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Article(id=1192149556004532367, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1192149543010582589, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250205, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1742140800000, receivedDateStr=2025-03-17, revisedDate=null, revisedDateStr=null, acceptedDate=1746460800000, acceptedDateStr=2025-05-06, onlineDate=1762160203219, onlineDateStr=2025-11-03, pubDate=1756915200000, pubDateStr=2025-09-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1762160203219, onlineIssueDateStr=2025-11-03, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1762160203219, creator=13701087609, updateTime=1762160203219, updator=13701087609, issue=Issue{id=1192149543010582589, tenantId=1146029695717560320, journalId=1192105938417971205, year='2025', volume='65', issue='10', pageStart='4241', pageEnd='4713', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1762160200113, creator=13701087609, updateTime=1762160638682, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1192151382586175735, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1192149543010582589, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1192151382586175736, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1192149543010582589, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=4517, endPage=4527, ext={EN=ArticleExt(id=1192149556994388122, articleId=1192149556004532367, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Pathogenicity and fungicide resistance of Lichtheimiaramosa LYSF001: analysis of virulence genes and evaluation of sensitivity to fungicides, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

[Objective] To investigate the pathogenicity of Lichtheimiaramosa LYSF001 and ITS resistance to common fungicides, providing experimental evidence for the diagnosis and treatment of fungal infections in animals. [Methods] The strain was identified by morphological characterization and molecular biology techniques. PCR was employed to detect virulence genes in strain LYSF001, and an animal infection model was established to evaluate the pathogenicity of the strain. Histopathological examination, Grocott’s methenamine silver (GMS) staining, and ITS sequences amplification were employed to analyze pathological changes in the livers and kidneys of infected animals. Additionally, antimicrobial susceptibility tests were conducted to assess the sensitivity of strain LYSF001 to caspofungin, amphotericin B, and itraconazole. [Results] Strain LYSF001 was identified as L. ramosa. PCR analysis revealed that strain LYSF001 carried five virulence genes (CalA, PKP2, LaeA, Alp2, and AspF1). Animal experiments demonstrated that the strain led to the mortality of 40% and caused visceral hyperemia in mice, with the most significant pathological changes (inflammatory cell infiltration and tissue necrosis) observed in the livers and kidneys. Severe infections led to animal mortality. GMS staining revealed the presence of dark-colored hyphae in the heart and liver, and ITS sequences amplification further confirmed fungal infection. The antimicrobial susceptibility test results indicated that strain LYSF001 was resistant to caspofungin but sensitive to amphotericin B and itraconazole. [Conclusion] L. ramosa LYSF001 exhibits strong pathogenicity, capable of causing severe infections and even death in animals. Additionally, the strain showed resistance to caspofungin but sensitivity to amphotericin B and itraconazole. The findings provide important experimental evidence and technical support for the diagnosis and treatment of fungal infections in animals.

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E-mail: WANG Li,
WEI Yong,
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These authors contributed equally to this work.

, authorsList=Keyu ZOU, Yanwen YANG, Liu XING, Jingyun CHEN, Li WANG, Yong WEI), CN=ArticleExt(id=1192149822481248834, articleId=1192149556004532367, tenantId=1146029695717560320, journalId=1192105938417971205, language=CN, title=分枝横梗霉LYSF001的毒力基因分析与抗真菌药物敏感性评估, columnId=1192149544164012138, journalTitle=微生物学报, columnName=研究报告, runingTitle=null, highlight=null, articleAbstract=

【目的】 探讨分枝横梗霉菌株LYSF001的致病性及其对常见抗真菌药物的耐药性，为动物真菌感染的诊断与治疗提供实验依据。 【方法】 通过形态学特征和分子生物学技术分离并鉴定菌株；利用PCR技术检测菌株LYSF001的毒力基因，并构建动物感染模型以评估其致病性；采用组织病理学检查、GMS染色及ITS序列扩增技术分析感染动物的肝脏和肾脏病理变化；通过药敏试验评估菌株LYSF001对卡泊芬净、两性霉素B和伊曲康唑的敏感性。 【结果】 菌株LYSF001鉴定为分枝横梗霉；检测出菌株LYSF001携带5个毒力基因，分别为CalA、PKP2、LaeA、Alp2和AspF1；该菌株感染小鼠后造成40%的死亡率，试验小鼠内脏充血，其中肝脏和肾脏的病理变化最为显著，表现为炎症细胞浸润和组织坏死；GMS染色显示心脏、肝脏中存在深色菌丝，ITS序列扩增进一步证实了真菌感染；药敏试验证实菌株LYSF001对两性霉素B及伊曲康唑表现出敏感性，然而该菌株对卡泊芬净存在耐药性。 【结论】 本研究证实分枝横梗霉菌株LYSF001具有较强的致病性，可引发动物严重感染甚至死亡；同时，该菌株对两性霉素B和伊曲康唑敏感，但对卡泊芬净耐药。这为动物真菌感染的诊断与治疗提供了重要的实验依据和技术支持。

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作者贡献声明

邹珂宇：论文撰写，数据收集与监管，统计分析等；杨彦文：审阅文章、数据收集与监管，数据分析；邢柳：软件程序、提出概念；陈景运：软件程序、提出概念；王利：项目管理、监督管理；魏勇：获取基金、项目管理。

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Clinical and Vaccine Immunology, 2009, 16(10): 1485-1492., articleTitle=Immune responses of human immature dendritic cells can be modulated by the recombinant Aspergillus fumigatus antigen Aspf1, refAbstract=null), Reference(id=1192160978214208082, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, doi=null, pmid=null, pmcid=null, year=2018, volume=9, issue=1, pageStart=1062, pageEnd=1073, url=null, language=null, rfNumber=[25], rfOrder=26, authorNames=LIU H, XU WJ, SOLIS NV, WOOLFORD C, MITCHELL AP, FILLER SG, journalName=Virulence, refType=null, unstructuredReference=LIU H, XU WJ, SOLIS NV, WOOLFORD C, MITCHELL AP, FILLER SG. Functional convergence of gliP and aspf1 in Aspergillus fumigatus pathogenicity[J]. 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Mycology, 2024, 15(1): 120-128., articleTitle=Estimating poultry aspergillosis prevalence and diagnostic accuracy of histopathological and mycological culture in Côte d’Ivoire using Bayesian latent class analysis, refAbstract=null), Reference(id=1192160978381980246, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, doi=null, pmid=null, pmcid=null, year=2009, volume=7, issue=null, pageStart=138, pageEnd=null, url=null, language=null, rfNumber=[27], rfOrder=28, authorNames=PIANCASTELLI C, GHIDINI F, DONOFRIO G, JOTTINI S, TADDEI S, CAVIRANI S, CABASSI CS, journalName=Reproductive Biology and Endocrinology, refType=null, unstructuredReference=PIANCASTELLI C, GHIDINI F, DONOFRIO G, JOTTINI S, TADDEI S, CAVIRANI S, CABASSI CS. Isolation and characterization of a strain of Lichtheimia corymbifera (ex Absidia corymbifera) from a case of bovine abortion[J]. Reproductive Biology and Endocrinology, 2009, 7: 138., articleTitle=Isolation and characterization of a strain of Lichtheimia corymbifera (ex Absidia corymbifera) from a case of bovine abortion, refAbstract=null)], funds=[Fund(id=1192160974862959120, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, awardId=sccxtd-2024-14, language=EN, fundingSource=the Sichuan Meat Sheep Innovation Team—Epidemic Prevention Position(sccxtd-2024-14), fundOrder=null, country=null), Fund(id=1192160974951039506, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, awardId=sccxtd-2024-14, language=CN, fundingSource=四川省肉羊创新团队——防疫岗位(sccxtd-2024-14), fundOrder=null, country=null), Fund(id=1192160975013954068, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, awardId=ZYN2025149, language=EN, fundingSource=the Southwest Minzu University for the Fundamental Research Funds for the Central Universities(ZYN2025149), fundOrder=null, country=null), Fund(id=1192160975068480022, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, awardId=ZYN2025149, language=CN, fundingSource=西南民族大学中央高校基本科研业务费专项资金(ZYN2025149), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1192160970567991729, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, xref=1, ext=[AuthorCompanyExt(id=1192160970576380337, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, companyId=1192160970567991729, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1Key Laboratory of Qinghai-Xizang Plateau Animal Genetic Resource Reservation and Exploitation of Ministry of Education, Southwest Minzu University, Chengdu, Sichuan, China), AuthorCompanyExt(id=1192160970580574642, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, companyId=1192160970567991729, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1西南民族大学，青藏高原动物遗传资源保护与利用教育部重点实验室，四川 成都)]), AuthorCompany(id=1192160970639294900, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, xref=2, ext=[AuthorCompanyExt(id=1192160970643489204, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, companyId=1192160970639294900, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2Sichuan Animal Science Academy, Chengdu, Sichuan, China), AuthorCompanyExt(id=1192160970651877813, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, companyId=1192160970639294900, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2四川省畜牧科学研究院，四川 成都)])], figs=[ArticleFig(id=1192160974028292605, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, language=EN, label=Figure 1, caption=Characterization of the strain LYSF001. A: The culture of strain LYSF001 at 37 ℃ for seven days; B: Observation of strain LYSF001 morphology under upright microscope (1: 400×; 2: 1 000×); C: Observation of strain LYSF001 morphology under SEM (1: 3 000×; 2: 40 000×); D: The growth curves of strain LYSF001 at 37 ℃、OD600 (1, 2, 3: Three repetitions of strain LYSF001); E: The growth condition and growth curves of strain LYSF001 at 18, 28, and 38 ℃, respectively., figureFileSmall=tXD67bSUj4H4KVxc1zgkrw==, figureFileBig=LP7AKnR4/kytYFde5lDlsg==, tableContent=null), ArticleFig(id=1192160974095401471, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, language=CN, label=图1, caption=菌株LYSF001的特征。A：菌株LYSF001在37 ℃下培养7 d的菌落形态；B：正置显微镜下观察菌株LYSF001的形态(1：200×；2：1 000×)；C：扫描电子显微镜(SEM)下观察菌株LYSF001的形态(1：3 000×；2：40 000×)；D：菌株LYSF001在37 ℃、OD600下的生长曲线(1、2、3：菌株LYSF001的3个重复)；E：菌株LYSF001在18、28和38 ℃下的生长情况及生长曲线。, figureFileSmall=tXD67bSUj4H4KVxc1zgkrw==, figureFileBig=LP7AKnR4/kytYFde5lDlsg==, tableContent=null), ArticleFig(id=1192160974170898945, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, language=EN, label=Figure 2, caption=Identification of the strain LYSF001. A: Identification of ITS sequences of isolated fungi (Lane M: DL2000 DNA marker; Lane 1: PCR product of strain LYSF001); B: Phylogenetic tree analysis of strain LYSF001 (The numbers in parentheses represent the ITS sequence registry numbers of each species in the database, and the values at the branch nodes reflect the statistical support of the corresponding branches; The scale bar indicates the sequence differences displayed on the phylogenetic tree, with a scale of 0.50); C: Virulence gene identification of strain LYSF001 (Lane M: DL2000 DNA marker; Lane 1: CotH; Lane 2: PrtT; Lane 3: CalA; Lane 4: PKP2; Lane 5: LaeA; Lane 6: Alp2; Lane 7: AspF1; -: No specific target band; +: Specific target band)., figureFileSmall=3NZlUbEoVFU7emxzvakv0g==, figureFileBig=pk0asEhMZiN83F5VFZTmPw==, tableContent=null), ArticleFig(id=1192160974233813507, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, language=CN, label=图2, caption=菌株LYSF001基因鉴定。A：分离真菌ITS序列的鉴定结果(泳道M：DL2000 DNA marker；泳道1：菌株LYSF001的PCR产物)；B：菌株LYSF001的系统发育树分析(图中圆括号内标注的数字代表各物种ITS序列在数据库中的注册编号，分支节点处数值则反映对应分支的统计支持率；标尺表示发育树可显示的序列差异度，比例尺为0.50)；C：菌株LYSF001的毒力基因鉴定(泳道M：DL2000 DNA marker；泳道1：CotH；泳道2：PrtT；泳道3：CalA；泳道4：PKP2；泳道5：LaeA；泳道6：Alp2；泳道7：AspF1；-：无特异性目的条带；+：有特异性目的条带)。, figureFileSmall=3NZlUbEoVFU7emxzvakv0g==, figureFileBig=pk0asEhMZiN83F5VFZTmPw==, tableContent=null), ArticleFig(id=1192160974288339460, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, language=EN, label=Figure 3, caption=Histopathological examination. A: Results of ITS sequence detection in different organs of mice (Lane M: DL000 DNA marker; Lane 1: Positive control; Lane 2: Heart; Lane 3: Liver; Lane 4: Spleen; Lane 5: Lungs; Lane 6: Kidneys; Lane 7: Small intestine); B: Pathological tissue sections of mice (400×); C: Transmission electron microscope observation of mouse tissues (1, 3, 5, 7: 6 000×; 2, 4, 6, 8: 12 000×)., figureFileSmall=swzZjYdwyWRq+dML8Ke72Q==, figureFileBig=ov8WVsnXmCxqQDhZor9rOg==, tableContent=null), ArticleFig(id=1192160974393197062, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, language=CN, label=图3, caption=组织病理学检查。A：小鼠不同器官中ITS序列检测结果(泳道M：DL2000 DNA marker；泳道1：阳性对照；泳道2：心脏；泳道3：肝脏；泳道4：脾脏；泳道5：肺；泳道6：肾脏；泳道7：小肠)；B：小鼠病理组织切片(400×)；C：小鼠组织的透射电子显微镜观察(1、3、5、7：6 000×；2、4、6、8：12 000×)。, figureFileSmall=swzZjYdwyWRq+dML8Ke72Q==, figureFileBig=ov8WVsnXmCxqQDhZor9rOg==, tableContent=null), ArticleFig(id=1192160974460305928, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, language=EN, label=Table 1, caption=

Primers for virulence genes of strain LYSF001

, figureFileSmall=null, figureFileBig=null, tableContent=

基因

Gene

引物序列

Primer sequences (5′→3′)

序列大小

Fragment size (bp)

退火温度

Annealing temperature (℃)

CotHF: GCGGATTTCACCGCTTCTTC1 30955.0
R: CGCTTTTTCCGCGAATACCC
PKR2F: CTGAACTCCACGTGGGCATA58353.0
R: GACCAATCACTGAGCCGTCA
CalAF: ATTGAACGTCACACCCCGTT62855.0
R: ATCGAAGGCGAGCGATTACA
PrtTF: AGGCATACCACCACCAAT17554.0
R: GCTCAGTAACCAGCCATCT
LaeAF: CGCCTACACTGGCTCGTT37457.0
R: AGGGTTGGATGGGACTGA
Alp2F: CAACGGACACGGAACTCA13051.0
R: CAGAACGACATCGGACATAG
AspF1F: ACTGGTTCACTAACGGCTACG20151.0
R: GGTCTTCCTTGGGTTTCTTC
), ArticleFig(id=1192160974531609098, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, language=CN, label=表1, caption=

菌株LYSF001毒力基因引物

, figureFileSmall=null, figureFileBig=null, tableContent=

基因

Gene

引物序列

Primer sequences (5′→3′)

序列大小

Fragment size (bp)

退火温度

Annealing temperature (℃)

CotHF: GCGGATTTCACCGCTTCTTC1 30955.0
R: CGCTTTTTCCGCGAATACCC
PKR2F: CTGAACTCCACGTGGGCATA58353.0
R: GACCAATCACTGAGCCGTCA
CalAF: ATTGAACGTCACACCCCGTT62855.0
R: ATCGAAGGCGAGCGATTACA
PrtTF: AGGCATACCACCACCAAT17554.0
R: GCTCAGTAACCAGCCATCT
LaeAF: CGCCTACACTGGCTCGTT37457.0
R: AGGGTTGGATGGGACTGA
Alp2F: CAACGGACACGGAACTCA13051.0
R: CAGAACGACATCGGACATAG
AspF1F: ACTGGTTCACTAACGGCTACG20151.0
R: GGTCTTCCTTGGGTTTCTTC
), ArticleFig(id=1192160974644855307, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, language=EN, label=Table 2, caption=

Results of drug sensitivity test

, figureFileSmall=null, figureFileBig=null, tableContent=

抗真菌药物

Antifungal drugs

浓度

Content (µg/tablet)

敏感

Sensitive (S) (mm)

中度敏感

Intermediary (I) (mm)

耐药

Drug resistant (R) (mm)

抑菌圈直径

Inhibition zone diameter (mm)

结果

Results

两性霉素B

Amphotericin B

10≥1513-14<1230.00±1.00S

伊曲康唑

Itraconazole

8≥1714-16<1324.66±0.15S

卡泊芬净

Carbenicillin

5≥1714-16<138.00±0.10R

伏立康唑

Voriconazole

1≥1714-16<1313.33±0.20I
), ArticleFig(id=1192160974720352781, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1192149556004532367, language=CN, label=表2, caption=

药物敏感性测试结果

, figureFileSmall=null, figureFileBig=null, tableContent=

抗真菌药物

Antifungal drugs

浓度

Content (µg/tablet)

敏感

Sensitive (S) (mm)

中度敏感

Intermediary (I) (mm)

耐药

Drug resistant (R) (mm)

抑菌圈直径

Inhibition zone diameter (mm)

结果

Results

两性霉素B

Amphotericin B

10≥1513-14<1230.00±1.00S

伊曲康唑

Itraconazole

8≥1714-16<1324.66±0.15S

卡泊芬净

Carbenicillin

5≥1714-16<138.00±0.10R

伏立康唑

Voriconazole

1≥1714-16<1313.33±0.20I
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分枝横梗霉LYSF001的毒力基因分析与抗真菌药物敏感性评估
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邹珂宇 1 , 杨彦文 1 , 邢柳 1 , 陈景运 1 , 王利 1 , 魏勇 2
微生物学报 | 研究报告 2025,65(10): 4517-4527
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微生物学报 | 研究报告 2025, 65(10): 4517-4527
分枝横梗霉LYSF001的毒力基因分析与抗真菌药物敏感性评估
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邹珂宇1, 杨彦文1, 邢柳1, 陈景运1, 王利1 , 魏勇2
作者信息
  • 1西南民族大学，青藏高原动物遗传资源保护与利用教育部重点实验室，四川 成都
  • 2四川省畜牧科学研究院，四川 成都
Pathogenicity and fungicide resistance of Lichtheimiaramosa LYSF001: analysis of virulence genes and evaluation of sensitivity to fungicides
Keyu ZOU1, Yanwen YANG1, Liu XING1, Jingyun CHEN1, Li WANG1 , Yong WEI2
Affiliations
  • 1Key Laboratory of Qinghai-Xizang Plateau Animal Genetic Resource Reservation and Exploitation of Ministry of Education, Southwest Minzu University, Chengdu, Sichuan, China
  • 2Sichuan Animal Science Academy, Chengdu, Sichuan, China
出版时间: 2025-09-04 doi: 10.13343/j.cnki.wsxb.20250205
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摘要
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【目的】 探讨分枝横梗霉菌株LYSF001的致病性及其对常见抗真菌药物的耐药性，为动物真菌感染的诊断与治疗提供实验依据。 【方法】 通过形态学特征和分子生物学技术分离并鉴定菌株；利用PCR技术检测菌株LYSF001的毒力基因，并构建动物感染模型以评估其致病性；采用组织病理学检查、GMS染色及ITS序列扩增技术分析感染动物的肝脏和肾脏病理变化；通过药敏试验评估菌株LYSF001对卡泊芬净、两性霉素B和伊曲康唑的敏感性。 【结果】 菌株LYSF001鉴定为分枝横梗霉；检测出菌株LYSF001携带5个毒力基因，分别为CalA、PKP2、LaeA、Alp2和AspF1；该菌株感染小鼠后造成40%的死亡率，试验小鼠内脏充血，其中肝脏和肾脏的病理变化最为显著，表现为炎症细胞浸润和组织坏死；GMS染色显示心脏、肝脏中存在深色菌丝，ITS序列扩增进一步证实了真菌感染；药敏试验证实菌株LYSF001对两性霉素B及伊曲康唑表现出敏感性，然而该菌株对卡泊芬净存在耐药性。 【结论】 本研究证实分枝横梗霉菌株LYSF001具有较强的致病性，可引发动物严重感染甚至死亡；同时，该菌株对两性霉素B和伊曲康唑敏感，但对卡泊芬净耐药。这为动物真菌感染的诊断与治疗提供了重要的实验依据和技术支持。

分枝横梗霉  /  毒力基因  /  致病性  /  耐药性

[Objective] To investigate the pathogenicity of Lichtheimiaramosa LYSF001 and ITS resistance to common fungicides, providing experimental evidence for the diagnosis and treatment of fungal infections in animals. [Methods] The strain was identified by morphological characterization and molecular biology techniques. PCR was employed to detect virulence genes in strain LYSF001, and an animal infection model was established to evaluate the pathogenicity of the strain. Histopathological examination, Grocott’s methenamine silver (GMS) staining, and ITS sequences amplification were employed to analyze pathological changes in the livers and kidneys of infected animals. Additionally, antimicrobial susceptibility tests were conducted to assess the sensitivity of strain LYSF001 to caspofungin, amphotericin B, and itraconazole. [Results] Strain LYSF001 was identified as L. ramosa. PCR analysis revealed that strain LYSF001 carried five virulence genes (CalA, PKP2, LaeA, Alp2, and AspF1). Animal experiments demonstrated that the strain led to the mortality of 40% and caused visceral hyperemia in mice, with the most significant pathological changes (inflammatory cell infiltration and tissue necrosis) observed in the livers and kidneys. Severe infections led to animal mortality. GMS staining revealed the presence of dark-colored hyphae in the heart and liver, and ITS sequences amplification further confirmed fungal infection. The antimicrobial susceptibility test results indicated that strain LYSF001 was resistant to caspofungin but sensitive to amphotericin B and itraconazole. [Conclusion] L. ramosa LYSF001 exhibits strong pathogenicity, capable of causing severe infections and even death in animals. Additionally, the strain showed resistance to caspofungin but sensitivity to amphotericin B and itraconazole. The findings provide important experimental evidence and technical support for the diagnosis and treatment of fungal infections in animals.

Lichtheimiaramosa  /  virulence genes  /  pathogenicity  /  fungicide resistance
邹珂宇, 杨彦文, 邢柳, 陈景运, 王利, 魏勇. 分枝横梗霉LYSF001的毒力基因分析与抗真菌药物敏感性评估. 微生物学报, 2025 , 65 (10) : 4517 -4527 . DOI: 10.13343/j.cnki.wsxb.20250205
Keyu ZOU, Yanwen YANG, Liu XING, Jingyun CHEN, Li WANG, Yong WEI. Pathogenicity and fungicide resistance of Lichtheimiaramosa LYSF001: analysis of virulence genes and evaluation of sensitivity to fungicides[J]. Acta Microbiologica Sinica, 2025 , 65 (10) : 4517 -4527 . DOI: 10.13343/j.cnki.wsxb.20250205
正文
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毛霉病(Mucormycosis)是由毛霉目(Mucorales)霉菌引起的一种严重且致命的侵袭性真菌感染，主要感染免疫功能低下的个体，其病理特征为具有血管侵袭倾向，可导致血栓形成、组织坏死及高死亡率[1]。毛霉病是全球第三大常见的侵袭性真菌病，发病率仅次于念珠菌病和曲霉病[2]，该病主要由毛霉(Mucoromycotina)中的毛霉属(Mucor)、根霉属(Rhizomucor)、根毛霉属(Rhizopus)和横梗霉属(Lichtheimia)，以及被孢霉目(Mortierellomycotina)中的被孢霉属(Mortierella)引起[3]。
作为毛霉病的重要病原体，横梗霉属(Lichtheimia)真菌是一类广泛分布的腐生真菌，常见于土壤和农产品中，尤其对免疫功能低下的个体具有显著影响[4]。分枝横梗霉(Lichtheimiaramosa)可引起人类鼻部坏疽、肺毛霉病、新生儿败血症、坏死性筋膜炎及急性化脓性坏死性炎症[5-9]。在伊朗，分枝横梗霉曾导致毛霉病[6]；在印度，该菌引发了新生儿败血症和坏死性筋膜炎[7]。中国也有报道分枝横梗霉引起的鼻部坏疽和肺毛霉病病例[5,9]。在荷兰，一名38岁终末期肾病的男性腹膜透析患者感染了分枝横梗霉，病理检查显示其肾脏出现急性化脓性坏死性炎症，并伴有广泛菌丝侵袭[8]。此外，分枝横梗霉还在日本引起犊牛纹状体坏死，在土耳其导致牛角质化病变[10-11]。由于不同报道中描述的症状多样，可能与感染途径有关，因此仅凭临床症状不应作为诊断分枝横梗霉感染的依据。鉴于分枝横梗霉感染进展迅速且具有较高的死亡率和发病率，准确诊断对有效治疗至关重要。从分子层面研究致病性霉菌的致病性，多是研究菌株的毒力基因，例如，毒力蛋白CotH通过类似血管紧张素的机制参与毛霉的致病作用[12]；PKR2可作为细胞生长过程的阻遏物；作为核糖体毒性蛋白，AspF1在侵染宿主细胞后，通过特异性抑制核糖体功能阻断蛋白质合成过程，最终诱导宿主细胞发生程序性死亡[13]。此外，在治疗过程中使用靶向抗真菌药物也极为重要。目前用于治疗分枝横梗霉感染的常见抗真菌药物包括艾沙康唑、纳他霉素、两性霉素B、卡泊芬净、泊沙康唑、酮康唑、伏立康唑、伊曲康唑和米卡芬净等[5-9,14-15]。
目前关于分枝横梗霉的研究主要集中在感染后的治疗方面，而对其致病性的了解较少。本研究通过检测菌株LYSF001的毒力基因并进行动物致病性实验，探讨其致病性，并评估多种药物的抗真菌效果。
1 材料与方法
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1.1 伦理审批声明
本研究所有动物实验均按照西南民族大学动物护理与伦理委员会的指导原则进行，编号为SMU-CAVS-230601001。
1.2 菌株与动物
分枝横梗霉菌株从患病的黑山羊中分离获得。20只健康的昆明小鼠(KM小鼠)，每只体重在18-22 g之间，购自成都达硕实验动物有限公司。
1.3 菌株分离
向样本中加入磷酸盐缓冲液(phosphate buffered saline, PBS)后，将其接种至马铃薯葡萄糖琼脂(potato dextrose agar, PDA)培养基中，于37 ℃培养7 d。随后用无菌接种针将纯化后的菌株转移至新鲜的PDA培养基中，继续在37 ℃下培养7 d后，使用乳酸酚棉蓝染色液对样本进行染色，并在显微镜下观察菌丝形态。选取合适的菌落，用2.5%戊二醛固定，经蒸馏水冲洗后，用梯度乙醇脱水。真空干燥2 h后，使用离子喷涂仪在样本表面喷涂铂金，并在扫描电子显微镜(FEI公司)下观察。
1.4 菌株不同温度培养
挑选培养5 d的菌落，挑取菌丝体接种至马铃薯葡萄糖琼脂培养基，分别在18、28和38 ℃恒温培养箱中培养，每个处理设置3个生物学重复；培养期间定时观察，每24 h采用十字交叉法测定菌落生长直径并记录数据，连续观察7 d。实验设置3个300 mL马铃薯葡萄糖液体培养基的锥形瓶，各接种1 mL孢子悬液，于37 ℃恒温振荡培养。在不同培养时间取样，采用分光光度法测定600 nm波长处的吸光度值[16]；以培养时间为自变量，OD600值为因变量，通过测定数据构建菌株LYSF001的生长动力学曲线。
1.5 基因序列扩增与系统发育分析
参照十六烷基三甲基溴化铵提取液(hexadecyl trimethyl ammonium bromide, CTAB) (北京雷根生物技术有限公司)说明书提取菌株总DNA，使用通用引物ITS1 (5′-TCCGTAGGTGA ACCTGCGG-3′)和ITS4 (5′-TCCTCCGCTTATT GATATGC-3′)进行ITS序列扩增。PCR反应体系(25 µL)：Premix Taq 12.5 µL，ITS1、ITS4引物(10 µmol/L)各1 µL，DNA模板1 µL，ddH2O 9.5 µL。PCR反应程序：98 ℃ 2 min；98 ℃ 10 s，56 ℃ 10 s，72 ℃ 30 s，共35个循环；72 ℃ 2 min。采用1%琼脂糖凝胶电泳对扩增产物进行分离纯化后进行测序分析。将获得的ITS序列与NCBI GenBank中真菌的ITS序列进行BLAST比对，并使用MEGA 11.0软件进行系统发育树分析。PCR反应体系所用试剂购自成都微克生物技术有限公司，引物合成和测序由生工生物工程(上海)股份有限公司完成。
1.6 毒力基因检测
参照1.5节的方法提取菌株的总DNA，对菌株LYSF001的毒力基因进行扩增，基因序列见表1。PCR反应程序：98 ℃ 2 min；98 ℃ 10 s，退火15 s，72 ℃ 20 s，共35个循环；72 ℃ 2 min。
1.7 动物实验
将冻存的LYSF001菌种接种于马铃薯葡萄糖琼脂培养基，37 ℃恒温培养7 d。采用无菌磷酸盐缓冲液洗涤孢子后，经纱布过滤8次。获取新鲜孢子悬液后，使用血球计数板测定孢子密度，最终调整孢子悬液浓度为5×107 CFU/mL。将20只昆明小鼠随机分为对照组(C组)和处理组 (T组) (每组n=10)。实验期间，小鼠单独饲养于实验室动物房，温度为28 ℃，正常饮食，适应7 d后进行为期15 d的实验。处理组小鼠腹腔注射0.3 mL分枝横梗霉悬液(5×107 CFU/mL)，对照组小鼠腹腔注射0.3 mL生理盐水。
1.8 RNA提取与实时定量PCR
参照AG RNAex Pro RNA提取试剂盒(四川艾瑞科生物技术有限公司)说明书，提取不同器官的总RNA；参照cDNA合成试剂盒(四川艾瑞科生物技术有限公司)说明书将RNA逆转录成cDNA。使用通用引物ITS1和ITS4进行ITS序列扩增。PCR反应体系同1.5节。PCR反应程序：98 ℃ 2 min；98 ℃ 10 s，56 ℃ 10 s，72 ℃ 30 s，共35个循环；72 ℃ 2 min。
1.9 解剖学与组织病理学检查
通过颈椎脱位法处死所有小鼠，观察并记录不同器官的解剖学结构。采集组织样本并用4%多聚甲醛固定。组织样本经乙醇梯度脱水，二甲苯透明处理25 min后，用石蜡包埋并切片(4-5 μm)用于组织学染色。切片经苏木精和伊红染色后，用中性树胶封片，并在正置显微镜下采集图像。
1.10 透射电子显微镜检查
肝脏和肾脏组织保存于3%戊二醛中，随后用1%锇酸固定2 h。经4 ℃丙酮梯度脱水后，组织样本包埋于树脂中。使用超薄切片机(Leica公司)切割50 nm超薄切片，并在室温下用醋酸铀和柠檬酸铅染色。透射电子显微镜(transmission electron microscope, TEM)图像通过透射电子显微镜(Jeol公司)采集。
1.11 GMS染色
使用银染色剂(gomori methenamine silver, GMS)染色试剂盒(Leagene Biotechnology公司)对切片进行染色。切片经梯度脱水后，用中性树脂封片，并在正置显微镜(Olympus公司)下采集图像。
1.12 药物敏感性试验
本研究选取两性霉素B、伊曲康唑、伏立康唑及卡泊芬净4种抗真菌制剂作为药敏试验药物。将菌株LYSF001的接种悬液制备于无菌PBS中，悬液来自28 ℃培养7 d的PDA培养基。采用纱布过滤获取新鲜孢子悬液后，利用血球计数板进行孢子密度测定，并根据测定结果调整至5×106 CFU/mL。将接种物均匀涂布于平板表面。所有药敏试验均在Mueller-Hinton琼脂上进行，于28 ℃培养1 d后测量抑菌圈直径(以mm为单位)。
1.13 统计分析
所有实验数据均采用SPSS 26.0进行统计学处理。通过单因素方差分析方法评估组间差异，统计学显著性水平设定为P<0.05。实验结果的可视化呈现使用GraphPad Prism 5.0软件完成。
2 结果与分析
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2.1 菌株的培养及鉴定
在PDA培养基中，37 ℃培养7 d。初期(1-2 d)：菌株LYSF001的菌落呈白色絮状，边缘不规则，表面蓬松。中期(3-5 d)：菌丝密度增加，中心区域逐渐转为灰白色，背面开始出现浅褐色色素沉积。成熟期(7 d)：菌落整体呈浓密白色，背面发展为深褐色，并伴随明显的褶皱纹理(图1A)。光学显微镜下可见无隔膜菌丝，分枝频繁，菌丝壁光滑。部分菌丝末端分化出匍匐菌丝，为典型的接合菌生长模式。孢囊梗从匍匐菌丝的节点处垂直伸出，直立或轻微弯曲，表面光滑无附属结构。顶端膨大形成囊轴，呈半球形至圆锥形，与孢子囊连接处界限清晰，孢子囊呈梨形或球形。囊内可见大量孢囊孢子，呈椭圆形或球形，无色透明，表面光滑(图1B)。释放的孢囊孢子散落在周围，部分孢子仍附着于囊轴表面，呈松散聚集状态(图2B)。根据形态学特征初步鉴定菌株LYSF001为毛霉菌中的横梗霉。扫描电镜分析显示：在低倍数下(3 000×)，该菌株呈现典型的无隔多核菌丝结构，菌丝较宽且存在明显的差异；孢子囊着生于直立或微弯的孢囊梗末端，呈球形或梨形，表面具有放射状突起；高倍数下(40 000×)可见椭圆形孢子，其外壁光滑且纹饰稀少，孢子囊形态均一，未见附属结构，并形成特征性的反向包绕胚柄赤道环结构(图1C)，上述显微特征均为横梗霉的典型形态学指征。菌株生长曲线显示，在37 ℃条件下，0-12 h孢子萌发缓慢，12-48 h菌丝生物量迅速增加，OD600值显著上升，约60 h达到生长稳定期，生长速率趋于平缓(图1D)。温度适应性实验显示菌株在38 ℃下生长更快，菌丝颜色更深，而在18 ℃下生长较慢(图1E)。
2.2 菌株基因及毒力基因鉴定
菌株LYSF001的ITS序列长度为865 bp (图2A)。通过比对NCBI数据库中的ITS序列，发现菌株LYSF001与分枝横梗霉MT279293.1的相似度为98.9% (图2B)。共鉴定出5个毒力基因序列，包括CalA、PKP2、LaeA、Alp2和AspF1 (图2C)。
2.3 组织病理学检查
在处理组中，小鼠表现出精神萎靡、毛发粗糙和食欲减退的症状。实验期间，处理组中有4只小鼠死亡，死亡率为40%，而对照组小鼠保持健康且全部存活。处理组小鼠的尸检结果显示，内脏器官粘连，腹腔内有淡黄色液体沉积，肝脏和肾脏出血。在不同器官中检测到ITS序列，且与菌株LYSF001分离鉴定结果一致(图3A)。
肝脏和肾脏的肉眼病理学变化提示需要进一步的组织学检查。GMS染色结果显示，分枝横梗霉在肝脏和肾脏切片中呈棕黑色，在明亮的背景中显著可见。HE染色揭示了多种病理变化：在肝脏中，肝窦扩张并充血，肝索排列紊乱，伴有炎性细胞浸润；肾脏组织中间质血管扩张充血，肾小管上皮细胞呈现肿胀及退行性改变，伴随管腔缩窄(图3B)。
尽管通过GMS和HE染色在光学显微镜下可以详细观察组织结构和细胞变化，但透射电子显微镜(TEM)进一步揭示了超微结构的变化。对照组中，细胞核内染色质分布均匀，线粒体超微结构清晰完整，基质密度高，线粒体嵴清晰可见。在处理组中，肝脏中线粒体数量显著增多，颜色明显加深，基质电子密度增高，嵴间隙变窄并呈现聚集状态；同时观察到肾小管上皮细胞发生变性(图3C)。
2.4 药物敏感性试验
四种抗真菌药物的抑菌圈直径范围总结于表2。两性霉素B (10 µg)纸片的抑菌圈直径最大，其次是伊曲康唑(8 µg)、卡泊芬净(5 µg)和伏立康唑(1 µg)。药敏试验结果表明，LYSF001菌株对两性霉素B与伊曲康唑呈敏感性，伏立康唑显示中等敏感度，而卡泊芬净则表现为完全耐药。
3 讨论
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在本研究中，菌株LYSF001的形态特征与以往研究报道的分枝横梗霉相似[6-7,10]。为了获得更准确和快速的鉴定结果，进一步采用分子生物学方法对菌株LYSF001进行了鉴定。结合ITS和mNGS技术可以提高菌株鉴定的速度和准确性[17-21]。本研究通过PCR分析鉴定出LYSF001菌株的5个毒力基因(CalA、PKP2、LaeA、Alp2和AspF1)，这些基因在真菌感染中可能通过调节次级代谢产物、诱导免疫细胞凋亡等机制发挥重要作用。例如，LaeA基因已被证实通过调控真菌的次级代谢产物和毒力因子来增强其致病性，并可能作为抗真菌药物的潜在靶点[22]。此外，AspF1基因作为一种核毒素[23]，能够抑制宿主细胞的蛋白质合成并诱导免疫细胞凋亡。这些发现与以往研究中关于分枝横梗霉毒力基因的功能描述一致[24-25]，进一步证实了菌株LYSF001的致病潜力，这些毒力基因也可能成为未来抗真菌药物的治疗靶点。
分枝横梗霉最常见的感染部位为皮肤、软组织以及肺部。此外，仅有少数病例涉及播散性感染或鼻脑感染[4]。在其他动物的真菌感染报道中，分枝横梗霉可引起家禽的呼吸道感染[26]和牛的流产[27]。在本研究的动物实验中，菌株LYSF001通过腹腔注射感染小鼠后，主要累及肝脏和肾脏，表现为器官充血、炎症细胞浸润及组织坏死。这一结果与Spithoven等[8]报道的分枝横梗霉感染病例相似，后者也观察到肾脏的急性化脓性坏死性炎症及广泛菌丝侵袭。然而，与William等[7]报道的新生儿坏死性筋膜炎病例不同，本研究中未观察到皮肤和软组织的显著病变，这可能与感染途径和宿主免疫状态的差异有关。
药敏试验证实菌株LYSF001对两性霉素B及伊曲康唑表现出敏感性，然而该菌株对卡泊芬净存在耐药性。这一结果与Gupta等[19]的研究一致，后者也指出两性霉素B和伊曲康唑是治疗毛霉病的首选药物。然而，LYSF001菌株对卡泊芬净的耐药性提示，临床治疗中需谨慎选择抗真菌药物，避免因药物耐药性导致治疗失败。
尽管本研究提供了分枝横梗霉LYSF001的致病性和耐药性数据，但仍存在一些局限性。首先，本研究是对该菌株致病性的初步探究，动物实验仅使用了昆明小鼠作为模型，未能完全模拟山羊等自然宿主的感染情况。其次，实验中的感染途径为腹腔注射，与实际感染途径(如呼吸道或皮肤感染)可能存在差异，这可能影响研究结果的普适性。此外，本研究未对毒力基因的功能进行深入验证，未来可通过基因敲除或过表达实验进一步探讨这些基因在致病过程中的具体作用。
4 结论
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本研究通过形态学观察、分子生物学鉴定及动物实验，系统探讨了分枝横梗霉(Lichtheimiaramosa) LYSF001的致病性及耐药性。本研究证实菌株LYSF001为分枝横梗霉，且具有显著的致病性。检测到该菌株的5个毒力基因，分别为CalA、PKP2、LaeA、Alp2和AspF1，该菌株能够引发实验小鼠的内脏充血，肝脏和肾脏的炎症反应尤为严重，感染后甚至导致试验小鼠死亡。此外，菌株LYSF001对两性霉素B和伊曲康唑敏感，但对卡泊芬净表现出耐药性。这些发现为分枝横梗霉感染的诊断和治疗提供了重要的实验依据。
基金
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  • 四川省肉羊创新团队——防疫岗位(sccxtd-2024-14)
  • 西南民族大学中央高校基本科研业务费专项资金(ZYN2025149)
文献
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文章信息
doi: 10.13343/j.cnki.wsxb.20250205
  • 接收时间：2025-03-17
  • 首发时间：2025-11-03
  • 出版时间：2025-09-04
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  • 收稿日期：2025-03-17
  • 录用日期：2025-05-06
基金
the Sichuan Meat Sheep Innovation Team—Epidemic Prevention Position(sccxtd-2024-14)
四川省肉羊创新团队——防疫岗位(sccxtd-2024-14)
the Southwest Minzu University for the Fundamental Research Funds for the Central Universities(ZYN2025149)
西南民族大学中央高校基本科研业务费专项资金(ZYN2025149)
作者信息
    1西南民族大学，青藏高原动物遗传资源保护与利用教育部重点实验室，四川 成都
    2四川省畜牧科学研究院，四川 成都
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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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