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Article(id=1297571024070004824, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1297570992835023717, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20260128, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1770652800000, receivedDateStr=2026-02-10, revisedDate=null, revisedDateStr=null, acceptedDate=1774368000000, acceptedDateStr=2026-03-25, onlineDate=1787294639841, onlineDateStr=2026-08-21, pubDate=1785772800000, pubDateStr=2026-08-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1787294639841, onlineIssueDateStr=2026-08-21, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1787294639841, creator=13701087609, updateTime=1787294639841, updator=13701087609, issue=Issue{id=1297570992835023717, tenantId=1146029695717560320, journalId=1192105938417971205, year='2026', volume='66', issue='8', pageStart='3681', pageEnd='4288', issueExtLink='null', onlineDate='null', pubDate='1785772800000', pubDateStr='2026-08-04', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1787294632395, creator='13701087609', updateTime=1787294931551, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1297572247670124783, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1297570992835023717, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1297572247670124784, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1297570992835023717, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=3994, endPage=4007, ext={EN=ArticleExt(id=1297571024292302937, articleId=1297571024070004824, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Isolation, identification, and pathogenicity analysis of a porcine deltacoronavirus strain CHN/GSYD/2023, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

Porcine deltacoronavirus (PDCoV), a major porcine intestinal coronavirus, has caused enormous economic losses to the pig industry. Up to now, there is no commercial vaccine available. [Objective] To obtain the epidemic strain of PDCoV and study its biological characteristics and pathogenicity, thereby providing biological materials for the development of an effective vaccine against PDCoV as soon as possible. [Methods] LLC-PK1 cells were used to isolate the virus from PDCoV-positive samples from a pig farm in Gansu Province. The virus was identified by observation of cytopathic effect (CPE), indirect immunofluorescence assay (IFA), and whole genome sequencing. [Results] After blind passage of the PDCoV-positive LLC-PK1 cells to P6, the cells became larger and rounded, and then shrunk and shed into single or clustered dense particles, typical features of CPE. The IFA results confirmed successful isolation of the PDCoV strain CHN/GSYD/2023. The whole genome sequencing and phylogenetic analysis showed that CHN/GSYD/2023 isolated in this study was more closely related to the strain from China and was located on a different clade from the strain CHN/XJYN/2016 preserved in our laboratory. After 5-day-old piglets were infected with CHN/GSYD/2023, 100% (5/5) of piglets became diseased. Autopsy and histopathology of piglets showed thinning small intestinal wall, a large amount of effusion in the intestinal cavity, ileum intestinal villus necrosis and other pathological damage. [Conclusion] A PDCoV strain CHN/GSYD/2023 was successfully isolated from PDCoV-positive samples in a pig farm in Gansu. The strain can be stably proliferated and passaged in LLC-PK1 cells, and it is highly pathogenic to suckling piglets.

, authors=Ruiming YU1, Liping ZHANG2, Peng ZHOU2, Zhongwang ZHANG2, Li PAN2, Huichen GUO2, Ligang YUAN1, Xinsheng LIU2, authorsList=Ruiming YU, Liping ZHANG, Peng ZHOU, Zhongwang ZHANG, Li PAN, Huichen GUO, Ligang YUAN, Xinsheng LIU, authorCompany=null, correspAuthors=Ligang YUAN, Xinsheng LIU, authorNote=null, correspAuthorsNote=
E-mail: YUAN Ligang, ;
LIU Xinsheng,
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猪德尔塔冠状病毒(porcine deltacoronavirus, PDCoV)是一种重要的猪肠道冠状病毒，给养猪业带来了重大经济损失，截至目前尚无可用的商品化疫苗。 【目的】 获得PDCoV流行毒株，并对其生物学特性和致病性进行初步研究，为尽快研发出有效防控PDCoV的疫苗提供生物材料。 【方法】 使用LLC-PK1细胞分离来自甘肃某猪场PDCoV阳性病料中的病毒，并通过观察细胞病变效应(cytopathic effect, CPE)、间接免疫荧光试验(indirect immunofluorescence assay, IFA)、病毒全基因组测序等方法进行鉴定，同时评价分离毒株在仔猪上的致病性。 【结果】 PDCoV阳性病料接种LLC-PK1细胞后，连续盲传至第6代(passage 6, P6)，细胞出现变大、变圆、皱缩脱落成单个或成簇密集颗粒的典型CPE；IFA鉴定PDCoV阳性，表明成功分离出PDCoV CHN/GSYD/2023毒株。全基因组序列与系统发育分析表明，本研究分离的CHN/GSYD/2023毒株与中国毒株亲缘关系更近，与本实验室保存的CHN/XJYN/2016毒株位于不同的进化分支上；5日龄仔猪感染CHN/GSYD/2023毒株后，仔猪发病率为100% (5/5)。剖检及组织病理学显示，仔猪小肠肠壁变薄，肠腔内含大量积液，回肠绒毛坏死等病理损伤。 【结论】 本研究从甘肃地区某猪场PDCoV阳性样品中成功分离出PDCoV CHN/GSYD/2023毒株。该毒株可在LLC-PK1细胞中稳定增殖传代，对哺乳仔猪具有高致病性。

, authors=于瑞明1, 张莉萍2, 周鹏2, 张中旺2, 潘丽2, 郭慧琛2, 袁莉刚1, 刘新生2, authorsList=于瑞明, 张莉萍, 周鹏, 张中旺, 潘丽, 郭慧琛, 袁莉刚, 刘新生, authorCompany=null, correspAuthors=袁莉刚, 刘新生, authorNote=

作者贡献声明

于瑞明：实验设计、实验操作、初稿写作；张莉萍：数据收集和处理；周鹏：数据分析、动物实验操作；张中旺：数据分析及文献查找；潘丽、郭慧琛：监督指导、经费支持；袁莉刚：监督指导、稿件润色修改；刘新生：数据分析、经费支持、稿件润色修改。

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Journal of Clinical Microbiology, 2015, 53(5): 1537-1548., articleTitle=Isolation and characterization of porcine deltacoronavirus from pigs with diarrhea in the United States, refAbstract=null), Reference(id=1297571036334149838, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, doi=null, pmid=null, pmcid=null, year=2018, volume=33, issue=2, pageStart=131, pageEnd=141, url=null, language=null, rfNumber=[21], rfOrder=20, authorNames=Xu ZC, Zhong HL, Zhou QF, Du YP, Chen L, Zhang Y, Xue CY, Cao YC, journalName=Virologica Sinica, refType=null, unstructuredReference=Xu ZC, Zhong HL, Zhou QF, Du YP, Chen L, Zhang Y, Xue CY, Cao YC. A highly pathogenic strain of porcine deltacoronavirus caused watery diarrhea in newborn piglets[J]. Virologica Sinica, 2018, 33(2): 131-141., articleTitle=A highly pathogenic strain of porcine deltacoronavirus caused watery diarrhea in newborn piglets, refAbstract=null), Reference(id=1297571036401258703, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, doi=null, pmid=null, pmcid=null, year=2020, volume=37, issue=9, pageStart=2641, pageEnd=2654, url=null, language=null, rfNumber=[22], rfOrder=21, authorNames=He WT, Ji X, He W, Dellicour S, Wang SL, Li GR, Zhang LT, Gilbert M, Zhu HN, Xing G, Veit M, Huang Z, Han GZ, Huang YW, Suchard MA, Baele G, Lemey P, Su S, journalName=Molecular Biology and Evolution, refType=null, unstructuredReference=He WT, Ji X, He W, Dellicour S, Wang SL, Li GR, Zhang LT, Gilbert M, Zhu HN, Xing G, Veit M, Huang Z, Han GZ, Huang YW, Suchard MA, Baele G, Lemey P, Su S. Genomic epidemiology, evolution, and transmission dynamics of porcine deltacoronavirus[J]. 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Microorganisms, 2024, 12(2): 416., articleTitle=Genetic and evolutionary analysis of porcine deltacoronavirus in Guangxi Province, Southern China, from 2020 to 2023, refAbstract=null), Reference(id=1297571036879409361, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, doi=null, pmid=null, pmcid=null, year=2025, volume=16, issue=null, pageStart=2446742, pageEnd=null, url=null, language=null, rfNumber=[24], rfOrder=23, authorNames=Yao X, Lu WH, Qiao WT, Zhang YQ, Zhang BY, Li HX, Li JL, journalName=Virulence, refType=null, unstructuredReference=Yao X, Lu WH, Qiao WT, Zhang YQ, Zhang BY, Li HX, Li JL. The highly pathogenic strain of porcine deltacoronavirus disrupts the intestinal barrier and causes diarrhea in newborn piglets[J]. Virulence, 2025, 16: 2446742., articleTitle=The highly pathogenic strain of porcine deltacoronavirus disrupts the intestinal barrier and causes diarrhea in newborn piglets, refAbstract=null), Reference(id=1297571037097513170, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, doi=null, pmid=null, pmcid=null, year=2026, volume=17, issue=null, pageStart=2605366, pageEnd=null, url=null, language=null, rfNumber=[25], rfOrder=24, authorNames=Wang YN, Ma XY, Zhang H, Xue XH, Li JX, Jiang YP, Cui W, Liu DD, Zhou B, Wang L, journalName=Virulence, refType=null, unstructuredReference=Wang YN, Ma XY, Zhang H, Xue XH, Li JX, Jiang YP, Cui W, Liu DD, Zhou B, Wang L. Molecular characterization and pathogenicity of a novel Chinese porcine deltacoronavirus strain CH/HLJ/20 isolated from diarrheic piglets[J]. Virulence, 2026, 17: 2605366., articleTitle=Molecular characterization and pathogenicity of a novel Chinese porcine deltacoronavirus strain CH/HLJ/20 isolated from diarrheic piglets, refAbstract=null), Reference(id=1297571037214953683, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, doi=null, pmid=null, pmcid=null, year=2025, volume=2025, issue=null, pageStart=4084814, pageEnd=null, url=null, language=null, rfNumber=[26], rfOrder=25, authorNames=Zhang YQ, Wang B, Wei WQ, Lu W, Du EZ, Liu Y, Yang YL, Huang YW, journalName=Transboundary and Emerging Diseases, refType=null, unstructuredReference=Zhang YQ, Wang B, Wei WQ, Lu W, Du EZ, Liu Y, Yang YL, Huang YW. Genetic and pathogenic characterization of a porcine deltacoronavirus strain isolated in Zhejiang Province, China[J]. Transboundary and Emerging Diseases, 2025, 2025: 4084814., articleTitle=Genetic and pathogenic characterization of a porcine deltacoronavirus strain isolated in Zhejiang Province, China, refAbstract=null)], funds=[Fund(id=1297571033935007924, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, awardId=NCTIP-XD/C 03, language=EN, fundingSource=National Center of Technology Innovation for Pigs Project(NCTIP-XD/C 03), fundOrder=null, country=null), Fund(id=1297571034002116789, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, awardId=NCTIP-XD/C 03, language=CN, fundingSource=国家生猪技术创新中心项目(NCTIP-XD/C 03), fundOrder=null, country=null), Fund(id=1297571034060837046, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, awardId=24ZD13NA008, language=EN, fundingSource=Major Science and Technology Project of Gansu Province(24ZD13NA008), fundOrder=null, country=null), Fund(id=1297571034161500343, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, awardId=24ZD13NA008, language=CN, fundingSource=甘肃省科技重大专项(24ZD13NA008), fundOrder=null, country=null), Fund(id=1297571034220220600, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, awardId=23ZDNA007, language=EN, fundingSource=Major Science and Technology Project of Gansu Province(23ZDNA007), fundOrder=null, country=null), Fund(id=1297571034283135161, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, awardId=23ZDNA007, language=CN, fundingSource=甘肃省科技重大专项(23ZDNA007), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1297571027698077803, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, xref=1., ext=[AuthorCompanyExt(id=1297571027706466412, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, companyId=1297571027698077803, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1.College of Veterinary Medicine, Gansu Agricultural University, Lanzhou, Gansu, China), AuthorCompanyExt(id=1297571027714855021, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, companyId=1297571027698077803, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1.甘肃农业大学 动物医学院，甘肃 兰州)]), AuthorCompany(id=1297571027786158190, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, xref=2., ext=[AuthorCompanyExt(id=1297571027794546799, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, companyId=1297571027786158190, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2.State Key Laboratory of Animal Disease Control and Prevention, Lanzhou Veterinary Research Institute, Chinese Academy for Agricultural Sciences, Lanzhou, Gansu, China), AuthorCompanyExt(id=1297571027798741104, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, companyId=1297571027786158190, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2.中国农业科学院兰州兽医研究所，动物疫病防控全国重点实验室，甘肃 兰州)])], figs=[ArticleFig(id=1297571032177594528, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=EN, label=Figure 1, caption=Isolation and identification of PDCoV CHN/GSYD/2023 strain. A-C: The CPE of PDCoV CHN/GSYD/2023 P5, P6 and P7 was observed under a microscope (200×); D-F: After LLCPK1 cells were infected with PDCoV CHN/GSYD/2023 P5, P6 and P7 for 24 h. Mouse anti-PDCoV S protein monoclonal antibody was used as the primary antibody, and Alexa Fluor 488-labeled goat anti-mouse IgG was used as the secondary antibody to detect PDCoV antigen. The green fluorescence signal was observed under a fluorescence microscope., figureFileSmall=LodVr/nAFHOdyk6B2bLcTw==, figureFileBig=SEl+qKjx3671SZRkazta2A==, tableContent=null), ArticleFig(id=1297571032248897697, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=CN, label=图1, caption=PDCoV CHN/GSYD/2023毒株的分离与鉴定, figureFileSmall=LodVr/nAFHOdyk6B2bLcTw==, figureFileBig=SEl+qKjx3671SZRkazta2A==, tableContent=null), ArticleFig(id=1297571032483778722, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=EN, label=Figure 2, caption=Proliferation kinetics of PDCoV CHN/GSYD/2023 strain in LLC-PK1 cells., figureFileSmall=+SayU1whNlkPHqWyZXYTgg==, figureFileBig=ZUH7LmU6iBc+7WhL6bjI+Q==, tableContent=null), ArticleFig(id=1297571032542498979, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=CN, label=图2, caption=PDCoV CHN/GSYD/2023毒株在LLC-PK1细胞中的增殖动力学, figureFileSmall=+SayU1whNlkPHqWyZXYTgg==, figureFileBig=ZUH7LmU6iBc+7WhL6bjI+Q==, tableContent=null), ArticleFig(id=1297571032634773668, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=EN, label=Figure 3, caption=Phylogenetic tree analysis of PDCoV CHN/GSYD/2023 strain. The phylogenetic tree was constructed based on multiple sequence alignment, and the neighbor-joining method was used for inference. The number on each node in the figure represents the bootstrap value (percentage) of the branch, reflecting the confidence of the corresponding branch. The serial number in the brackets is the GenBank accession number, which is used to identify each strain. The strain labeled with black dots is the strain isolated in this study. The scale represents the genetic distance., figureFileSmall=LSaS8iieOEAB04LLwI/5QQ==, figureFileBig=SQ3Jzlv1c5FPVjfgI9+5zA==, tableContent=null), ArticleFig(id=1297571032693493925, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=CN, label=图3, caption=PDCoV CHN/GSYD/2023毒株系统发育树分析, figureFileSmall=LSaS8iieOEAB04LLwI/5QQ==, figureFileBig=SQ3Jzlv1c5FPVjfgI9+5zA==, tableContent=null), ArticleFig(id=1297571032760602790, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=EN, label=Figure 4, caption=Amino acid sequence alignment analysis of S protein of different PDCoV strains., figureFileSmall=vEA2qsvCFmxdd/+/KEO4/Q==, figureFileBig=lKq9Y/Xhrnui01iEbvxCuQ==, tableContent=null), ArticleFig(id=1297571032836100263, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=CN, label=图4, caption=PDCoV不同毒株S蛋白氨基酸序列比对分析, figureFileSmall=vEA2qsvCFmxdd/+/KEO4/Q==, figureFileBig=lKq9Y/Xhrnui01iEbvxCuQ==, tableContent=null), ArticleFig(id=1297571032936763560, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=EN, label=Figure 5, caption=Analysis of amino acid mutation sites of S protein of PDCoV CHN/GSYD/2023 strain. A: Schematic diagram of the PDCoV S protein; B-C: The S protein mutation site of PDCoV CHN/GSYD/2023 strain was displayed in the three-dimensional structure of S protein, and the amino acid mutation residues were displayed in a spherical structure, in which the red spherical structure represented the unique mutation of CHN/GSYD/2023 strain. Important amino acid mutations are shown by local amplification schematics., figureFileSmall=u3SM1pzHVz4G1eJGCze3dg==, figureFileBig=cktUJD8IXbHqMHp5o/o7lg==, tableContent=null), ArticleFig(id=1297571033041621161, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=CN, label=图5, caption=PDCoV CHN/GSYD/2023毒株S蛋白氨基酸突变位点分析, figureFileSmall=u3SM1pzHVz4G1eJGCze3dg==, figureFileBig=cktUJD8IXbHqMHp5o/o7lg==, tableContent=null), ArticleFig(id=1297571033117118634, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=EN, label=Figure 6, caption=The fecal consistency scores and detoxification kinetics curve of piglets. A: The fecal consistency scores of piglets. Clinical signs of diarrhea were scored as normal=0; pasty=1; semiliquid=2; liquid=3; B: The detoxification kinetics curve of piglets. PDCoV copies was calculated according to formula lg X=(41.58-CT value)/3.46., figureFileSmall=2BRJ+PyxF4KGe4YX+A8aDw==, figureFileBig=TfhrbNfWsnHSEZc7lrfdQQ==, tableContent=null), ArticleFig(id=1297571033196810411, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=CN, label=图6, caption=仔猪腹泻评分及排毒动力学曲线, figureFileSmall=2BRJ+PyxF4KGe4YX+A8aDw==, figureFileBig=TfhrbNfWsnHSEZc7lrfdQQ==, tableContent=null), ArticleFig(id=1297571033272307884, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=EN, label=Figure 7, caption=Clinical symptoms, necropsy, histopathological and immunohistochemical analysis of piglets. A-D: Examination of piglets in PDCoV CHN/GSYD/2023 group; E-H: Examination of piglets in PDCoV CHN/XJYN/2016 group; I-L: Examination of piglets in MEM group., figureFileSmall=zheNYyDUKXlwNxJzIqynVg==, figureFileBig=pkiJScE0r26LR6ugMwtyYg==, tableContent=null), ArticleFig(id=1297571033356193965, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=CN, label=图7, caption=仔猪临床症状、剖检、组织病理学和免疫组化分析, figureFileSmall=zheNYyDUKXlwNxJzIqynVg==, figureFileBig=pkiJScE0r26LR6ugMwtyYg==, tableContent=null), ArticleFig(id=1297571033414914222, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=EN, label=Figure 8, caption=Ratio of villus height to crypt depth in ileum of piglets. ns: P>0.05; ***: P<0.001., figureFileSmall=T7oMyWA/NcSKoeZnO7V62Q==, figureFileBig=5US2qVlIuq5yEp/FxTV3LQ==, tableContent=null), ArticleFig(id=1297571033482023087, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=CN, label=图8, caption=仔猪回肠组织的肠绒毛高度和隐窝深度的比值, figureFileSmall=T7oMyWA/NcSKoeZnO7V62Q==, figureFileBig=5US2qVlIuq5yEp/FxTV3LQ==, tableContent=null), ArticleFig(id=1297571033574297776, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=EN, label=Table 1, caption=

Information of PDCoV reference strains

, figureFileSmall=null, figureFileBig=null, tableContent=
Strain namesGenBank accession No.YearOrigin
CHN/AH/2004KP757890.12004China
CHN/AH-81/2018MN520193.12018China
CHN/CH-01/2016KX443143.22016China
CHN/CHGD/2016MH715491.12016China
CHN/GD-1/2018MN520202.12018China
CHN/GS/2016MF642322.12016China
CHN/HB/2014KP757891.12014China
CHN/HKU15-155/2010NC_039208.12010China
CHN/HN/2014KT336560.12014China
CHN/HNZK-04/2018MH708124.12018China
CHN/Hunan/2014KY513724.12014China
CHN/JS/2014KP757892.12014China
CHN/JXJGS01/2016KY293677.12016China
CHN/NH/2015KU981059.12015China
CHN/QH/2017MF642325.12017China
CHN/SD/2014MF431743.12014China
CHN/Tianjin/2016KY065120.12016China
CHN/XJYN/2016MN064712.12016China
JPN/HKD/2016LC260045.12016Japan
JPN/IWT/2014LC260041.12014Japan
KOR/DH1/2016KY354363.12016Korea
KOR/DH2/2016KY354364.12016Korea
LAO/BTL-0115/2016KX118627.12016Laos
THA/S5015L/2015KU051649.12015Thailand
THA/TT-1115/2015KU984334.12015Thailand
USA/Arkansas61/2015KR150443.12015United States of America
USA/HKU15-SD3424/2014KJ584356.12014United States of America
USA/IA-8734/2014KJ567050.12014United States of America
USA/IL/2014KP981395.12014United States of America
HT/Human-0081-4/2014MW685622.12014Haiti
HT/Human-0256-1/2015MW685623.12015Haiti
HT/Human-0329-4/2015MW685624.12015Haiti
USA/Illinois134/2014KJ601778.12014United States of America
USA/Indiana453/2014KR265851.12014United States of America
USA/Iowa136/2015KX022602.12015United States of America
USA/Nebraska210/2014KR265861.12014United States of America
USA/NorthCarolina452/2014KR265858.12014United States of America
USA/OhioCVM1/2014KJ769231.12014United States of America
VNM/HaNoi6/2015KX834351.12015Vietnam
), ArticleFig(id=1297571033658183857, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=CN, label=表1, caption=

PDCoV参考毒株信息

, figureFileSmall=null, figureFileBig=null, tableContent=
Strain namesGenBank accession No.YearOrigin
CHN/AH/2004KP757890.12004China
CHN/AH-81/2018MN520193.12018China
CHN/CH-01/2016KX443143.22016China
CHN/CHGD/2016MH715491.12016China
CHN/GD-1/2018MN520202.12018China
CHN/GS/2016MF642322.12016China
CHN/HB/2014KP757891.12014China
CHN/HKU15-155/2010NC_039208.12010China
CHN/HN/2014KT336560.12014China
CHN/HNZK-04/2018MH708124.12018China
CHN/Hunan/2014KY513724.12014China
CHN/JS/2014KP757892.12014China
CHN/JXJGS01/2016KY293677.12016China
CHN/NH/2015KU981059.12015China
CHN/QH/2017MF642325.12017China
CHN/SD/2014MF431743.12014China
CHN/Tianjin/2016KY065120.12016China
CHN/XJYN/2016MN064712.12016China
JPN/HKD/2016LC260045.12016Japan
JPN/IWT/2014LC260041.12014Japan
KOR/DH1/2016KY354363.12016Korea
KOR/DH2/2016KY354364.12016Korea
LAO/BTL-0115/2016KX118627.12016Laos
THA/S5015L/2015KU051649.12015Thailand
THA/TT-1115/2015KU984334.12015Thailand
USA/Arkansas61/2015KR150443.12015United States of America
USA/HKU15-SD3424/2014KJ584356.12014United States of America
USA/IA-8734/2014KJ567050.12014United States of America
USA/IL/2014KP981395.12014United States of America
HT/Human-0081-4/2014MW685622.12014Haiti
HT/Human-0256-1/2015MW685623.12015Haiti
HT/Human-0329-4/2015MW685624.12015Haiti
USA/Illinois134/2014KJ601778.12014United States of America
USA/Indiana453/2014KR265851.12014United States of America
USA/Iowa136/2015KX022602.12015United States of America
USA/Nebraska210/2014KR265861.12014United States of America
USA/NorthCarolina452/2014KR265858.12014United States of America
USA/OhioCVM1/2014KJ769231.12014United States of America
VNM/HaNoi6/2015KX834351.12015Vietnam
), ArticleFig(id=1297571033725292722, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=EN, label=Table 2, caption=

Clinical symptoms of piglets infected with PDCoV strain

, figureFileSmall=null, figureFileBig=null, tableContent=
GroupNumberPDCoV strainIncidenceCT value
G15PDCoV CHN/GSYD/20235/5 (100)20.37±3.25
G25PDCoV CHN/XJYN/20165/5 (100)21.37±2.78
G35MEM0/5 (0)34.15±1.27
), ArticleFig(id=1297571033834344627, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1297571024070004824, language=CN, label=表2, caption=

仔猪感染PDCoV CHN/GSYD/2023毒株后的临床症状

, figureFileSmall=null, figureFileBig=null, tableContent=
GroupNumberPDCoV strainIncidenceCT value
G15PDCoV CHN/GSYD/20235/5 (100)20.37±3.25
G25PDCoV CHN/XJYN/20165/5 (100)21.37±2.78
G35MEM0/5 (0)34.15±1.27
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猪德尔塔冠状病毒CHN/GSYD/2023株的分离鉴定与致病性分析
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于瑞明 1 , 张莉萍 2 , 周鹏 2 , 张中旺 2 , 潘丽 2 , 郭慧琛 2 , 袁莉刚 1 , 刘新生 2
微生物学报 | 研究报告 2026,66(8): 3994-4007
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微生物学报 |研究报告 2026 , 66 (8) : 3994 -4007
猪德尔塔冠状病毒CHN/GSYD/2023株的分离鉴定与致病性分析
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于瑞明1, 张莉萍2, 周鹏2, 张中旺2, 潘丽2, 郭慧琛2, 袁莉刚1 , 刘新生2
作者信息
  • 1.甘肃农业大学 动物医学院，甘肃 兰州
  • 2.中国农业科学院兰州兽医研究所，动物疫病防控全国重点实验室，甘肃 兰州
通讯作者:
袁莉刚, 刘新生
作者简介:

作者贡献声明

于瑞明：实验设计、实验操作、初稿写作；张莉萍：数据收集和处理；周鹏：数据分析、动物实验操作；张中旺：数据分析及文献查找；潘丽、郭慧琛：监督指导、经费支持；袁莉刚：监督指导、稿件润色修改；刘新生：数据分析、经费支持、稿件润色修改。

Isolation, identification, and pathogenicity analysis of a porcine deltacoronavirus strain CHN/GSYD/2023
Ruiming YU1, Liping ZHANG2, Peng ZHOU2, Zhongwang ZHANG2, Li PAN2, Huichen GUO2, Ligang YUAN1 , Xinsheng LIU2
Affiliations
  • 1.College of Veterinary Medicine, Gansu Agricultural University, Lanzhou, Gansu, China
  • 2.State Key Laboratory of Animal Disease Control and Prevention, Lanzhou Veterinary Research Institute, Chinese Academy for Agricultural Sciences, Lanzhou, Gansu, China
    • Corresponding Author:
    E-mail: YUAN Ligang, ;
    LIU Xinsheng,
出版时间: 2026-08-04 doi: 10.13343/j.cnki.wsxb.20260128
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摘要
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猪德尔塔冠状病毒(porcine deltacoronavirus, PDCoV)是一种重要的猪肠道冠状病毒，给养猪业带来了重大经济损失，截至目前尚无可用的商品化疫苗。 【目的】 获得PDCoV流行毒株，并对其生物学特性和致病性进行初步研究，为尽快研发出有效防控PDCoV的疫苗提供生物材料。 【方法】 使用LLC-PK1细胞分离来自甘肃某猪场PDCoV阳性病料中的病毒，并通过观察细胞病变效应(cytopathic effect, CPE)、间接免疫荧光试验(indirect immunofluorescence assay, IFA)、病毒全基因组测序等方法进行鉴定，同时评价分离毒株在仔猪上的致病性。 【结果】 PDCoV阳性病料接种LLC-PK1细胞后，连续盲传至第6代(passage 6, P6)，细胞出现变大、变圆、皱缩脱落成单个或成簇密集颗粒的典型CPE；IFA鉴定PDCoV阳性，表明成功分离出PDCoV CHN/GSYD/2023毒株。全基因组序列与系统发育分析表明，本研究分离的CHN/GSYD/2023毒株与中国毒株亲缘关系更近，与本实验室保存的CHN/XJYN/2016毒株位于不同的进化分支上；5日龄仔猪感染CHN/GSYD/2023毒株后，仔猪发病率为100% (5/5)。剖检及组织病理学显示，仔猪小肠肠壁变薄，肠腔内含大量积液，回肠绒毛坏死等病理损伤。 【结论】 本研究从甘肃地区某猪场PDCoV阳性样品中成功分离出PDCoV CHN/GSYD/2023毒株。该毒株可在LLC-PK1细胞中稳定增殖传代，对哺乳仔猪具有高致病性。

猪德尔塔冠状病毒  /  分离鉴定  /  致病性

Porcine deltacoronavirus (PDCoV), a major porcine intestinal coronavirus, has caused enormous economic losses to the pig industry. Up to now, there is no commercial vaccine available. [Objective] To obtain the epidemic strain of PDCoV and study its biological characteristics and pathogenicity, thereby providing biological materials for the development of an effective vaccine against PDCoV as soon as possible. [Methods] LLC-PK1 cells were used to isolate the virus from PDCoV-positive samples from a pig farm in Gansu Province. The virus was identified by observation of cytopathic effect (CPE), indirect immunofluorescence assay (IFA), and whole genome sequencing. [Results] After blind passage of the PDCoV-positive LLC-PK1 cells to P6, the cells became larger and rounded, and then shrunk and shed into single or clustered dense particles, typical features of CPE. The IFA results confirmed successful isolation of the PDCoV strain CHN/GSYD/2023. The whole genome sequencing and phylogenetic analysis showed that CHN/GSYD/2023 isolated in this study was more closely related to the strain from China and was located on a different clade from the strain CHN/XJYN/2016 preserved in our laboratory. After 5-day-old piglets were infected with CHN/GSYD/2023, 100% (5/5) of piglets became diseased. Autopsy and histopathology of piglets showed thinning small intestinal wall, a large amount of effusion in the intestinal cavity, ileum intestinal villus necrosis and other pathological damage. [Conclusion] A PDCoV strain CHN/GSYD/2023 was successfully isolated from PDCoV-positive samples in a pig farm in Gansu. The strain can be stably proliferated and passaged in LLC-PK1 cells, and it is highly pathogenic to suckling piglets.

porcine deltacoronavirus  /  isolation and identification  /  pathogenicity
于瑞明, 张莉萍, 周鹏, 张中旺, 潘丽, 郭慧琛, 袁莉刚, 刘新生. 猪德尔塔冠状病毒CHN/GSYD/2023株的分离鉴定与致病性分析. 微生物学报, 2026 , 66 (8) : 3994 -4007 . DOI: 10.13343/j.cnki.wsxb.20260128
Ruiming YU, Liping ZHANG, Peng ZHOU, Zhongwang ZHANG, Li PAN, Huichen GUO, Ligang YUAN, Xinsheng LIU. Isolation, identification, and pathogenicity analysis of a porcine deltacoronavirus strain CHN/GSYD/2023[J]. Acta Microbiologica Sinica, 2026 , 66 (8) : 3994 -4007 . DOI: 10.13343/j.cnki.wsxb.20260128
正文
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冠状病毒(coronavirus, CoV)是一类具有包膜的单股正链RNA病毒，属于尼多病毒目(Nidovirales)，冠状病毒科(Coronaviridae)[1-2]。根据基因组序列，冠状病毒可分为α、β、γ和δ共4个属。猪德尔塔冠状病毒(porcine deltacoronavirus, PDCoV)属于δ冠状病毒属，病毒粒子呈球形，直径约100 nm，有囊膜及末端凸起的刺突，基因组大小约25.4 kb，是目前已知最小的冠状病毒[3]。PDCoV基因组的组成、排列与其他冠状病毒相似，两端包含5′非翻译区(untranslated region, UTR)和3′ UTR，8个开放阅读框(open reading frames, ORFs)分别编码多聚蛋白pp1a/pp1b、纤突蛋白(spike, S)、包膜蛋白(envelope, E)、膜蛋白(membrane, M)、核衣壳蛋白(nucleocapsid, N)、非结构蛋白6 (nonstructural protein 6, NS6)和非结构蛋白7 (nonstructural protein 7, NS7)[4]。PDCoV S蛋白呈同源三聚体结构，以刺突状镶嵌于病毒颗粒表面，包含S1和S2 2个亚基。其中3个S1亚基组成头部，负责识别并结合宿主细胞表面受体；3个S2亚基组成茎部，介导病毒包膜与宿主细胞膜的融合[5-6]。S蛋白具有高度免疫原性，在诱导机体产生中和抗体方面最为重要，因此也是疫苗研发的重要靶点[7]。
PDCoV是引起仔猪病毒性腹泻的重要病原之一[8]。自2012年首次报道以来[9]，PDCoV已在全球范围内广泛传播，严重危害多国生猪养殖产业[10-11]。PDCoV主要感染哺乳仔猪，其引起的仔猪呕吐、腹泻和脱水等临床症状与病理变化与传染性胃肠炎病毒(transmissible gastroenteritis virus, TGEV)、猪流行性腹泻病毒(porcine epidemic diarrhea virus, PEDV)、猪急性腹泻综合征冠状病毒(swine acute diarrhea syndrome coronavirus, SADS-CoV)及猪轮状病毒(porcine rotavirus, PoRV)所致者极为相似；且PDCoV与PEDV或TGEV等病原混合感染可导致更为严重的临床症状和更高的致死率[11-12]。研究表明，PDCoV在实验条件下可感染鸡、火鸡、牛等动物，具有跨物种传播潜力[13-14]。然而，最新研究表明PDCoV可感染儿童，研究人员从3名海地急性未分化发热儿童血浆样本中分离到PDCoV，基因组和进化分析表明，人类感染至少源于2个不同病毒谱系的跨种传播事件，这些病毒在编码Nsp15和S蛋白的基因中具有相同的突变特征；结构分析进一步预测，S1亚基受体结合域中的S1亚基的1个氨基酸变化可能影响蛋白质的柔韧性及其与宿主细胞受体的结合[15]。因此，PDCoV与其他猪肠道病毒的共感染特性及其跨物种传播能力使其成为不容忽视的经济和公共卫生风险。
PDCoV在适应环境变化过程中不断进化，频繁变异导致其跨物种传播能力持续增强。因此，分离和鉴定新的流行毒株尤为迫切。本研究针对当前PDCoV流行毒株，通过病毒分离、鉴定及致病性分析阐明流行毒株的遗传变异规律，旨在为PDCoV流行病学监测及疫苗研发提供数据支撑。
1 材料与方法
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1.1 病料
病料样本采自甘肃永登县某猪场腹泻猪的粪便及肠道内容物，于-80 ℃以下超低温冰箱中保存备用。
1.2 细胞、病毒和实验动物
猪肾细胞LLC-PK1细胞购自ATCC (CL-101TM)，由中国农业科学院兰州兽医研究所传代并保存。PDCoV CHN/XJYN/2016毒株由本实验室前期分离并保存[16]。PDCoV抗原、抗体均为阴性的5日龄仔猪购自甘肃省某猪场。所有实验猪在兰州兽医研究所ABSL-Ⅱ实验室内饲养，所有实验猪均接受人文关怀，并于实验结束时实施安乐死。本研究所有动物实验均获中国农业科学院兰州兽医研究所动物伦理委员会批准，编号为LVRIAEC-2025-134。
1.3 主要试剂和仪器
EastepTM Super Total RNA Extraction Kit，Promega公司；One Step PrimeScriptTM RT-PCR Kit (Perfect Real Time)，宝生物工程(大连)有限公司；MEM培养基、青链霉素双抗、PBS、胎牛血清、Trypsin (2.5%)、HEPES、NEAA，Gibco公司；牛血清白蛋白(bovine serum albumin, BSA)，Invitrogen公司；HiPure Viral RNA Kit，Magen Biotechnology公司；PDCoV抗S蛋白单克隆抗体由本实验室制备并保存[17]。
低温高速离心机，Eppendorf公司；CFX96TM Real-Time System，Bio-Rad公司；倒置生物显微镜，Nikon公司；FormaTM Steri-CycleTM CO2孵育器和大屏幕数码倒置显微镜，ThermoFisher Scientific公司；生物安全柜，苏州安泰公司。
1.4 病料处理
将从甘肃永登县某猪场采集的临床腹泻样品用含1%青链霉素双抗的MEM培养基10倍稀释后充分振荡，4 ℃、3 500 r/min离心30 min，吸取上清液经0.22 μm无菌滤膜过滤，分装后于-80 ℃冰箱保存备用。
1.5 RT-qPCR
按照EastepTM Super Total RNA Extraction Kit操作说明提取病毒RNA，利用TaqMan实时荧光定量One Step PrimScriptTM RT-PCR Kit检测病料中PDCoV病毒滴度。所用PDCoV N基因特异性引物为PDCoV-N-Forward (5′-ACGTCG TAAGACCCAGCATC-3′)和PDCoV-N-Reverse (5′-CCCACCTGAAAGTTGCTCTC-3′)，探针为PDCoV-N-probe (5′-CY5-GTATGGCTGATCCTC GCATCATGGC-BHQ2-3′)。RT-qPCR反应体系(20 μL)：2×One Step RT-PCR Buffer Ⅲ 10 μL，TaKaRa Ex Taq HS (5 U/μL) 0.4 μL，PrimeScript RT Enzyme Mix Ⅱ 0.4 μL，上、下游引物(10 µmol/L)各0.4 μL，PDCoV-N-probe 0.6 μL，Nuclease-Free Water 5.8 μL，RNA 2 μL。反应条件：42 ℃ 5 min，95 ℃ 10s；95 ℃ 5 s，57 ℃ 20 s，共40次循环。
1.6 病毒的分离和传代培养
将LLC-PK1细胞接种于T25细胞瓶中，置于37 ℃、5% CO2培养箱中培养至细胞密度达70%-80%时用无菌PBS洗涤3次备用。将处理好的病料上清液用2 mL含1% NEAA、1% HEPES和20 µg/mL Trypsin的MEM稀释，混匀后加入T25培养瓶中，置于37 ℃、5% CO2培养箱中孵育1 h，加入2 mL MEM (含20 µg/mL胰酶)继续培养，同时设置空白细胞作为阴性对照。每天观察细胞病变情况。在前几代，病毒感染细胞后CPE较少甚至不出现，此种情况下需于接种后72 h收集病毒，于-80 ℃中反复冻融3次后继续盲传，直至出现明显细胞病变。待细胞病变达90%时收获病毒。
1.7 间接免疫荧光
以病毒生长培养基作为阴性对照，将LLC-PK1细胞接种于6孔板中培养至单层，弃去培养液，用PBS洗涤细胞3次。以PDCoV CHN/GSYD/2023毒株感染LLC-PK1细胞，感染复数(multiplicity of infection, MOI)为0.01。接种24 h后，弃去培养基并用PBS洗涤细胞3次。用4%多聚甲醛固定细胞30 min，加入1 mL 0.25% Triton X-100渗透，室温静置10 min。PBS洗涤3次后，用5% BSA室温封闭1 h。每孔加入1:1 000稀释的小鼠抗PDCoV S蛋白单克隆抗体(本实验室制备和保存)，37 ℃孵育1 h。随后加入稀释的Alexa Fluor 488标记的山羊抗小鼠IgG抗体，37 ℃孵育1 h。最后用4′,6-二脒基-2-苯基吲哚(4′,6-diamidino-2-phenylindole, DAPI)染色5 min，PBS洗涤3次，于荧光显微镜下观察并拍照。
1.8 半数组织培养物感染量测定
将冻融收集的病毒液室温融化，3 500 r/min离心2 min，取上清，用MEM培养基(胰酶浓度20 µg/mL)将病毒进行连续10倍稀释，即10-1，10-2，…(根据病毒大致滴度确定稀释倍数)；然后用排枪将细胞接种至96孔板中，细胞汇合度控制在80%左右，PBS洗涤2次，再以无血清MEM洗涤1次。每个稀释度取100 μL加入96孔板，每个稀释度设8个重复孔，再补加100 μL MEM，终体积为200 μL。设置空白对照，即加入200 μL MEM。置于37 ℃培养，每天观察细胞病变并记录病变孔数，以Reed-Muench法计算半数组织培养物感染量(50% tissue culture infectious dose, TCID50)。
1.9 病毒复制动力学
将MOI为0.01的病毒液接种至长满单层的LLC-PK1细胞T25培养瓶中，体系中胰酶终浓度为20 μg/mL。分别于感染后0、6、12、24、36、48、60、72 h (hours post-infection, hpi)收获病毒培养物，测定各时间点TCID50，绘制病毒生长曲线。
1.10 病毒全基因组高通量测序
高通量测序(又称二代测序、深度测序)技术，通过大规模平行测序方式可快速无差别地测定样本中所有核酸。根据HiPure Viral RNA Kit操作说明提取PDCoV CHN/GSYD/2023第6代(passage 6, P6)细胞培养物中的病毒RNA。核酸经文库构建、上机测序后转化为生物信息学软件可识别的数据格式，再经软件流程质控、去除rRNA、宿主序列及细菌等冗余序列后获得初步病毒数据，最终经组装、比对、注释等分析流程获得病毒基因组信息。
1.11 系统发育树分析
将PDCoV CHN/GSYD/2023毒株与GenBank中39株PDCoV完整基因组序列(表1)进行系统发育分析。系统发育树采用MEGA 7.0基于距离的邻接法构建，以1 000个重复数据集进行了bootstrap分析。
1.12 仔猪致病性实验
15头5日龄健康新生仔猪购自无PDCoV感染或疫苗接种史的商品化猪场。接种前采集仔猪血清检测PDCoV特异性抗体，同时采集直肠拭子进行RT-qPCR检测，确认PDCoV、PEDV、TGEV和PoRV等主要猪肠道病毒均为阴性。所有仔猪随机分为3组，每组5头，分笼分圈饲养。G1组每头仔猪口服1 mL CHN/GSYD/2023-P6病毒液(104.91 TCID50/mL)，G2组每头仔猪口服1 mL CHN/XJYN/2016-P6病毒液(104.80 TCID50/mL)，G3组每头仔猪口服1 mL MEM作为对照。实验期间，每天监测并记录仔猪临床体征(腹泻、呕吐、厌食等)，并对粪便稠度评分：固体为0分，糊状为1分，半流质(轻度腹泻)为2分，液体(重度腹泻)为3分。实验结束后采集肠道组织进行组织病理学检查。
1.13 组织病理学和免疫组化分析
感染7 d后对试验仔猪实施安乐死并解剖，取回肠组织以4%多聚甲醛常温固定48 h，经脱水、石蜡包埋、切片、HE染色、封片后，于显微摄像系统下采集图像。同时，采用PDCoV S蛋白特异性单克隆抗体(本实验室保存)对肠道组织进行免疫组化(immunohistochemistry, IHC)分析。
2 结果与分析
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2.1 PDCoV CHN/GSYD/2023毒株的分离鉴定
将RT-qPCR检测PDCoV阳性的粪便样品处理后接种于LLC-PK1细胞，进行连续盲传。结果显示，传至第5代(passage 5, P5)时未观察到明显细胞病变效应(cytopathic effect, CPE)；传至P6时，显微镜下可见细胞变大、变圆，出现单个或成簇存在的密集颗粒细胞及细胞皱缩、脱落等典型CPE；继续传至P7，仍可观察到明显CPE (图1A-1C)。将PDCoV CHN/GSYD/2023毒株P5、P6和P7以MOI=0.01接种至铺满LLC-PK1细胞的35 mm培养皿中，以小鼠抗PDCoV S蛋白特异性单克隆抗体进行间接免疫荧光试验(indirect immunofluorescence assay, IFA)鉴定。结果表明，24 hpi后，P5、P6和P7感染的LLC-PK1细胞中均可观察到绿色荧光，表明CHN/GSYD/2023毒株P5、P6和P7可在LLC-PK1细胞中稳定复制(图1D-1F)。
2.2 PDCoV CHN/GSYD/2023毒株的增殖动力学
将PDCoV CHN/GSYD/2023或CHN/XJYN/2016毒株P6接种于LLC-PK1细胞，于0、6、12、24、36、48、60、72 hpi收取病毒液，测定各时间点TCID50，比较分析2株病毒的增殖动力学。结果显示，PDCoV CHN/GSYD/2023毒株感染LLC-PK1细胞后可持续增殖，60 hpi时病毒TCID50达峰值(104.91 TCID50/mL)，随后略有下降，与CHN/XJYN/2016毒株相比差异不显著(图2)。
2.3 PDCoV CHN/GSYD/2023毒株全基因组测序及系统进化分析
对PDCoV CHN/GSYD/2023毒株进行全基因组测序分析(GenBank登录号为PZ065654)，结果显示该毒株基因组全长为25 394 nt (不含polyA尾)。系统进化分析表明，所有PDCoV毒株遗传进化上亲缘关系密切，序列相似性高，提示其可能起源于共同祖先。美国、韩国和日本的PDCoV分离株聚于同一分支，亲缘关系较近，可能源于同一毒株；越南、老挝和泰国的分离株归于另一分支，且东南亚分离株与中国毒株亲缘关系较近。本研究分离的CHN/GSYD/2023毒株与本实验室保存的CHN/XJYN/2016毒株位于不同进化分支(图3)。
选有代表的毒株对其S蛋白氨基酸序列进行比对分析，结果发现S蛋白变异位点主要集中在NTD、CTD及S1和S2亚基裂解位点附近(图4)。根据突变位点在S蛋白三维结构中的位置分析(图5)，这些突变位点多位于S蛋白结构表面，而表面氨基酸对抗体识别至关重要，其频繁突变可能有助于病毒免疫逃逸。本研究分离的CHN/GSYD/2023毒株S蛋白存在11个独有突变位点(图5A、5B)，主要集中在NTD、CTD和膜融合肽附近，这些位点可能对该毒株的致病性具有重要影响。
2.4 PDCoV CHN/GSYD/2023毒株对哺乳仔猪具有高致病性
为确定PDCoV CHN/GSYD/2023毒株对仔猪的致病性，以P6对5日龄哺乳仔猪进行攻毒试验，并以本实验室保存的CHN/XJYN/2016毒株作为对照。结果显示，G1组和G2组所有仔猪均出现水样腹泻、呕吐、精神抑郁等临床症状，直肠拭子样本检测PDCoV RNA阳性；G3对照组所有仔猪健康状态良好，无腹泻、呕吐等临床症状，直肠拭子样本检测PDCoV RNA阴性(表2，图6)。剖检大体病理学检查发现，G1组和G2组仔猪小肠肠壁变薄，肠腔内含大量积液和气体(图7B、7F)；G3对照组仔猪肠道组织正常，未见病理变化(图7J)。免疫组化分析结果显示，G1组和G2组仔猪回肠肠上皮细胞上可见大量PDCoV抗原(图7C、7G)；G3组仔猪回肠肠上皮细胞未观察到PDCoV抗原(图7K)。组织病理学结果显示，G1组和G2组仔猪回肠可见肠绒毛坏死、固有层细胞坏死、黏膜下水肿、淋巴细胞和中性粒细胞浸润等病理损伤(图7D、7H)；而G3组仔猪回肠组织未观察到病理损伤(图7L)。G1组和G2组仔猪回肠组织肠绒毛高度与隐窝深度的比值均显著低于G3组(图8)。
3 讨论与结论
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PDCoV作为一种新发猪肠道冠状病毒，已给全球生猪养殖业带来巨大经济损失。对于病毒性疾病的防控，疫苗接种仍是目前最有效的手段，但目前尚无针对PDCoV的有效商品化疫苗[18]。为尽快研发能有效防控PDCoV的疫苗，本研究从腹泻仔猪病料中分离出1株具有高致病性的PDCoV毒株CHN/GSYD/2023。
PDCoV的体外分离中，细胞系的选择至关重要。已有研究表明，PDCoV可在LLC-PK1细胞(ATCC编号为CL-101)或ST细胞(ATCC编号为CRL1746)中稳定增殖和传代[19-21]。因此，根据本实验室已有的细胞系资源，本研究选择LLC-PK1细胞对PDCoV流行毒株进行分离鉴定和连续传代培养。将处理后的阳性病料接种LLC-PK1细胞后，连续盲传至P6，细胞出现变大、变圆，继而皱缩脱落成单个或成簇存在的密集颗粒等典型CPE (图1B)，这与Gao等[16]的研究结果一致。随后，通过IFA检测P5、P6和P7病毒S蛋白在LLC-PK1细胞中的表达，可见特异性荧光信号(图1D-1F)，表明成功分离到PDCoV CHN/GSYD/2023毒株。TCID50测定结果(图2)表明，PDCoV CHN/GSYD/2023毒株可在LLC-PK1细胞中稳定增殖，60 hpi时病毒TCID50达峰值(104.91 TCID50/mL)，随后有所下降，与CHN/XJYN/2016毒株相比差异不显著。
截至目前，PDCoV已在中国、美国、日本、韩国和东南亚地区分化出3个遗传分支[22-23]。自2014年以来，中国谱系一直是PDCoV进化传播的主要分支；同时，作为现有疫苗研发的参考谱系，紧跟其流行趋势和演变规律极为重要[24-25]。本研究对分离的PDCoV CHN/GSYD/2023毒株进行了全基因组测序，并与GenBank中39株PDCoV全基因组序列进行了比对和系统发育分析。结果表明，CHN/GSYD/2023毒株与中国毒株亲缘关系更近，与本实验室保存的CHN/XJYN/2016毒株位于不同的进化分支上，而CHN/XJYN/2016毒株与越南毒株亲缘关系更为接近(图3)。S蛋白氨基酸序列比对和突变位点分析表明，CHN/GSYD/2023毒株S蛋白存在11个独有突变位点(图5A、5B)，主要集中在NTD、CTD和膜融合肽附近，这些位点的突变是否影响其致病性，有待后续试验进一步验证。
已有研究表明，PDCoV毒株在临床致病性上存在差异，某些高致病毒株可引起新生仔猪急性水样腹泻、脱水甚至死亡，从而对养猪业造成重大经济损失[26]。本研究中，PDCoV CHN/GSYD/2023毒株和CHN/XJYN/2016毒株均可致5日龄哺乳仔猪100% (5/5)发病。剖检和组织病理学结果显示，仔猪小肠肠壁变薄，肠腔内含大量积液，回肠肠绒毛坏死，固有层细胞坏死，黏膜下水肿，淋巴细胞和中性粒细胞浸润等病理损伤。对照组仔猪健康状态良好，剖检未见肠道组织损伤。然而，本研究样本量较小(n=5)，具有一定局限性，后续研究将考虑增大样本量(如15头/组或设置多个重复组)，以获得更为稳健的数据。
综上所述，本研究从甘肃地区某猪场PDCoV阳性样品中成功分离出PDCoV CHN/GSYD/2023毒株。该毒株可在LLC-PK1细胞中稳定增殖传代，全基因组系统发育分析表明其与中国毒株亲缘关系更近，与本实验室保存的CHN/XJYN/2016毒株位于不同的进化分支上。该毒株感染5日龄哺乳仔猪可致100% (5/5)发病，并引起严重的肠道组织病理损伤。
基金
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  • 国家生猪技术创新中心项目(NCTIP-XD/C 03)
  • 甘肃省科技重大专项(24ZD13NA008)
  • 甘肃省科技重大专项(23ZDNA007)
文献
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2026年第66卷第8期
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doi: 10.13343/j.cnki.wsxb.20260128
  • 接收时间：2026-02-10
  • 首发时间：2026-08-21
  • 出版时间：2026-08-04
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  • 收稿日期：2026-02-10
  • 录用日期：2026-03-25
基金
National Center of Technology Innovation for Pigs Project(NCTIP-XD/C 03)
国家生猪技术创新中心项目(NCTIP-XD/C 03)
Major Science and Technology Project of Gansu Province(24ZD13NA008)
甘肃省科技重大专项(24ZD13NA008)
Major Science and Technology Project of Gansu Province(23ZDNA007)
甘肃省科技重大专项(23ZDNA007)
作者信息
    1.甘肃农业大学 动物医学院，甘肃 兰州
    2.中国农业科学院兰州兽医研究所，动物疫病防控全国重点实验室，甘肃 兰州

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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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