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Article(id=1228017376701116807, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1228017371202388759, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20240618, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1728403200000, receivedDateStr=2024-10-09, revisedDate=null, revisedDateStr=null, acceptedDate=1733068800000, acceptedDateStr=2024-12-02, onlineDate=1770711758065, onlineDateStr=2026-02-10, pubDate=1741017600000, pubDateStr=2025-03-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1770711758065, onlineIssueDateStr=2026-02-10, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1770711758065, creator=13701087609, updateTime=1770711758065, 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=1148, endPage=1161, ext={EN=ArticleExt(id=1228017377296708011, articleId=1228017376701116807, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Untargeted metabolomics reveals the mechanism of Prevotella copri in promoting atherosclerosis in mice, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

[Objective] To explore the potential mechanism by which Prevotella copri promotes atherosclerosis (AS) from the perspective of host-gut microbiota-metabolism. [Methods] ApoE-/- mice were randomized into four groups (n=8): control group (Chow group, fed with a normal diet), model group (AS group, fed with a high-fat diet), low-concentration P. copri group (P. copri-low group, administrated with P. copri at 109 CFU/mL by oral gavage daily from the first day of feeding with the high-fat diet), and high-concentration P. copri group (P. copri-high group, administrated with P. copri at 1011 CFU/mL by oral gavage daily from the first day of feeding with the high-fat diet). The body weight was measured and recorded weekly to evaluate the weight gain trend. After 5 weeks, oil red O staining was employed to evaluate the aortic plaque area, and enzyme-linked immunosorbent assay (ELISA) was employed to measure lipid levels, on the basis of which the impact of P. copri on AS progression was assessed. Additionally, qPCR was used to detect the abundance of P. copri in the gut, and untargeted metabolomics was employed to analyze the metabolite changes in the feces of mice. [Results] Compared with the Chow group, the AS group showed increases in the body weight, aortic plaque area, and plasma levels of low-density lipoprotein cholesterol (LDL-C), total cholesterol (TC), and triglycerides (TG) and a decline in the high-density lipoprotein cholesterol (HDL-C) level. The abundance of P. copri in the gut showed no significant difference between the P. copri-low group and the P. copri-high group, indicating that P. copri successfully colonized the gut in both groups. Based on this, the P. copri-low group was selected as the standard concentration group (P. copri group) for further analysis. Compared with the AS group, P. copri colonization in the gut significantly increased the body weight and aortic plaque area and exacerbated dyslipidemia. Metabolomic analysis revealed that P. copri transplantation led to significant increases in the content of several metabolites, including Cer(d18:1/18:1(9Z)), N-palmitoylsphingosine, genistein, adenine, and linoleic acid. KEGG pathway enrichment analysis further indicated that P. copri might contribute to the development and progression of AS through key pathways such as the regulation of ABC transporters, bile acid metabolism, and neuroactive ligand-receptor interactions. [Conclusion] P. copri may exacerbate inflammation and lipid metabolism imbalance by regulating sphingolipid signaling, purine metabolism, and linoleic acid metabolism, thereby promoting the progression of AS.

, correspAuthors=Jingbin GUO, authorNote=null, correspAuthorsNote=
*E-mail:
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【目的】 从宿主-肠道菌群-代谢的角度，探讨普雷沃氏菌(Prevotella copri)促进动脉粥样硬化(atherosclerosis, AS)的可能作用机制。 【方法】 将ApoE-/-小鼠随机分为4组，每组8只：对照组(Chow组)，喂以普通饲料；模型组(AS组)，喂以高脂饲料；低浓度P. copri移植组(P. copri-low组)，自高脂饮食开始的第1天起每天灌胃109 CFU/mL的P. copri；高浓度P. copri移植组(P. copri-high组)，自高脂饮食开始的第1天起每天灌胃1011 CFU/mL的P. copri。每周测量并记录体重变化，评估体重增长趋势。5周后，应用油红O染色评估主动脉斑块面积，结合酶联免疫吸附法(enzyme-linked immunosorbent assay, ELISA)检测血脂水平，评估P. copri对AS进展的影响。同时，利用实时定量PCR (quantitative PCR, qPCR)检测P. copri在肠道中的丰度，并通过非靶向代谢组学技术分析小鼠粪便代谢物的变化。 【结果】 AS组小鼠的体重、主动脉斑块面积显著增加，血浆低密度脂蛋白胆固醇(low-density lipoprotein cholesterol, LDL-C)、总胆固醇(total cholesterol, TC)和甘油三酯(triglycerides, TG)水平显著升高，高密度脂蛋白胆固醇(high-density lipoprotein cholesterol, HDL-C)水平显著降低，与Chow组相比均存在显著差异。P. copri-low组和P. copri-high组在肠道中的P. copri丰度无显著差异，表明两组P. copri均能在小鼠肠道成功定殖。基于此，后续实验选取P. copri-low组作为标准浓度组(P. copri组)进行分析。与AS组相比，P. copri成功定殖显著增加了小鼠的体重和主动脉斑块面积，并加重了血脂紊乱。代谢组学分析显示，P. copri移植导致多种代谢物显著升高，包括Cer(d18:1/18:1(9Z))、棕榈酰鞘氨醇(N-palmitoylsphingosine)、染料木素(genistein)、腺嘌呤(adenine)和亚油酸(linoleic acid)。KEGG通路富集分析进一步揭示，P. copri通过调控ABC转运蛋白、胆汁酸代谢和神经活性配体-受体相互作用等关键通路参与AS的发生发展。 【结论】 P. copri通过调控鞘脂信号通路、嘌呤代谢和亚油酸代谢，可能加剧炎症和脂质代谢失衡，从而促进AS的进展。

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作者贡献声明

李泽桦：负责研究设计、实验实施、数据分析和论文撰写；曾宇宏：参与研究设计、提供实验技术支持，并协助数据分析；郝卿鋆：负责样品处理、实验数据收集，并参与论文讨论与完善；郭敬宾：指导实验设计和研究方向，审阅论文并提供关键性意见。

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A: Serum total cholesterol (TC) levels; B: Serum triglycerides (TG) levels; C: Serum low-density lipoprotein cholesterol (LDL-C) levels; D: Serum high-density lipoprotein cholesterol (HDL-C) levels. *: P<0.05; **: P<0.01; ***: P<0.001., figureFileSmall=YGA+OCehtbPKAl+NzXl9fQ==, figureFileBig=fmLLZfD5mml4llBZvfvQBA==, tableContent=null), ArticleFig(id=1228088874992332949, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=CN, label=图3, caption=Prevotella copri 对AS小鼠血脂水平的影响, figureFileSmall=YGA+OCehtbPKAl+NzXl9fQ==, figureFileBig=fmLLZfD5mml4llBZvfvQBA==, tableContent=null), ArticleFig(id=1228088876389036191, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=EN, label=Figure 4, caption=Effect of Prevotella copri on atherosclerotic plaque area in ApoE-/- mice (n=8). A: Representative images of en face oil red O staining of aorta and quantification of lesion area; B: Representative oil red O staining of aortic sinus (scale bar=500 µm) and quantification of lesion area. ***: P<0.001., figureFileSmall=Duy0c2eeH0DtnFBGs+nFqw==, figureFileBig=oWwI3odvLIy1ovwJdEoPpA==, tableContent=null), ArticleFig(id=1228088876569391277, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=CN, label=图4, caption=Prevotella copri 对AS小鼠主动脉油红O染色的影响, figureFileSmall=Duy0c2eeH0DtnFBGs+nFqw==, figureFileBig=oWwI3odvLIy1ovwJdEoPpA==, tableContent=null), ArticleFig(id=1228088876661665969, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=EN, label=Figure 5, caption=Partial least squares discriminant analysis (PLS-DA) score plots in positive ion mode (A) and negative ion mode (B)., figureFileSmall=C3+wdEEj4J5ze4yYwE1EFw==, figureFileBig=fOp7eZvwck43bn/ptdUG6w==, tableContent=null), ArticleFig(id=1228088876795883704, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=CN, label=图5, caption=正离子模式和负离子模式的PLS-DA得分比较, figureFileSmall=C3+wdEEj4J5ze4yYwE1EFw==, figureFileBig=fOp7eZvwck43bn/ptdUG6w==, tableContent=null), ArticleFig(id=1228088876904935619, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=EN, label=Figure 6, caption=Partial least squares discriminant analysis (OPLS-DA) score plots and permutation test results in positive and negative ion modes. A: OPLS-DA score plot in positive ion mode; B: OPLS-DA score plot in negative ion mode; C: Permutation test results in positive ion mode; D: Permutation test results in negative ion mode., figureFileSmall=8ei/nP90u82hoKFgqM8PIg==, figureFileBig=EAKqUmYfXNAgZ4WNh6IbmQ==, tableContent=null), ArticleFig(id=1228088877013987531, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=CN, label=图6, caption=正离子模式和负离子模式的OPLS-DA得分图及置换检验结果, figureFileSmall=8ei/nP90u82hoKFgqM8PIg==, figureFileBig=EAKqUmYfXNAgZ4WNh6IbmQ==, tableContent=null), ArticleFig(id=1228088877139816658, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=EN, label=Figure 7, caption=Volcano plots in positive ion mode (A) and negative ion mode (B)., figureFileSmall=2DS/9TLakLr75iHKIPI2Cw==, figureFileBig=utNi2SapIjRsFF5cKDG6WA==, tableContent=null), ArticleFig(id=1228088877261451487, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=CN, label=图7, caption=正离子模式和负离子模式的火山图, figureFileSmall=2DS/9TLakLr75iHKIPI2Cw==, figureFileBig=utNi2SapIjRsFF5cKDG6WA==, tableContent=null), ArticleFig(id=1228088877336948962, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=EN, label=Figure 8, caption=Hierarchical clustering of differential metabolites in positive ion mode (A) and negative ion mode (B)., figureFileSmall=T6W+ANz6W8ITdAzd30agJA==, figureFileBig=nxUO+a/jOfJoqN40xLfNNg==, tableContent=null), ArticleFig(id=1228088877429223656, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=CN, label=图8, caption=正离子模式和负离子模式下差异代谢物的层次聚类结果, figureFileSmall=T6W+ANz6W8ITdAzd30agJA==, figureFileBig=nxUO+a/jOfJoqN40xLfNNg==, tableContent=null), ArticleFig(id=1228088877534081267, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=EN, label=Figure 9, caption=Correlation analysis results of differential metabolites in positive ion mode (A) and negative ion mode (B)., figureFileSmall=TDTvLlwUff0m+ojZSP6MbQ==, figureFileBig=OSuLqyWY1lhs47Nn13ThDw==, tableContent=null), ArticleFig(id=1228088877617967356, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=CN, label=图9, caption=正离子模式和负离子模式的差异代谢物相关性分析结果, figureFileSmall=TDTvLlwUff0m+ojZSP6MbQ==, figureFileBig=OSuLqyWY1lhs47Nn13ThDw==, tableContent=null), ArticleFig(id=1228088877697659141, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=EN, label=Figure 10, caption=KEGG pathway enrichment analysis results in positive ion mode., figureFileSmall=Milb9rIecbLDMrNizt2znA==, figureFileBig=aJY7f320TOr7Voft9U+cWw==, tableContent=null), ArticleFig(id=1228088877823488268, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=CN, label=图10, caption=正离子模式KEGG通路富集分析结果, figureFileSmall=Milb9rIecbLDMrNizt2znA==, figureFileBig=aJY7f320TOr7Voft9U+cWw==, tableContent=null), ArticleFig(id=1228088877915762964, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=EN, label=Figure 11, caption=Correlation between fecal metabolites and AS-related indicators. Data are presented as mean±SD. *: P<0.05; **: P<0.01; ***: P<0.001., figureFileSmall=9hx2JIdt4hbXerA7OckJvw==, figureFileBig=apnEA6z503hbCBXdJY06KQ==, tableContent=null), ArticleFig(id=1228088878045786396, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=CN, label=图11, caption=粪便代谢物与AS相关指标的关联图, figureFileSmall=9hx2JIdt4hbXerA7OckJvw==, figureFileBig=apnEA6z503hbCBXdJY06KQ==, tableContent=null), ArticleFig(id=1228088878154838308, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=EN, label=Table 1, caption=

Significant differential metabolites in feces under positive ion mode

, figureFileSmall=null, figureFileBig=null, tableContent=
NumberMetaboliteVIPP-valueTrendFold change
1Cer(d18:1/18:1(9Z))2.4790.001↑1.706
21-eicosatrienoyl-sn-glycero-3-phosphoethanolamine1.2840.009↑2.065
3All cis-(6,9,12)-linolenic acid2.2670.011↓0.673
4N-(omega)-hydroxyarginine1.5510.011↑3.144
5Genistein1.1810.014↑1.745
6Adenine14.5790.017↑3.194
7Linoleic acid2.7560.022↑1.932
8S-methyl-5′-thioadenosine8.2200.024↑2.868
9N-palmitoylsphingosine2.1470.024↑1.698
10Isoetharine1.7750.025↑2.646
11Sphinganine11.4070.026↑2.408
12Val-Pro1.4410.026↑1.891
13Dimethylaminopurine1.4860.029↑2.895
14Adenosine7.2150.037↑1.759
15N-acetylserotonin1.0230.039↑1.501
16Cholecalciferol (vitamin D3)2.3120.041↑1.507
171-palmitoylglycerol2.0130.045↑2.688
18Indole-2-carboxylic acid9.8310.045↑121.305
19Trans-vaccenic acid1.3820.048↑1.791
), ArticleFig(id=1228088878280667439, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=CN, label=表1, caption=

正离子模式下粪便的显著性差异代谢物

, figureFileSmall=null, figureFileBig=null, tableContent=
NumberMetaboliteVIPP-valueTrendFold change
1Cer(d18:1/18:1(9Z))2.4790.001↑1.706
21-eicosatrienoyl-sn-glycero-3-phosphoethanolamine1.2840.009↑2.065
3All cis-(6,9,12)-linolenic acid2.2670.011↓0.673
4N-(omega)-hydroxyarginine1.5510.011↑3.144
5Genistein1.1810.014↑1.745
6Adenine14.5790.017↑3.194
7Linoleic acid2.7560.022↑1.932
8S-methyl-5′-thioadenosine8.2200.024↑2.868
9N-palmitoylsphingosine2.1470.024↑1.698
10Isoetharine1.7750.025↑2.646
11Sphinganine11.4070.026↑2.408
12Val-Pro1.4410.026↑1.891
13Dimethylaminopurine1.4860.029↑2.895
14Adenosine7.2150.037↑1.759
15N-acetylserotonin1.0230.039↑1.501
16Cholecalciferol (vitamin D3)2.3120.041↑1.507
171-palmitoylglycerol2.0130.045↑2.688
18Indole-2-carboxylic acid9.8310.045↑121.305
19Trans-vaccenic acid1.3820.048↑1.791
), ArticleFig(id=1228088878385525043, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=EN, label=Table 2, caption=

Significant differential metabolites in feces under negative ion mode

, figureFileSmall=null, figureFileBig=null, tableContent=
NumberMetaboliteVIPP-valueTrendFold change
1PGF3a11.0220.007↑6.290
2d-mannose1.4680.024↓0.652
3Ponasterone A1.4410.029↑1.585
4Chenodeoxycholate16.7970.040↑1.896
5Adenine3.8280.046↑3.167
), ArticleFig(id=1228088878498771258, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1228017376701116807, language=CN, label=表2, caption=

负离子模式下粪便的显著性差异代谢物

, figureFileSmall=null, figureFileBig=null, tableContent=
NumberMetaboliteVIPP-valueTrendFold change
1PGF3a11.0220.007↑6.290
2d-mannose1.4680.024↓0.652
3Ponasterone A1.4410.029↑1.585
4Chenodeoxycholate16.7970.040↑1.896
5Adenine3.8280.046↑3.167
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基于非靶向代谢组学的普雷沃氏菌对动脉粥样硬化小鼠作用机制
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李泽桦 1 , 曾宇宏 2 , 郝卿鋆 1 , 郭敬宾 1, *
微生物学报 | 研究报告 2025,65(3): 1148-1161
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微生物学报 | 研究报告 2025, 65(3): 1148-1161
基于非靶向代谢组学的普雷沃氏菌对动脉粥样硬化小鼠作用机制
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李泽桦1, 曾宇宏2, 郝卿鋆1, 郭敬宾1, *
作者信息
  • 1 南方医科大学珠江医院 心血管内科，广东 广州
  • 2 南方医科大学珠江医院 设备器材科，广东 广州
Untargeted metabolomics reveals the mechanism of Prevotella copri in promoting atherosclerosis in mice
Zehua LI1, Yuhong ZENG2, Qingyun HAO1, Jingbin GUO1, *
Affiliations
  • 1 Department of Cardiology, Zhujiang Hospital, Southern Medical University, Guangzhou, Guangdong, China
  • 2 Department of Equipment and Materials, Zhujiang Hospital, Southern Medical University, Guangzhou, Guangdong, China
出版时间: 2025-03-04 doi: 10.13343/j.cnki.wsxb.20240618
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【目的】 从宿主-肠道菌群-代谢的角度，探讨普雷沃氏菌(Prevotella copri)促进动脉粥样硬化(atherosclerosis, AS)的可能作用机制。 【方法】 将ApoE-/-小鼠随机分为4组，每组8只：对照组(Chow组)，喂以普通饲料；模型组(AS组)，喂以高脂饲料；低浓度P. copri移植组(P. copri-low组)，自高脂饮食开始的第1天起每天灌胃109 CFU/mL的P. copri；高浓度P. copri移植组(P. copri-high组)，自高脂饮食开始的第1天起每天灌胃1011 CFU/mL的P. copri。每周测量并记录体重变化，评估体重增长趋势。5周后，应用油红O染色评估主动脉斑块面积，结合酶联免疫吸附法(enzyme-linked immunosorbent assay, ELISA)检测血脂水平，评估P. copri对AS进展的影响。同时，利用实时定量PCR (quantitative PCR, qPCR)检测P. copri在肠道中的丰度，并通过非靶向代谢组学技术分析小鼠粪便代谢物的变化。 【结果】 AS组小鼠的体重、主动脉斑块面积显著增加，血浆低密度脂蛋白胆固醇(low-density lipoprotein cholesterol, LDL-C)、总胆固醇(total cholesterol, TC)和甘油三酯(triglycerides, TG)水平显著升高，高密度脂蛋白胆固醇(high-density lipoprotein cholesterol, HDL-C)水平显著降低，与Chow组相比均存在显著差异。P. copri-low组和P. copri-high组在肠道中的P. copri丰度无显著差异，表明两组P. copri均能在小鼠肠道成功定殖。基于此，后续实验选取P. copri-low组作为标准浓度组(P. copri组)进行分析。与AS组相比，P. copri成功定殖显著增加了小鼠的体重和主动脉斑块面积，并加重了血脂紊乱。代谢组学分析显示，P. copri移植导致多种代谢物显著升高，包括Cer(d18:1/18:1(9Z))、棕榈酰鞘氨醇(N-palmitoylsphingosine)、染料木素(genistein)、腺嘌呤(adenine)和亚油酸(linoleic acid)。KEGG通路富集分析进一步揭示，P. copri通过调控ABC转运蛋白、胆汁酸代谢和神经活性配体-受体相互作用等关键通路参与AS的发生发展。 【结论】 P. copri通过调控鞘脂信号通路、嘌呤代谢和亚油酸代谢，可能加剧炎症和脂质代谢失衡，从而促进AS的进展。

普雷沃氏菌  /  动脉粥样硬化  /  代谢组学  /  ApoE-/-小鼠

[Objective] To explore the potential mechanism by which Prevotella copri promotes atherosclerosis (AS) from the perspective of host-gut microbiota-metabolism. [Methods] ApoE-/- mice were randomized into four groups (n=8): control group (Chow group, fed with a normal diet), model group (AS group, fed with a high-fat diet), low-concentration P. copri group (P. copri-low group, administrated with P. copri at 109 CFU/mL by oral gavage daily from the first day of feeding with the high-fat diet), and high-concentration P. copri group (P. copri-high group, administrated with P. copri at 1011 CFU/mL by oral gavage daily from the first day of feeding with the high-fat diet). The body weight was measured and recorded weekly to evaluate the weight gain trend. After 5 weeks, oil red O staining was employed to evaluate the aortic plaque area, and enzyme-linked immunosorbent assay (ELISA) was employed to measure lipid levels, on the basis of which the impact of P. copri on AS progression was assessed. Additionally, qPCR was used to detect the abundance of P. copri in the gut, and untargeted metabolomics was employed to analyze the metabolite changes in the feces of mice. [Results] Compared with the Chow group, the AS group showed increases in the body weight, aortic plaque area, and plasma levels of low-density lipoprotein cholesterol (LDL-C), total cholesterol (TC), and triglycerides (TG) and a decline in the high-density lipoprotein cholesterol (HDL-C) level. The abundance of P. copri in the gut showed no significant difference between the P. copri-low group and the P. copri-high group, indicating that P. copri successfully colonized the gut in both groups. Based on this, the P. copri-low group was selected as the standard concentration group (P. copri group) for further analysis. Compared with the AS group, P. copri colonization in the gut significantly increased the body weight and aortic plaque area and exacerbated dyslipidemia. Metabolomic analysis revealed that P. copri transplantation led to significant increases in the content of several metabolites, including Cer(d18:1/18:1(9Z)), N-palmitoylsphingosine, genistein, adenine, and linoleic acid. KEGG pathway enrichment analysis further indicated that P. copri might contribute to the development and progression of AS through key pathways such as the regulation of ABC transporters, bile acid metabolism, and neuroactive ligand-receptor interactions. [Conclusion] P. copri may exacerbate inflammation and lipid metabolism imbalance by regulating sphingolipid signaling, purine metabolism, and linoleic acid metabolism, thereby promoting the progression of AS.

Prevotella copri  /  atherosclerosis  /  metabolomics  /  ApoE-/- mice
李泽桦, 曾宇宏, 郝卿鋆, 郭敬宾. 基于非靶向代谢组学的普雷沃氏菌对动脉粥样硬化小鼠作用机制. 微生物学报, 2025 , 65 (3) : 1148 -1161 . DOI: 10.13343/j.cnki.wsxb.20240618
Zehua LI, Yuhong ZENG, Qingyun HAO, Jingbin GUO. Untargeted metabolomics reveals the mechanism of Prevotella copri in promoting atherosclerosis in mice[J]. Acta Microbiologica Sinica, 2025 , 65 (3) : 1148 -1161 . DOI: 10.13343/j.cnki.wsxb.20240618
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根据2019年世界卫生组织的统计，全球约有1 790万人因心血管疾病相关因素死亡，占当年总死亡人数的32%。近30年来，我国在医疗可及性和质量方面取得了显著进展，多项心血管技术已达到或接近世界领先水平。然而，由于不健康生活方式的广泛存在以及人口老龄化的加速，我国心血管疾病的患病率和死亡率仍在持续上升，已成为居民主要的致死原因之一[1]。动脉粥样硬化(atherosclerosis, AS)是心血管疾病的核心病理基础，在引发动脉粥样硬化性心血管疾病(atherosclerotic cardiovascular disease, ASCVD)中发挥着关键作用。深入探讨AS的发病机制，对于理解ASCVD的发生与发展，以及制定有效的治疗策略，具有重要的临床和学术意义。
AS的发生机制复杂，涉及脂质沉积、炎症和氧化应激等多种病理因素[2]。近年来，研究发现这些病理过程不仅受机体内在代谢的调控，还可能与肠道菌群及其代谢产物密切相关[3]。肠道菌群作为一个复杂的生态系统，由细菌、真菌、古生菌和病毒共同组成，其中细菌占比超过99%，在维持肠道微生态平衡和宿主代谢中发挥着核心作用[4]。越来越多的研究显示，肠道菌群的失调与ASCVD及其相关危险因素之间存在密切联系[5-6]。因此，肠道菌群作为潜在的研究与干预靶点，有望为心血管疾病的治疗提供新的思路。
普雷沃氏菌(Prevotella)是一种短杆状、厌氧性革兰氏阴性菌，隶属于普氏菌科普雷沃氏菌属[7]。在以植物为主食的非西方人群中，Prevotella的检出率较高，其中Prevotella copri是该菌属中丰度最高的菌种[8]。研究表明，P. copri能够直接或间接驱动炎症反应，从而促进炎症相关疾病的发生[9]。Li等[10]研究发现，P. copri显著促进AS的进展。然而，关于P. copri干预后代谢谱的变化及其相关机制的研究仍然不足。本研究基于超高效液相色谱-四极杆飞行时间质谱(ultra-high performance liquid chromatography-quadrupole time of flight mass spectrometry, UHPLC-QTOF-MS)的非靶向代谢组学分析，探讨P. copri单菌移植对小鼠粪便代谢组的影响，以期为进一步揭示AS的发病机制提供科学依据。
1 材料与方法
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1.1 主要试剂和仪器
P. copri，北纳生物公司；戊巴比妥钠、乙酸铵，Sigma-Aldrich公司；乙腈，Merck公司；小鼠低密度脂蛋白胆固醇(low-density lipoprotein cholesterol, LDL-C)、总胆固醇(total cholesterol, TC)、甘油三酯(triglycerides, TG)和高密度脂蛋白胆固醇(high-density lipoprotein cholesterol, HDL-C)的酶联免疫吸附法(enzyme-linked immunosorbent assay, ELISA)试剂盒，上海科兴商贸有限公司；4%多聚甲醛固定液，上海碧云天生物技术股份有限公司；油红O染液，武汉赛维尔生物科技有限公司；粪便DNA提取试剂盒，方舟生物安全科技(广州)有限公司；荧光定量检测试剂盒，广州瑞真生物技术有限公司；普通饲料、高脂饲料(含18%脂肪和1.2%胆固醇)，广东省医学实验动物中心。
质谱仪，AB SCIEX公司；超高压液相色谱仪，Agilent公司；台式高速离心机，Eppendorf公司；ACQUITY UPLC HSS T3色谱柱(2.1 mm×100 mm, 1.8 µm)、ACQUITY UPLCBEH Amide色谱柱(2.1 mm×100 mm, 1.7 µm)，Waters公司；酶标仪，ThermoFisher Scientific公司；实时定量PCR仪，Applied Biosystem公司。
1.2 实验动物
本研究采用的雄性ApoE-/-小鼠购自江苏集萃药康生物科技股份有限公司，均为无特定病原体动物(specific pathogen free, SPF)级动物，8周龄，体重20-25 g，合格证编号SYXK(粤)2019-0215。动物实验在南方医科大学珠江医院实验动物中心的SPF级动物房内进行，并已通过南方医科大学珠江医院实验动物伦理审查委员会的批准，批准号为LAEC-2023-066。
1.3 动物分组及给药
动物饲养在恒温环境(24±2) ℃、12 h昼夜交替光照条件下，自由饮水和进食。实验开始前，小鼠适应性饲养1周。随后，将ApoE-/-小鼠随机分为4组：对照组(Chow组)、模型组(AS组)、低浓度P. copri移植组(P. copri-low组)和高浓度P. copri移植组(P. copri-high组)，每组8只。Chow组喂以普通饲料，AS组喂以高脂饲料。自高脂饮食开始的第1天起，P. copri-low组和P. copri-high组分别按照109 CFU/mL、1011 CFU/mL的剂量每日灌胃P. copri，连续5周；AS组小鼠则每日灌胃等体积的生理盐水作为对照。其间，各组小鼠每周称重并记录体重。5周喂养结束后，通过腹部按摩收集小鼠末次灌胃后12 h的粪便样品，并采用实时定量PCR (quantitative PCR, qPCR)检测P. copri在肠道中的定殖情况。基于定殖效果，在后续的药理和代谢组学实验中，选取了P. copri-low组作为标准组(P. copri组)进行进一步分析。
1.4 小鼠粪便中 Prevotella copri 丰度的验证
称取250 mg小鼠粪便样本，使用粪便DNA提取试剂盒提取总DNA。通过qPCR检测P. copri的基因丰度(正向引物序列为5′-GGAG GCAGCAGTGAGGAATA-3′，反向引物序列为5′-ACGCTTAATCTCCGGCCTAC-3′)。qPCR反应体系(20 µL)：SYBR Green qPCR Master Mix 10 µL，上、下游引物(终浓度为0.25 µmol/L)各0.5 µL，模板DNA 1 µL，无核酸酶水8 µL。实验中以16S rRNA基因作为内参(正向引物序列为5′-ACTCCTACGGGAGGCAGCAGT-3′，反向引物序列为5′-ATTACCGCGGCTGCTGGC-3′)，采用2-ΔΔCt方法计算P. copri的相对丰度变化。
1.5 小鼠血脂的检测
在收集粪便样本后，各组小鼠经2%戊巴比妥钠麻醉，通过眼眶后静脉丛采集血液样本。血液样本在4 ℃静置2 h，随后以3 000 r/min离心15 min，分离血清并收集上清液。根据ELISA试剂盒的说明书，分别测定血清中的TG、TC、LDL-C和HDL-C水平。
1.6 小鼠主动脉斑块面积的评估
采血后，将实验小鼠处死，并通过左心室缓慢灌注PBS溶液进行冲洗。随后，从左锁骨下动脉至髂骨分叉处解剖出完整的主动脉和心脏组织，并置于4%多聚甲醛溶液中固定过夜。固定后的主动脉沿纵向切开，并进行油红O染色；固定后的心脏从心底部向主动脉方向切取10 μm厚的冰冻切片，在距三尖瓣100 μm处取横断面，选择包含3个主动脉瓣的切片进行油红O染色。通过显微镜采集图像，利用Image-Pro Plus软件(v6.0.0.260，Media Cybernetics公司)分析图像数据。计算油红O染色阳性面积与血管总面积的比值(油红O阳性面积/血管总面积×100%)以评估整体主动脉的斑块面积；计算油红O染色阳性面积占主动脉根部横截面总面积的百分比以评估主动脉窦的斑块面积。
1.7 小鼠粪便样品的处理
称取60 mg粪便样本，加入200 µL水，充分匀浆并涡旋60 s。随后加入800 µL甲醇-乙腈混合溶液(1:1，体积比)，再次涡旋60 s以确保充分混匀。样本进行低温超声处理30 min，并重复该过程2次以提高提取效率。处理后的样本于-20 ℃静置1 h以促进蛋白沉淀，14 000 r/min离心20 min，收集上清液用于后续分析。
1.8 色谱-质谱分析
收集的上清液样品通过UHPLC配备的HILIC色谱柱进行分离，柱温为25 ℃，流速为0.3 mL/min，流动相A为水中含25 mmol/L乙酸铵和25 mmol/L氨水，流动相B为乙腈。梯度洗脱程序为：0-0.5 min，B维持在95%；0.5-7 min，B线性降至65%；7-8 min，B线性降至40%；8-9 min，B维持在40%；9-9.1 min，B线性回升至95%；9.1-12 min，B维持在95%。样品分离后进行质谱分析。样本检测完成后，使用质谱仪对代谢物进行鉴定，并收集质控样品的一级和二级谱图。二级质谱采用信息依赖采集(information dependent acquisition, IDA)模式，在高灵敏度模式下采集数据，解离电位±60 V，碰撞能量(35±15) eV。最后，采集的数据通过自建的MetDDA和LipDDA数据库进行代谢物结构鉴定。
1.9 数据处理
原始数据使用ProteoWizard软件转换为.mzXML格式，并利用XCMS程序进行峰对齐、保留时间校正及峰面积提取。代谢物的结构鉴定基于精确质量数匹配(<25 ppm)和二级质谱图匹配进行。数据经Pareto-scaling预处理后，进行多维统计分析，包括有监督的偏最小二乘判别分析(partial least squares discriminant analysis, PLS-DA)和正交偏最小二乘判别分析(orthogonal partial least squares discriminant analysis, OPLS-DA)。单变量分析采用Student’s t检验和倍数变化分析，并使用R软件绘制火山图以展示差异代谢物的分布。代谢物显著性通过最佳拟合模型中的变量重要性投影(variable importance in projection, VIP)值进行评估。根据预设的VIP值(VIP>1)和P值(P<0.05)筛选出显著差异代谢物。随后，通过HMDB数据库(Human Metabolome Database, https://hmdb.ca/)对代谢物进行注释验证，并结合KEGG数据库(Kyoto Encyclopedia of Genes and Genomes, https://www.kegg.jp/)对差异代谢物进行代谢通路分析。
1.10 统计学分析
使用SPSS软件(v24.0，IBM公司)进行统计分析，显著性水平设定为P<0.05。两组间差异采用Student’s t检验，3组及以上的差异通过单因素方差分析评估。对于满足方差齐性的情况，采用Tukey检验进行多重比较；若不满足方差齐性，则使用Dunnett-t3检验。符合正态分布的计量数据以均数±标准差(mean±SD)表示。数据可视化使用GraphPad Prism软件(v9.0，GraphPad Software公司)完成。
2 结果与分析
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2.1 Prevotella copri 在小鼠肠道中成功定殖
如图1所示，与Chow组相比，AS组小鼠粪便中P. copri的丰度显著增加(P<0.001)。在P. copri移植后，P. copri-low组和P. copri-high组小鼠粪便中的P. copri丰度均显著高于AS组(P<0.001；P<0.001)。然而，P. copri-low组和P. copri-high组之间的P. copri丰度无显著差异(P>0. 05)，提示P. copri的定殖效应可能在达到阈值后趋于稳定。因此，在后续实验中选取了P. copri-low组作为标准浓度组进行进一步分析。这些结果表明，AS组小鼠粪便中P. copri丰度显著升高；而在P. copri移植组小鼠肠道中，P. copri成功定殖且丰度进一步增加。
2.2 Prevotella copri 促进AS小鼠的体重增长
如图2所示，AS组小鼠的体重显著高于Chow组(P<0.001)。P. copri移植后，小鼠的体重进一步显著增加(P<0.01)。这些结果表明，AS组小鼠的体重显著增加，且P. copri移植进一步加剧了AS小鼠的体重增长。
2.3 Prevotella copri 促进AS小鼠的血脂紊乱
如图3所示，与Chow组相比，AS组小鼠的TC、TG和LDL-C水平显著升高(P<0.001、P<0.001和P<0.01)，而HDL-C水平显著降低(P<0.001)。进一步比较显示，与AS组相比，P. copri移植后的小鼠TC、TG和LDL-C水平进一步显著升高(P<0.05、P<0.001和P<0.001)，而HDL-C水平显著降低(P<0.01)。这些结果表明，AS组小鼠表现出显著的血脂紊乱，而P. copri移植进一步恶化其血脂代谢失衡。
2.4 Prevotella copri 促进小鼠的AS斑块负荷
如图4所示，Chow组小鼠的主动脉未见明显的红染区域。与Chow组相比，AS组小鼠的整体主动脉和主动脉窦斑块负荷显著增加(均为P<0.001)。进一步比较显示，与AS组相比，P. copri组小鼠的整体主动脉和主动脉窦斑块面积进一步显著增大(均为P<0.001)。这些结果表明，高脂饮食成功诱导了AS模型的形成，而P. copri移植能够进一步加重小鼠的AS斑块负荷。
2.5 Prevotella copri 对AS小鼠粪便代谢物的影响
2.5.1 PLS-DA得分
本研究中建立的PLS-DA模型得分图如图5所示。通过7次循环交叉验证获得的模型评价参数显示，P. copri组与AS组在正离子模式下(R2X=0.494，R2Y=0.996，Q2=0.623)和负离子模式下(R2X=0.414，R2Y=0.966，Q2=0.452)均具有良好的分类效果，表明模型具有较高的稳定性。
2.5.2 OPLS-DA得分和置换检验
本研究中建立的OPLS-DA模型得分图如图6A、6B所示。经过7次循环交叉验证后获得的模型评价参数表明，P. copri组与AS组在正离子模式(R2X=0.494，R2Y=0.996，Q2=0.452)和负离子模式(R2X=0.531，R2Y=0.996，Q2=0.573)中均显示出明显的分离，表明模型具有良好的稳定性和预测能力。置换检验如图6C、6D所示，所有Q2值从左到右均低于最右侧的原始蓝色Q2值，说明模型具备较高的稳健性和可靠性，且未出现过拟合现象。
2.5.3 单变量统计分析
在比较两组样本之间的差异代谢物时，火山图常作为一种有效的单变量分析工具。如图7所示，粉色点代表筛选出的差异代谢物。在正离子模式下，P. copri组与AS组的对比中共识别出740种差异代谢物，其中590种呈上调趋势，150种呈下调趋势；在负离子模式下，共识别出936种差异代谢物，其中737种呈上调趋势，199种呈下调趋势。
2.5.4 显著性差异代谢物
在正离子模式下，P. copri组与AS组的对比中共识别出19种显著性差异代谢物(表1)，其中18种代谢物呈上调趋势，1种代谢物呈下调趋势；在负离子模式下，两组对比共有5种显著性差异代谢物(表2)，其中4种代谢物呈上调趋势，1种代谢物呈下调趋势。
2.5.5 差异代谢物聚类分析
如图8所示，同组样本在聚类分析中聚集于同一簇，表明组内样本具有较高的一致性。此外，聚集在同一簇的代谢物具有相似的表达模式，并可能参与代谢通路中相近或相关的反应步骤。
2.5.6 差异代谢物相关性分析
图9展示了正负离子模式下P. copri组显著性差异代谢物之间的相关性。通过相关性分析，评估显著差异代谢物之间的关联性，从而揭示它们在生物状态变化过程中彼此间的相互作用关系。
2.5.7 差异代谢物KEGG通路及其富集分析
如图10所示，本研究发现差异代谢物显著富集于多个关键KEGG通路，包括ABC转运蛋白、嘌呤代谢、鞘脂信号通路、亚油酸代谢和类风湿性关节炎等相关通路。
2.6 差异代谢物与AS相关指标的关联分析
如图11所示，通过对代谢组学分析中显著富集的代谢物与AS相关药理指标(如斑块面积、血脂水平等)进行关联分析，结果显示，Cer(d18:1/18:1(9Z))和adenine等代谢物与主动脉斑块面积及血浆LDL-C、TC和TG水平呈显著正相关，而与HDL-C水平呈显著负相关；相反，all cis-(6,9,12)-linolenic acid等代谢物与主动脉斑块面积及血浆LDL-C、TC和TG水平呈显著负相关，并与HDL-C水平呈显著正相关。这些发现揭示了P. copri可能通过调控上述代谢物，促进AS病变的发生和发展。
3 讨论与结论
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越来越多的研究表明，肠道菌群及其代谢产物是心血管疾病预防和治疗的重要靶标[11-12]。在以植物性饮食为主的非西方人群中，Prevotella型肠道菌群具有较高的流行率，其中P. copri是最为丰富的菌种。P. copri的丰度在类风湿性关节炎、炎症性肠病和2型糖尿病等多种疾病中显著升高，并通过直接或间接途径促进这些炎症性疾病的发展[9,13-15]。本研究初步揭示了P. copri对AS的调控作用。然而，具体机制仍需进一步研究。
代谢组学作为研究代谢产物动态变化的重要技术，已在部分疾病的生物标志物研究中取得了显著进展[16-18]。本研究通过粪便代谢组学进一步揭示了P. copri在AS中的潜在致病机制。(1) 鞘脂代谢通路的激活：Cer(d18:1/18:1(9Z))和棕榈酰鞘氨醇(N-palmitoylsphingosine)等鞘氨醇类脂及其衍生物显著升高。这些代谢物不仅在炎症反应中发挥关键作用，还与脂质代谢紊乱密切相关。鞘氨醇类脂的积累可能通过激发动脉壁的炎症反应和脂质沉积，加剧AS的进展[19]。P. copri移植后这些代谢物显著升高，表明其可能通过激活鞘脂代谢通路促进AS的发展。(2) 脂肪酸代谢的双重作用：亚油酸(linoleic acid)和反式-11-十八烯酸(trans-vaccenic acid)等多不饱和脂肪酸广泛参与炎症和氧化应激反应[20]。亚油酸及其代谢产物(如花生四烯酸)在炎症反应中具有双重调控作用，而P. copri可能通过调节这些脂肪酸代谢，加重AS中的炎症和氧化应激[21]。这些变化表明P. copri通过脂肪酸代谢通路影响AS的病理进展。(3) 特定代谢物的促炎潜力：染料木素(genistein)虽具有抗炎和抗氧化特性，但在P. copri移植后的特定炎症环境中，可能通过雌激素受体途径发挥促炎作用[22]。这一现象进一步揭示了肠道菌群对特定代谢产物双重特性调控的复杂性。(4) 嘌呤代谢的扰动：腺嘌呤(adenine)和腺苷(adenosine)是嘌呤代谢的重要成分，尽管adenosine在局部环境中具有抗炎作用，其长期升高可能导致血管收缩和血小板聚集，诱发心血管不良反应[23]。通过关联分析，我们发现P. copri组显著富集的代谢物与AS相关指标(如斑块面积、血脂水平)密切相关，表明P. copri可能通过改变代谢物平衡加速AS的病理进程。
在KEGG通路富集分析中，我们发现P. copri可能通过多种代谢通路推动AS的进展，特别是ABC转运蛋白、嘌呤代谢和鞘脂信号通路等关键代谢途径。P. copri的调控可能导致胆固醇在动脉壁的积累以及炎症反应的增强，从而加剧AS的发展。此外，胆汁酸代谢的失衡也可能导致血清胆固醇水平的升高，而亚油酸代谢和溶酶体功能的改变则表明，P. copri可能通过增加炎症和氧化应激，扰乱脂质代谢平衡，进一步推动AS的病理进程。综上所述，本研究揭示了P. copri在AS中的多重致病机制，特别是通过调控代谢通路和炎症反应，显著加速了AS的病理进展。这些发现为理解肠道菌群在心血管疾病中的作用提供了重要线索，并为未来开发基于微生物的AS干预策略提供了科学依据。
基金
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  • 国家自然科学基金(82200442)
  • 广州市科技计划(2025A04J3768)
文献
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doi: 10.13343/j.cnki.wsxb.20240618
  • 接收时间：2024-10-09
  • 首发时间：2026-02-10
  • 出版时间：2025-03-04
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  • 收稿日期：2024-10-09
  • 录用日期：2024-12-02
基金
National Natural Science Foundation of China(82200442)
国家自然科学基金(82200442)
Science and Technology Program of Guangzhou(2025A04J3768)
广州市科技计划(2025A04J3768)
作者信息
    1 南方医科大学珠江医院 心血管内科，广东 广州
    2 南方医科大学珠江医院 设备器材科，广东 广州

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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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