱毒活疫苗，对人体具有一定的感染性。S2株是中国兽医药品监察所专家于1952年从进口母猪流产胎儿中分离出的一株猪种布鲁氏菌，经过7年的人工致弱处理[5-6]。S2疫苗株在我国家畜布鲁氏菌病的防控中得到了广泛应用。与其他布鲁氏菌病疫苗相比，S2疫苗具有较高的安全性，可通过口服免疫方式避免引起怀孕动物流产[6-7]。然而，由于S2疫苗具有完整的脂多糖结构，导致家畜免疫后产生的自然感染抗体与免疫抗体无法有效区分。
BP26蛋白是一个可分泌的周质蛋白，生物信息学建模分析表明，其三级结构呈现为通道状组件，由16个BP26分子构成，其中8个分子连接成环形成八聚体，而2个八聚体相互作用使环中心形成一个大的内腔通道。此结构与参与感染的噬菌体蛋白具有相似性，预示着BP26蛋白可能参与布鲁氏菌的感染过程[2]。BP26蛋白在布鲁氏菌中高度保守，能够刺激保护性T细胞产生迟发型变态反应，是一种可诱导产生抗体的优势抗原[8]。同时，BP26蛋白还具有区分感染动物与免疫动物血清学反应的潜力[9]。目前，针对BP26蛋白的布鲁氏菌病检测方法众多，其作为优势抗原诱导产生高滴度抗体的能力已得到广泛认可。
1 材料与方法
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1.1 材料
B. suis S2菌株、粗糙型布鲁氏菌RM6/66株、羊种布鲁氏菌B. melitensis M28株均由国家兽医微生物菌(毒)种保藏中心鉴定保存；同源重组质粒pEX18AP (蔗糖诱导的自杀质粒)由本实验室保存；大肠埃希氏菌DH5α和BL21(DE3)感受态细胞均购自北京索莱宝科技有限公司；小鼠布鲁氏菌病抗体阴性血清、布鲁氏菌病抗体阳性血清、BP26蛋白多抗阳性血清均由本实验室保存；限制性内切酶Kpn I和BamH I均购自NEB公司；胰蛋白胨大豆琼脂培养基(tryptone soy agar, TSA)和胰蛋白胨大豆肉汤培养基(tryptone soy broth, TSB)均购自BD公司；质粒小量提取试剂盒和琼脂糖凝胶DNA回收试剂盒均购自Omega公司；细菌基因组提取试剂盒购自西安天隆科技有限公司。6-8周龄的雌性BALB/c小鼠及350-400 g雌性豚鼠均购自北京维通利华实验动物技术有限公司。
1.2 引物设计与合成
布鲁氏菌S2染色体I基因组在NCBI网站的序列号为CP006962，其上包含bp26基因(BSS2_I1433)。根据基因序列设计上、下游同源臂引物(D-S2-Bp26F-F/R和D-S2-Bp26R-F/R)、鉴定筛选引物(YZ1-Bp26-F/R)以及bp26基因载体鉴定引物(18Ap-F/R)，所有引物均由生工生物工程(上海)股份有限公司合成，引物序列见表1。
1.3 同源重组质粒pEX18AP-bp26-DFR的构建
取保存的冻干B. suis S2菌株溶解后，于TSA平板上划线，37 ℃培养48 h后挑取复苏的单菌落，接种于15 mL TSB培养基培养至对数期。使用细菌基因组提取试剂盒提取S2基因组。以该基因组为模板，分别使用引物D-S2-Bp26F-F/R和D-S2-Bp26R-F/R，通过PCR扩增bp26基因的上、下游片段。PCR反应体系(50 µL)：2×PrimeSTAR Max Premix 25 µL，上、下游引物(10 µmol/L)各1 µL，S2 DNA模板1 µL，ddH2O 22 µL。PCR反应条件：95 ℃预变性5 min；98 ℃变性10 s，60 ℃退火5 s，72 ℃延伸5 s，30个循环；72 ℃终延伸10 min。切取目的产物大小位置的琼脂糖电泳凝胶后，按照琼脂糖凝胶DNA回收试剂盒说明书对扩增产物进行纯化回收。以胶回收纯化后的DNA产物为模板，通过PCR扩增将上、下游同源臂进行融合。PCR反应体系(50 µL)：2×Premix TaqTM 25 µL，上、下游同源臂各1 µL，上、下游引物(10 µmol/L)各1 µL、ddH2O 21 µL。PCR反应条件：94 ℃预变性2 min；94 ℃变性30 s，60 ℃退火5 s，72 ℃延伸1 min，每个循环退火温度下降0.5 ℃，共进行20个循环；94 ℃变性30 s，50 ℃退火5 s，72 ℃延伸1 min，共进行10个循环；72 ℃终延伸10 min。切取目的产物大小位置的琼脂糖电泳凝胶后，使用核酸胶回收试剂盒纯化回收bp26基因的上、下游融合片段。
将pEX18AP质粒和bp26基因的上、下游融合片段在37 ℃用Kpn I和BamH I内切酶双酶切4 h。胶回收纯化质粒和片段的酶切产物，在16 ℃用连接酶连接2 h后，转入DH5α感受态细胞中。提取质粒，得到同源重组质粒pEX18AP-p26-DFR。
1.4 布鲁氏菌S2Δbp26缺失株的构建与筛选
取10 μL同源重组质粒pEX18AP-bp26-DFR，加入90 μL制备的布鲁氏菌S2株感受态细胞中混匀，以2 200 V、200 Ω的条件进行电转。电转后迅速向电击杯中加入1 mL无抗TSB，轻柔吹打混匀后将所有液体移入无菌EP管中，于37 ℃、180 r/min复壮培养12 h。将复壮的菌液在常温下以8 000 r/min离心8 min后重悬菌体，在含氨苄青霉素的TSA培养基上37 ℃培养48 h，进行第1次单交换筛选。同时，在含氨苄青霉素的TSA培养基和5%蔗糖的无抗性TSA培养基上随机选择长出的单菌落进行37 ℃双划线培养48 h。挑取仅在含氨苄青霉素的TSA培养基上生长的单菌落，在TSB培养基中培养24 h。以相同方式离心重悬菌体并进行1 000倍稀释，将稀释液在含5%蔗糖的TSA培养基上37 ℃培养24 h，挑取单菌落后同样进行双划线培养。再次挑取仅在含5%蔗糖的TSA培养基上生长的单菌落使用YZ1-Bp26-F/R引物进行PCR扩增鉴定。PCR反应体系(25 µL)：2×Premix Taq™ 12.5 µL，上、下游引物(10 µmol/L)各1 µL，灭活菌液模板1 µL，ddH2O 9.5 µL。PCR反应条件：94 ℃预变性5 min；94 ℃变性30 s，60 ℃退火30 s，72 ℃延伸1 min，30个循环；72 ℃终延伸10 min。将鉴定阳性的菌株命名为S2Δbp26，并连续培养至第20代，每隔5代以相同体系及步骤进行PCR验证，并将各代次菌株均于-80 ℃保存。
1.5 生长曲线测定及表型鉴定结果
取S2和S2Δbp26缺失株分别接种于无抗性TSB培养液中，在37 ℃、200 r/min培养24 h后进行平板计数。将菌液稀释至2.7×105 CFU/mL后转接入无抗性TSB培养液中，以相同条件在摇床中培养。从转接时开始计时，每隔2 h吸取1 mL菌液于酶标仪中测定600 nm处的吸光度，直至第30 h。取S2、S2Δbp26-F1、S2Δbp26-F5、S2Δbp26-F10、S2Δbp26-F15、S2Δbp26-F20株，以及实验室保存的M111菌株，通过结晶紫染色及吖啶黄试验观察细菌形态[10-11]。
1.6 安全性试验
按照《中华人民共和国兽用生物制品规程》(2000年版)[12]要求，将20只18-20 g的雌性BALB/c小鼠随机平均分为4组，每组5只。分别在小鼠的腹股沟皮下注射0.25 mL的S2Δbp26-F1、S2Δbp26-F5、S2Δbp26-F10株和蛋白胨水，菌株接种剂量为每只2.5×108 CFU。接种后对小鼠的精神状态、食欲、饮水、体重、背毛、呼吸、粪便和行动观察6 d。
按照WOAH Terrestrial Manual 2022 Chapter 3.1.4[13]要求，将35只350-400 g的雌性豚鼠随机分为4组，实验组每组10只，空白对照组5只。分别在豚鼠的腿部肌肉处注射1 mL的S2Δbp26-F1、S2Δbp26-F5、S2Δbp26-F10株和蛋白胨水，菌株接种剂量为每只5×109 CFU。接种后对豚鼠的精神状态、食欲、饮水、体重、背毛、呼吸、粪便和行动观察14 d。
1.7 毒力试验
按照《中华人民共和国兽用生物制品规程》(2000年版)[12]相关要求，将20只350-400 g的雌性豚鼠随机平均分为4组，每组5只。分别在豚鼠的腹股沟皮下位置注射1 mL的S2、S2Δbp26-F1、S2Δbp26-F5和S2Δbp26-F10株。接种剂量为每只1×109 CFU。在接种后第15天对豚鼠进行高浓度CO2处死。将处死的豚鼠用75%乙醇浸泡30 s后取出，从腹腔左侧剪开外层皮肤和内层肌肉组织，将脾脏剪下并剥离脂肪和系膜后称重记录，全程保证无菌操作。将脾脏放入离心管中，加入2倍脾质量的0.1%蛋白胨水，将脾脏研磨成匀浆。对匀浆进行10倍梯度稀释，前3个梯度中每个梯度取100 μL匀浆溶液涂布在胰蛋白胨琼脂培养基中，进行3个重复。将涂布后的培养基置于37 ℃恒温培养72 h后数菌计数，计算平均值。
1.8 残余毒力试验(50%痊愈时间)
按照WOAH Terrestrial Manual 2022 Chapter 3.1.4[13]相关要求，将80只6周龄的雌性BALB/c 小鼠随机平均分为2组，每组40只。分别在小鼠的腹股沟皮下注射0.1 mL的1×109 CFU/mL剂量的S2疫苗株和S2Δbp26-F1菌株。在接种后的第3、5、7和9周，每组取10只小鼠摘眼球采血后脱颈处死，用75%乙醇浸泡30 s后取出，从腹腔左侧剪开外层皮肤和内层肌肉组织，将脾脏剪下并剥离脂肪和系膜后称重记录，全程保证无菌操作。向每个装有脾脏并做好标记的离心管中加入1 mL无菌0.1%蛋白胨水，研磨成匀浆，全部涂布在胰蛋白胨琼脂培养基中，培养3-5 d后记录脾脏分菌情况，并根据文献[14]报道的方法计算小鼠半数痊愈时间。
1.9 免疫保护力试验
将36只6周龄的雌性BALB/c小鼠随机平均分为6组，每组6只。按照WOAH Terrestrial Manual 2022 Chapter 3.1.4[13]推荐的免疫剂量、方式及攻毒剂量，分别在小鼠的腹股沟皮下注射0.1 mL的S2株和S2Δbp26-F1株，菌株免疫剂量分别为每只1×105 CFU，空白组注射0.1 mL的0.1%蛋白胨水。小鼠免疫30 d后使用0.1 mL的M28株进行腹腔内攻毒，剂量为每只2×105 CFU，攻毒前应使小鼠禁食禁水16 h。在攻毒后第15天和第45天，每种不同菌株和空白对照均随机取1组小鼠，脱颈处死后在75%乙醇中浸泡30 s，取出后从腹腔左侧剪开外层皮肤和内层肌肉组织，将脾脏剪下并剥离脂肪和系膜后称重记录。将脾脏分别装入离心管中并做好标记，向每个离心管中加入脾重9倍的0.1%蛋白胨水，研磨成匀浆。将组织匀浆进行10倍倍比稀释(10、100、1 000倍)，并各取100 µL涂布于TSA培养基上，37 ℃培养2-3 d后，选择菌落数少于300 CFU的平板进行菌落计数。
1.10 统计学分析
使用SPSS 17.0软件中的独立样本t-检验方法，分析试验组和对照组之间的平均差异。P<0.05被认为具有显著差异。
1.11 动物实验的场所和伦理审查说明
所有涉及布鲁氏菌疫苗株的操作(安全性试验、毒力试验、残余毒力试验)均在中国兽医药品监察所ABSL-2实验室内进行；涉及布鲁氏菌强毒株的操作(免疫保护力试验)在中国兽医药品监察所ABSL-3实验室内进行。
本研究方案已通过中国兽医药品监察所实验动物福利伦理审查委员会的审查，实验动物福利伦理审查编号分别为中监所(福)2023第00243号、中监所(福)2023第00244号、中监所(福)2023第00245号、中监所(福)2023第00246号。
2 结果与分析
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2.1 同源重组质粒pEX18AP-bp26-DFR的构建与鉴定结果
以提取的S2基因组为模板，通过PCR扩增得到523 bp的上游同源臂和506 bp的下游同源臂；以上、下游同源臂为模板，通过连接PCR扩增获得1 020 bp的融合片段，并将该融合片段克隆至pEX18AP载体中。经过Amp抗性筛选、PCR鉴定和双酶切验证，获得阳性重组质粒，命名为pEX18AP-bp26-DFR。阳性重组质粒的双酶切验证结果如图1所示。
2.2 布鲁氏菌S2Δbp26缺失株的筛选与遗传稳定性鉴定结果
将同源重组质粒pEX18AP-bp26-DFR电转化至S2菌株中，最终获得无Amp抗性，且能在含蔗糖培养基上生长的bp26基因缺失株，命名为S2Δbp26。采用引物YZ1-Bp26-F和YZ1-Bp26-R对5个S2Δbp26阳性克隆进行PCR鉴定。结果如图2所示，亲本S2疫苗株可以扩增出441 bp的片段，而缺失菌株无条带。
为了验证S2Δbp26的遗传稳定性，选取S2Δbp26缺失株的1号克隆进行连续传代，并每隔5个代次采用引物YZ1-Bp26-F和YZ1-Bp26-R进行PCR验证。图3结果显示，缺失株各代次均无条带，而亲本S2株可以扩增出441 bp的片段，表明S2Δbp26缺失株在体外连续传代20代时，其基因型遗传稳定。将得到的菌株于-80 ℃冷冻保存。
2.3 生长曲线测定与表型鉴定结果
在TSB正常培养环境和初始菌株浓度相同的情况下，每隔2 h对S2株和S2Δbp26缺失株在600 nm处的吸光度进行测量。结果表明，在各个不同时间点，2个菌株的生长趋势基本一致，见图4。
根据脂多糖是否含有完整的O链可将布鲁氏菌分为光滑型和粗糙型。使用结晶紫对细菌进行染色时，光滑型细菌不着色，保持白色菌落形态；而粗糙型细菌则被染成紫色。在吖啶黄试验中，光滑型细菌不形成沉淀，保持浑浊状态；而粗糙型细菌则因自凝现象形成沉淀[15-16]。本研究结果表明，S2Δbp26缺失株各代次在结晶紫染色中均未着色，且在吖啶黄试验中均未出现沉淀(图5、6)，这说明S2Δbp26缺失株的脂多糖结构完整，呈现光滑型表型，且各代次表型遗传稳定。
2.4 安全性试验结果
根据《中华人民共和国兽用生物制品规程》(2000年版)[12]及WOAH Terrestrial Manual 2022 Chapter 3.1.4[13]的相关要求，对接种S2株和S2Δbp26缺失株的小鼠和豚鼠进行观察，这些动物的精神状态、食欲、饮水、体重、背毛、呼吸、粪便和行动均未出现异常。小鼠在6 d内全部健康存活，豚鼠14 d内全部健康存活。这说明S2株和S2Δbp26缺失株均具有良好的安全性。
2.5 毒力试验结果
分别对接种S2株和S2Δbp26缺失株各代次15 d后的豚鼠脾脏质量和分菌情况进行评估，结果见图7和表2。接种15 d后，各组豚鼠脾脏质量无明显差异。分菌结果显示，豚鼠每克脾脏载菌量均小于2×105个，表明缺失株的毒力较低，符合《中华人民共和国兽用生物制品规程》(2000年版)[12]对疫苗株毒力的要求。
2.6 残余毒力试验结果
使用S2株和S2Δbp26-F1株接种6周龄BALB/c雌性小鼠，分别在接种后的第3周、第5周、第7周和第9周扑杀小鼠并无菌取脾脏，称重后进行研磨涂板。统计每周扑杀时脾脏无菌的小鼠数量，并根据WOAH Terrestrial Manual 2022 Chapter 3.1.4[13]中的方法，统计随时间累计的小鼠痊愈数量，计算半数小鼠痊愈时间(RT50)，结果见表3。结果显示，随时间增加小鼠痊愈率升高，在第7周和第9周时，所有小鼠脾脏均未分离出布鲁氏菌。统计结果表明缺失株小鼠的半数痊愈时间早于亲本株。
2.7 免疫保护力试验结果
为探究bp26基因缺失对菌株免疫保护能力的影响，使用S2株和S2Δbp26-F1株免疫6周龄BALB/c雌性小鼠，免疫30 d后用M28强毒株进行攻毒。攻毒后第15天和第45天小鼠脾脏载菌量结果如图8所示。在攻毒后第15天及第45天，免疫S2疫苗株和S2Δbp26-F1菌株的2组小鼠脾脏均未分离到细菌。与空白对照组相比，各免疫组小鼠的脾脏含菌量差异均具有统计学意义(P<0.01)。以上结果表明，S2疫苗株与S2Δbp26-F1菌株免疫后均能有效保护小鼠免受2×105剂量的布鲁氏菌野毒株攻击。
3 讨论与结论
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从布鲁氏菌病疫苗生产的批签发数据来看，S2疫苗株是我国使用最为广泛的一种疫苗，其安全性和免疫保护能力均表现良好，可用于接种猪、牛和羊，其是全球唯一可用于免疫猪的布鲁氏菌病弱毒疫苗。S2疫苗株能通过口服免疫途径为猪、牛和羊提供有效的免疫保护，其优势在于口服免疫不会导致母畜流产。因其低残留毒力、高安全性、接种便利以及高效的保护力，S2疫苗在中国及其他国家被广泛应用于家畜的布鲁氏菌病免疫接种。然而，尽管S2疫苗在控制布鲁氏菌病方面取得了显著成效，但它也存在一定的局限性，即其完整的脂多糖结构会干扰感染动物与免疫动物的血清学诊断[17]。
BP26蛋白是布鲁氏菌的一种优势抗原，在布鲁氏菌中高度保守，并能刺激保护性T细胞产生迟发型变态反应。张瑞慧[18]将BP26蛋白作为包被抗原建立了间接ELISA检测方法，该方法具有良好的敏感性和特异性。孙田华等[19]以BP26为包被抗原建立了胶体金试纸检测方法，为饲养场大规模检测提供了更为便捷有效的检测途径。许菲菲等[20]利用纯化的BP26蛋白标记胶体金制成金标抗原，成功研制出一种具有良好特异性和灵敏度的羊布鲁氏菌抗体金标检测试纸条。郭晓涵等[21]通过对比不同布鲁氏菌病免疫层析胶体金抗体试纸条，发现F14和BP26试纸条具有更强优势。除了血清学检测方法外，马世辉等[22]还基于bp26基因建立了环介导等温扩增检测法，该方法通过肉眼即可直观地进行可视化判断，为一线生产快速检测提供了新思路。
在原有布鲁氏菌病疫苗株基础上，通过缺失免疫原性蛋白编码基因构建新型基因缺失疫苗，是目前研发鉴别诊断疫苗的一种有效思路。例如，由天康生物股份有限公司和新疆畜牧科学院兽医研究所联合研发的布氏病菌活疫苗A19-Δvirb12疫苗，以及中国农业科学院哈尔滨兽医研究所开发的M5-90Δ26疫苗，均是在原有的A19疫苗和M5-90疫苗基础上，通过缺失免疫原性蛋白构建而成的。其中，M5-90Δ26疫苗与本研究中使用的缺失株缺失的是同一基因。上述2种疫苗均已获得新兽药证书，并已开始在我国应用于家畜布鲁氏菌病的防控工作中[23-24]。据M5-90Δ26产品说明书介绍，动物免疫M5-90Δ26后，血清虎红平板凝集试验结果呈阳性。在免疫后15-180日内，利用缺失株免疫的家畜不产生针对Bp26蛋白抗体的原理建立间接Bp26-ELISA (Bp26-iELISA)检测方法，可区分本疫苗免疫与野毒感染，该方法具有足够的敏感性和特异性。
在本研究中，为了破解S2疫苗难以进行血清学鉴别诊断的难题，对S2疫苗株的bp26基因进行了缺失，成功构建了S2Δbp26缺失株。与亲本菌株S2相比，该缺失株在体外培养条件下的生长速度无显著差异。体外连续传代结果显示，构建的缺失株基因型和表型均具有良好的稳定性。随后，按照《中华人民共和国兽用生物制品规程》(2000年版)[12]、WOAHTerrestrialManual2022[13]等规程的要求，分别采用安全性试验、毒力试验、残余毒力试验和免疫原性试验等对该缺失株进行了全面的评价。结果显示，S2Δbp26缺失株的安全性与免疫保护力均与S2疫苗株无显著性差异。这说明该缺失株具备作为兽用布鲁氏菌病疫苗候选株的潜力。与其他前人关于布鲁氏菌病疫苗候选株的研究相比，本研究对缺失株的生物学特性、安全性和有效性的评价更加全面和深入，所得结果也更加严谨和科学。
基金
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  • 国家重点研发计划(2022YFD1800600)
  • 国家重点研发计划(2022YFD1800601)
文献
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参考文献 引证文献
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2025年第65卷第3期
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doi: 10.13343/j.cnki.wsxb.20240645
  • 接收时间：2024-10-16
  • 首发时间：2026-02-10
  • 出版时间：2025-03-04
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  • 收稿日期：2024-10-16
  • 录用日期：2024-11-20
基金
National Key Research and Development Program of China(2022YFD1800600)
国家重点研发计划(2022YFD1800600)
National Key Research and Development Program of China(2022YFD1800601)
国家重点研发计划(2022YFD1800601)
作者信息
    1 中国兽医药品监察所，国家动物布鲁氏菌病参考实验室，北京
    2 吉林农业大学 动物医学院，吉林 长春
    3 北京三元集团畜牧兽医总站，北京

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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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