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Article(id=1226956554264556362, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226956547847275311, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250174, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1741017600000, receivedDateStr=2025-03-04, revisedDate=null, revisedDateStr=null, acceptedDate=1744041600000, acceptedDateStr=2025-04-08, onlineDate=1770458838287, onlineDateStr=2026-02-07, pubDate=1756915200000, pubDateStr=2025-09-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1770458838287, onlineIssueDateStr=2026-02-07, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1770458838287, creator=13701087609, updateTime=1770458838287, updator=13701087609, issue=Issue{id=1226956547847275311, tenantId=1146029695717560320, journalId=1192105938417971205, year='2025', volume='65', issue='9', pageStart='3821', pageEnd='4232', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1770458836757, creator=13701087609, updateTime=1770459153781, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1226957877613605816, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226956547847275311, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1226957877613605817, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226956547847275311, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=4119, endPage=4135, ext={EN=ArticleExt(id=1226956558593077252, articleId=1226956554264556362, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Effects of nicotinamide mononucleotide/nicotinamide riboside supplementation on gut microbiota and metabolites in mice, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

Nicotinamide mononucleotide (NMN) and nicotinamide riboside (NR) are two precursors of the redox coenzyme nicotinamide adenine dinucleotide (NAD). Due to their ability to effectively increase NAD levels without toxic side effects, they have garnered significant attention. [Objective] This study employed an integrated approach combining metabolomics and 16S rRNA gene sequencing to investigate and compare the effects of NMN or NR supplementation on growth performance, intestinal health, gut microbiota, and metabolites in mice. [Methods] Male C57 mice were randomly allocated into a control group and two experimental groups. The experimental groups received drinking water supplemented with NMN or NR via gavage. Non-targeted metabolomics analysis and 16S rRNA gene sequencing were conducted to systematically investigate the alterations in endogenous metabolites and gut microbiota composition. Furthermore, the expression levels of associated factors were quantified by PCR. [Results] Compared with the control group, supplementation with NMN or NR increased the richness and diversity and improved the composition of gut microbiota in mice, with NMN showing greater efficacy. Additionally, both NMN and NR supplementation significantly upregulated the expression levels of metabolites associated with anti-inflammatory and antioxidant activities. PCR results indicated that both NMN and NR suppressed the expression of pro-inflammatory cytokines. Furthermore, NMN supplementation significantly increased the number of goblet cells in the colon, thereby enhancing the intestinal barrier function. [Conclusion] Dietary supplementation with NMN or NR improves the gut microbiota composition in mice, elevates the levels of beneficial metabolites, and inhibits the expression of pro-inflammatory cytokines.

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*E-mail:
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烟酰胺单核苷酸(nicotinamide mononucleotide, NMN)和烟酰胺核苷(nicotinamide riboside, NR)是氧化还原辅酶烟酰胺腺嘌呤二核苷酸(nicotinamide adenine dinucleotide, NAD)的2种前体物质，因其能够有效增加机体NAD浓度且无毒副作用而受到广泛关注。 【目的】 联合代谢组学和16S rRNA基因测序技术，探讨饲喂NMN或NR对小鼠生长性能、肠道健康、肠道微生物群和代谢物的影响，并比较两者的功效。 【方法】 将雄性C57小鼠随机分为对照组和2个试验组，试验组分别灌胃添加NMN或NR的饮用水，利用非靶向代谢组学技术和16S rRNA基因测序技术联合分析小鼠内源性代谢物和肠道菌群的变化，并通过PCR检测相关因子的表达。 【结果】 与对照组相比，添加NMN或NR均可增加小鼠肠道微生物的丰富度和多样性，并改善微生物群的组成结构，其中添加NMN的效果更显著。与对照组相比，添加NMN和NR均可显著上调抗炎和抗氧化相关代谢物的水平；PCR检测结果表明，NMN和NR均可抑制促炎因子的表达。此外，添加NMN还可显著增加结肠杯状细胞的数量，从而增强肠道屏障功能。 【结论】 饲喂NMN或NR可改善小鼠肠道微生物群结构，增加有益代谢物，并抑制促炎因子的表达。

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作者贡献声明

倪冰：试验设计及开展，数据收集和统计分析及文章撰写等；朱莉莉：完成部分试验，数据收集，文章修改；陈好：文章修改；史新娥：试验指导，文章修改；靳建军：试验方案设计及指导，文章修改及审核。

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tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=CN, orderNo=3, keyword=烟酰胺单核苷酸), Keyword(id=1226964055655625470, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=CN, orderNo=4, keyword=烟酰胺核苷)], refs=[Reference(id=1226964060072227831, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, doi=null, pmid=null, pmcid=null, year=2019, volume=76, issue=3, pageStart=473, pageEnd=493, url=null, language=null, rfNumber=[1], rfOrder=0, authorNames=ADAK A, KHAN MR, journalName=Cellular and Molecular Life Sciences, refType=null, unstructuredReference=ADAK A, KHAN MR. 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Nature Metabolism, 2019, 1(1): 47-57., articleTitle=Slc12a8 is a nicotinamide mononucleotide transporter, refAbstract=null)], funds=[Fund(id=1226964057924744120, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, awardId=Z1090223017, language=EN, fundingSource=Fundamental Research Funds for the Central Universities(Z1090223017), fundOrder=null, country=null), Fund(id=1226964058050573250, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, awardId=Z1090223017, language=CN, fundingSource=中央高校基本科研业务费专项资金(Z1090223017), fundOrder=null, country=null), Fund(id=1226964058159625163, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, awardId=QCYRCXM-2022-212, language=EN, fundingSource=Qin Chuang Yuan Innovation and Entrepreneurship Talent Project(QCYRCXM-2022-212), fundOrder=null, country=null), Fund(id=1226964059564717014, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, awardId=QCYRCXM-2022-212, language=CN, fundingSource=陕西省秦创原引用高层次创新创业人才项目(QCYRCXM-2022-212), fundOrder=null, country=null), Fund(id=1226964059682157535, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, awardId=2022GD-TSLD-46, language=EN, fundingSource=Shaanxi Livestock and Poultry Breeding Double-chain Fusion Key Project(2022GD-TSLD-46), fundOrder=null, country=null), Fund(id=1226964059820569572, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, awardId=2022GD-TSLD-46, language=CN, fundingSource=陕西省畜禽育种“两链”融合重点项目(2022GD-TSLD-46), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1226964051561984489, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, xref=null, ext=[AuthorCompanyExt(id=1226964051570373098, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, companyId=1226964051561984489, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=Key Laboratory of Animal Genetic Breeding and Reproduction in Shaanxi Province, College of Animal Science and Technology, Northwest A&F University, Yangling, Shaanxi, China), AuthorCompanyExt(id=1226964051599733231, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, companyId=1226964051561984489, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=西北农林科技大学 动物科技学院，陕西省动物遗传育种与繁殖重点实验室，陕西 杨凌)])], figs=[ArticleFig(id=1226964055911478041, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=EN, label=Figure 1, caption=Effects of feeding NMN or NR on growth performance of mice. A: Changes in mouse body weight; B: Statistics of mouse food intake; C: Statistics of mouse muscle, fat and liver tissue weight (QU: Quadriceps femoris; GAS: Gastrocnemius; TA: Tibialis anterior; SOL: Soleus; EDL: Extensor digitorum longus; iWAT: Inguinal white adipose tissue; eWAT: Epididymal white adipose tissue; BAT: Brown adipose tissue); D: Statistics of mouse running distance; E: Statistics of mouse running time; F: Statistics of mouse grasping ability., figureFileSmall=ZgQO6odha4Un1aAsxTgf7A==, figureFileBig=D2ensd1FzLgyySc5qUV1BQ==, tableContent=null), ArticleFig(id=1226964056054084389, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=CN, label=图1, caption=饲喂NMN和NR对小鼠生长性能的影响。A：小鼠体重变化；B：小鼠采食量统计；C：小鼠肌肉、脂肪以及肝脏组织质量统计(QU：股四头肌；GAS：腓肠肌；TA：胫骨前肌；SOL：比目鱼肌；EDL：趾长伸肌；iWAT：腹股沟白色脂肪组织；eWAT：附睾白色脂肪组织；BAT：棕色脂肪组织)；D：小鼠跑步距离统计；E：小鼠跑步时间统计；F：小鼠抓握能力统计。, figureFileSmall=ZgQO6odha4Un1aAsxTgf7A==, figureFileBig=D2ensd1FzLgyySc5qUV1BQ==, tableContent=null), ArticleFig(id=1226964056171524913, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=EN, label=Figure 2, caption=Effects of feeding NMN or NR on intestinal morphology in mice. A: H&E staining results of mouse intestine; B: Alcian blue staining results of mouse colon; C: Statistics of mouse small intestinal villus height; D: Statistics of mouse small intestinal crypt depth; E: Ratio of mouse intestinal villus height to crypt depth (V/C); F: Statistics of mouse colon goblet cells., figureFileSmall=acl+nvUKY56X7aRHts+xKw==, figureFileBig=1YP49/qyqLI2sNhTM0BRzw==, tableContent=null), ArticleFig(id=1226964056301548349, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=CN, label=图2, caption=饲喂NMN和NR对小鼠肠道形态的影响。A：小鼠肠道H&E染色结果；B：小鼠结肠阿利新蓝染色结果；C：小鼠小肠绒毛高度统计；D：小鼠小肠隐窝深度统计；E：小鼠肠绒毛高度与隐窝深度比值(V/C)；F：小鼠结肠杯状细胞数量统计。, figureFileSmall=acl+nvUKY56X7aRHts+xKw==, figureFileBig=1YP49/qyqLI2sNhTM0BRzw==, tableContent=null), ArticleFig(id=1226964056431571786, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=EN, label=Figure 3, caption=Effects of feeding NMN or NR on intestinal inflammatory factors in mice. A: PCR expression results of tumor necrosis factor-α (TNF-α); B: PCR expression results of interleukin 6 (IL-6); C: PCR expression results of nuclear factor kappa-light-chain-enhancer of activated B cells; D: PCR expression results of interleukin-1β (IL-1β)., figureFileSmall=XXx3i5klcg2ip6oRhkAFqA==, figureFileBig=/Ou/gFhLye8v2uf/u/JHWQ==, tableContent=null), ArticleFig(id=1226964056578372439, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=CN, label=图3, caption=饲喂NMN和NR对小鼠肠道炎症因子的影响。A：肿瘤坏死因子(tumor necrosis factor-α, TNF-α)的PCR表达结果；B：白细胞介素6 (interleukin 6, IL-6) 的PCR表达结果；C：核因子-kB通路(nuclear factor kappa-light-chain-enhancer of activated B cells)的PCR表达结果；D：白细胞介素-1β (interleukin-1β, IL-1β)的PCR表达结果。, figureFileSmall=XXx3i5klcg2ip6oRhkAFqA==, figureFileBig=/Ou/gFhLye8v2uf/u/JHWQ==, tableContent=null), ArticleFig(id=1226964056670647131, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=EN, label=Figure 4, caption=Effects of feeding NMN and NR on intestinal microbial diversity in mice. A: Comparative analysis of gut microbial alpha diversity between the control group and the NMN-treated group in mice; B: Comparative analysis of gut microbial alpha diversity between the control group and the NR-treated group in mice; C: Comparative analysis of gut microbial alpha diversity among the control group, NMN-treated group, and NR-treated group in mice., figureFileSmall=5bssNRYgT+9VTdPsjMTpMw==, figureFileBig=IokLJlRgKr5V3MGVGfrE4g==, tableContent=null), ArticleFig(id=1226964056783893351, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=CN, label=图4, caption=饲喂NMN和NR对小鼠肠道微生物多样性的影响。A：对照组和NMN组小鼠肠道微生物α多样性结果比较；B：对照组和NR组小鼠肠道微生物α多样性结果比较；C：对照组、NMN组和NR组小鼠肠道微生物α多样性结果比较。, figureFileSmall=5bssNRYgT+9VTdPsjMTpMw==, figureFileBig=IokLJlRgKr5V3MGVGfrE4g==, tableContent=null), ArticleFig(id=1226964056871973739, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=EN, label=Figure 5, caption=Effects of feeding NMN and NR on intestinal microbial diversity in mice. A: Beta diversity results; B: Venn diagram of species composition; C: The species abundance at the phylum level in the NMN-fed group; D: The species abundance at the phylum level in the NR-fed group; E: The ratio of Bacillota to Bacteroidota in the NMN-fed group; F: The ratio of Bacillota to Bacteroidota in the NR-fed group; G: Results of species difference analysis at the phylum level; H: The species abundance at the genus level in the NMN-fed group; I: The species abundance at the genus level in the NR-fed group; J: Results of species difference analysis at the genus level., figureFileSmall=TVxGFRRZRl0ZRzano3nYyA==, figureFileBig=MCWm1E6lQGugdmFxXDNZRA==, tableContent=null), ArticleFig(id=1226964056955859830, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=CN, label=图5, caption=饲喂NMN和NR对小鼠肠道微生物多样性的影响。A：β多样性结果；B：物种组成韦恩图；C：饲喂NMN组门水平物种丰度；D：饲喂NR组门水平物种丰度；E：饲喂NMN组芽孢杆菌门(Bacillota)和拟杆菌门(Bacteroidota)的比值；F：饲喂NR组芽孢杆菌门(Bacillota)和拟杆菌门(Bacteroidota)的比值；G：门水平物种差异分析结果；H：饲喂NMN组属水平物种丰度；I：饲喂NR组属水平物种丰度；J：属水平物种差异分析结果。, figureFileSmall=TVxGFRRZRl0ZRzano3nYyA==, figureFileBig=MCWm1E6lQGugdmFxXDNZRA==, tableContent=null), ArticleFig(id=1226964057052328826, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=EN, label=Figure 6, caption=Effects of feeding NMN and NR on intestinal metabolites in mice. A: OPLS-DA score graph of the NMN-fed group; B:OPLS-DA score graph of the NR-fed group; C: Heat map of differential metabolites in the NMN-fed group; D: Heat map of differential metabolites in the NR-fed group; E: KEGG enrichment results of NMN-fed group; F: KEGG enrichment results of the NR-fed group., figureFileSmall=8x8j/hcsVVD9CjmffDT5UA==, figureFileBig=TNRUjBrGLNnwV8eggO1hTw==, tableContent=null), ArticleFig(id=1226964057152992132, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=CN, label=图6, caption=饲喂NMN和NR对小鼠肠道代谢物的影响。A：饲喂NMN组OPLS-DA得分图；B：饲喂NR组OPLS-DA得分图；C：饲喂NMN组差异代谢物热图；D：饲喂NR组差异代谢物热图；E：饲喂NMN组KEGG富集结果；F：饲喂NR组KEGG富集结果。, figureFileSmall=8x8j/hcsVVD9CjmffDT5UA==, figureFileBig=TNRUjBrGLNnwV8eggO1hTw==, tableContent=null), ArticleFig(id=1226964057245266827, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=EN, label=Figure 7, caption=Results of association analysis between intestinal microorganisms and metabolites. A: Clustering heat map of the correlation between intestinal microorganisms and metabolites in the NMN group vs. control group; B: Clustering heat map of the correlation between intestinal microorganisms and metabolites in the NR group vs. control group., figureFileSmall=Ifi1QoCSZXZEHSVT4U3n/Q==, figureFileBig=cdjF3zycQAZ85u7aqfRrQg==, tableContent=null), ArticleFig(id=1226964057333347221, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=CN, label=图7, caption=肠道微生物与代谢物关联分析结果。A：NMN组与对照组肠道微生物与代谢物相关性聚类热图；B：NR组与对照组肠道微生物与代谢物相关性聚类热图。, figureFileSmall=Ifi1QoCSZXZEHSVT4U3n/Q==, figureFileBig=cdjF3zycQAZ85u7aqfRrQg==, tableContent=null), ArticleFig(id=1226964057446593435, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=EN, label=Table 1, caption=

PCR primer used in this study

, figureFileSmall=null, figureFileBig=null, tableContent=
Primers namePrimer sequences (5′→3′)
TNF-α-FGGTGCCTATGTCTCAGCCTCTT
TNF-α-RGCCATAGAACTGATGAGAGGGAG
IL-1β-FGAAATGCCACCTTTTGACAGTG
IL-1β-RTGGATGCTCTCATCAGGACAG
NF-ĸB-FGACACGACAGAATCCTCAGCATCC
NF-ĸB-RCCACCAGCAGCAGCAGACATG
IL-6-FTGCAAGAGACTTCCATCCAGT
IL-6-RGTGAAGTAGGGAAGGCCG
β-actin-FGTCCACCTTCCAGCAGATGT
β-actin-RGAAAGGGTGTAAAACGCAGC
), ArticleFig(id=1226964057601782692, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226956554264556362, language=CN, label=表1, caption=

本研究所用PCR引物

, figureFileSmall=null, figureFileBig=null, tableContent=
Primers namePrimer sequences (5′→3′)
TNF-α-FGGTGCCTATGTCTCAGCCTCTT
TNF-α-RGCCATAGAACTGATGAGAGGGAG
IL-1β-FGAAATGCCACCTTTTGACAGTG
IL-1β-RTGGATGCTCTCATCAGGACAG
NF-ĸB-FGACACGACAGAATCCTCAGCATCC
NF-ĸB-RCCACCAGCAGCAGCAGACATG
IL-6-FTGCAAGAGACTTCCATCCAGT
IL-6-RGTGAAGTAGGGAAGGCCG
β-actin-FGTCCACCTTCCAGCAGATGT
β-actin-RGAAAGGGTGTAAAACGCAGC
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添加烟酰胺单核苷酸/烟酰胺核苷对小鼠肠道微生物及代谢物的影响
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倪冰 , 朱莉莉 , 陈好 , 史新娥 , 靳建军
微生物学报 | 研究报告 2025,65(9): 4119-4135
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微生物学报 | 研究报告 2025, 65(9): 4119-4135
添加烟酰胺单核苷酸/烟酰胺核苷对小鼠肠道微生物及代谢物的影响
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倪冰, 朱莉莉, 陈好, 史新娥, 靳建军
作者信息
  • 西北农林科技大学 动物科技学院，陕西省动物遗传育种与繁殖重点实验室，陕西 杨凌
Effects of nicotinamide mononucleotide/nicotinamide riboside supplementation on gut microbiota and metabolites in mice
Bing NI, Lili ZHU, Hao CHEN, Xin’e SHI, Jianjun JIN
Affiliations
  • Key Laboratory of Animal Genetic Breeding and Reproduction in Shaanxi Province, College of Animal Science and Technology, Northwest A&F University, Yangling, Shaanxi, China
出版时间: 2025-09-04 doi: 10.13343/j.cnki.wsxb.20250174
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摘要
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烟酰胺单核苷酸(nicotinamide mononucleotide, NMN)和烟酰胺核苷(nicotinamide riboside, NR)是氧化还原辅酶烟酰胺腺嘌呤二核苷酸(nicotinamide adenine dinucleotide, NAD)的2种前体物质，因其能够有效增加机体NAD浓度且无毒副作用而受到广泛关注。 【目的】 联合代谢组学和16S rRNA基因测序技术，探讨饲喂NMN或NR对小鼠生长性能、肠道健康、肠道微生物群和代谢物的影响，并比较两者的功效。 【方法】 将雄性C57小鼠随机分为对照组和2个试验组，试验组分别灌胃添加NMN或NR的饮用水，利用非靶向代谢组学技术和16S rRNA基因测序技术联合分析小鼠内源性代谢物和肠道菌群的变化，并通过PCR检测相关因子的表达。 【结果】 与对照组相比，添加NMN或NR均可增加小鼠肠道微生物的丰富度和多样性，并改善微生物群的组成结构，其中添加NMN的效果更显著。与对照组相比，添加NMN和NR均可显著上调抗炎和抗氧化相关代谢物的水平；PCR检测结果表明，NMN和NR均可抑制促炎因子的表达。此外，添加NMN还可显著增加结肠杯状细胞的数量，从而增强肠道屏障功能。 【结论】 饲喂NMN或NR可改善小鼠肠道微生物群结构，增加有益代谢物，并抑制促炎因子的表达。

肠道微生物  /  代谢组学  /  烟酰胺单核苷酸  /  烟酰胺核苷

Nicotinamide mononucleotide (NMN) and nicotinamide riboside (NR) are two precursors of the redox coenzyme nicotinamide adenine dinucleotide (NAD). Due to their ability to effectively increase NAD levels without toxic side effects, they have garnered significant attention. [Objective] This study employed an integrated approach combining metabolomics and 16S rRNA gene sequencing to investigate and compare the effects of NMN or NR supplementation on growth performance, intestinal health, gut microbiota, and metabolites in mice. [Methods] Male C57 mice were randomly allocated into a control group and two experimental groups. The experimental groups received drinking water supplemented with NMN or NR via gavage. Non-targeted metabolomics analysis and 16S rRNA gene sequencing were conducted to systematically investigate the alterations in endogenous metabolites and gut microbiota composition. Furthermore, the expression levels of associated factors were quantified by PCR. [Results] Compared with the control group, supplementation with NMN or NR increased the richness and diversity and improved the composition of gut microbiota in mice, with NMN showing greater efficacy. Additionally, both NMN and NR supplementation significantly upregulated the expression levels of metabolites associated with anti-inflammatory and antioxidant activities. PCR results indicated that both NMN and NR suppressed the expression of pro-inflammatory cytokines. Furthermore, NMN supplementation significantly increased the number of goblet cells in the colon, thereby enhancing the intestinal barrier function. [Conclusion] Dietary supplementation with NMN or NR improves the gut microbiota composition in mice, elevates the levels of beneficial metabolites, and inhibits the expression of pro-inflammatory cytokines.

gut microbiota  /  metabolomics  /  nicotinamide mononucleotide  /  nicotinamide riboside
倪冰, 朱莉莉, 陈好, 史新娥, 靳建军. 添加烟酰胺单核苷酸/烟酰胺核苷对小鼠肠道微生物及代谢物的影响. 微生物学报, 2025 , 65 (9) : 4119 -4135 . DOI: 10.13343/j.cnki.wsxb.20250174
Bing NI, Lili ZHU, Hao CHEN, Xin’e SHI, Jianjun JIN. Effects of nicotinamide mononucleotide/nicotinamide riboside supplementation on gut microbiota and metabolites in mice[J]. Acta Microbiologica Sinica, 2025 , 65 (9) : 4119 -4135 . DOI: 10.13343/j.cnki.wsxb.20250174
正文
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人类肠道微生物群主要由芽孢杆菌门(Bacillota)、拟杆菌门(Bacteroidota)、放线菌门(Actinomycetota)、梭杆菌门(Fusobacteriota)、假单胞菌门(Pseudomonadota)、疣微菌门(Verrucomicrobiota)和蓝细菌门(Cyanobacteriota) 7个菌门组成[1]。肠道微生物群具有促进食物分解，使其更容易吸收和消化、合成必需维生素、去除有毒化合物、抵抗病原体、维持肠黏膜完整性和调节免疫功能等[2]。肠道微生物群的失衡会影响肠黏膜完整性，引起微生物群落侵入黏膜细胞，诱导代谢性疾病的发生，如肥胖、2型糖尿病和非酒精性脂肪肝等疾病[3]。肠道微生物群对宿主生理机能的益处可归纳为三大类：为宿主提供能量和营养、调节免疫系统的发育和增强宿主防御能力。肠道微生物群的组成与功能的异常变化通常被称为菌群失调，可导致宿主-微生物群相互作用的破坏和宿主免疫系统的紊乱[4]。近年来，鉴于肠道微生物群的重要性和潜在价值，微生物群的组成及调节已成为肠道健康的重要研究方向。
烟酰胺腺嘌呤二核苷酸(nicotinamide adenine dinucleotide, NAD)是氧化还原反应中的重要辅酶，作为糖酵解、氧化磷酸化及三羧酸循环中关键酶的辅助因子发挥着重要作用[5]。NAD+是NAD的氧化型，与多种关键细胞功能直接或间接相关，包括保护线粒体功能、维持氧化还原稳态、发挥抗炎作用、抗衰老、调节细胞分化、影响基因组稳定性及表观遗传修饰等[6]。研究表明NAD+在肠杯状细胞中可以刺激黏蛋白产生，从而维持肠黏膜的完整性，对肠道免疫屏障起到促进作用[7]。根据现有研究，细胞内合成NAD+有3种途径，涉及许多不同的前体化合物。通过生物合成前体增加NAD+水平，可能有助于预防或缓解多种疾病，尤其是代谢疾病和与年龄相关的疾病。哺乳动物自身无法合成NAD+，因此必须通过摄入色氨酸(Trp)或多种膳食来源获取NAD前体，包括烟酰胺单核苷酸(nicotinamide mononucleotide, NMN)、烟酸(nicotinic acid, NA)、烟酰胺(nicotinamide, NAM)、烟酰胺核苷(nicotinamide riboside, NR)等[8]。已有研究表明，NR和NMN对机体的耐受性良好[9-10]，能有效提高各组织中的NAD+浓度，而NA则被报道会引起严重的恶心和潮红反应，NAM则可能导致肝毒性[11]。因此NAD的前体烟酰胺单核苷酸(NMN)和烟酰胺核苷(NR)受到了广泛关注。
NMN和NR对机体生理功能的有益作用已被充分证实。NMN给药可改善由高脂饮食诱导的肝脏胰岛素抵抗，从而发挥抗糖尿病作用；NMN还能提升骨骼肌、肝脏和心脏等代谢器官的线粒体功能，并对大脑的神经元功能具有正向改善作用[12-13]；NMN还对肠黏膜提供保护作用，积极促进肠道健康[14]。NR同样对改善小鼠的代谢健康状况显示出良好的效果，包括缓解肥胖、维持肌肉稳态以及增强小鼠的耐力[15]；NR还可促进成肌细胞的分化，并调节肠道微生物群的组成[16]。因此通过补充NR提高NAD含量被认为是一种有前景的策略，用于改善肝脏健康[17]。
基于现有研究，NMN和NR在调节机体新陈代谢及维持肠道健康方面具有广阔的应用潜力。补充NAD前体如NMN或NR可能通过调节肠道菌群的结构及改变代谢物的组成，从而改善动物的生长性能及相关生理功能。然而，目前尚不明确补充NMN与NR对肠道健康的改善效果是否存在差异，2种前体化合物对肠道微生物及代谢产物的影响是否存在差异尚不清楚。因此本研究以C57小鼠为实验对象，探讨NAD前体NMN和NR对小鼠肠道菌群、肠道代谢物及生长性能的影响，分析两者之间的异同，以期为NMN和NR在改善机体健康方面的应用提供理论参考。
1 材料与方法
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1.1 主要试剂和仪器
烟酰胺单核苷酸(NMN)、烟酰胺核苷(NR)均购自上海迈瑞尔生化科技有限公司；苏木精-伊红(hematoxylin-eosin, H&E)，西安依科生物技术有限公司；Soil DNA Kit，Omega Bio-Tek公司；MiSeq Reagent Kit v3，Illumina公司。
PCR引物由北京擎科生物科技股份有限公司合成，引物序列如表1所示。
跑步机，安徽正华生物仪器设备有限公司；质谱检测器，ThermoFisher Scientific公司。
1.2 试验动物分组及饲养方案
五周龄C57BL/6J雄性小鼠购自重庆腾鑫生物技术有限公司西安分公司，本研究所有动物实验获得西北农林科技大学动物伦理委员会批准，编号为IACUC2025-0309。
45只小鼠，随机分为3组，每组15只，分别为对照组(CON)、添加烟酰胺单核苷酸组(NMN)与添加烟酰胺核苷组(NR)。每组分为
5笼，每笼3只小鼠。适应性饲养1周后正式开始试验。试验组NR与NMN的添加量均为400 mg/(kg·d)[18]，溶剂为饮用水，每日固定时间进行灌胃，1周5次，对照组灌胃等量的饮用水。所有小鼠均饲养在西北农林科技大学实验动物中心(SPF级)，小鼠自由采食和饮水，温度条件为(25±2) °C，湿度为50%-60%。
1.3 生长性能测定
试验期间每周记录小鼠的体重及采食量。饲喂8周后，采用握力测试和跑步实验测试小鼠的运动耐力。握力测试：将小鼠置于铁丝网上，小鼠抓牢铁丝网后倾斜铁丝网并与地面垂直，测定小鼠跌落的时间。跑步实验：实验前小鼠在跑步机上以10 m/min的速度奔跑10 min，进行为期3 d的适应性训练。正式测试设置参数：电刺激强度0.2 mA，力竭标准为20次/30 s，起始速度由10 m/min缓慢增至20 m/min，直至小鼠力竭。
1.4 染色和形态学分析
麻醉后，通过颈椎脱臼实施安乐死。沿腹中线开腹，定位并剪取十二指肠、回肠和结肠，用预冷PBS冲洗。部分组织用4%多聚甲醛固定，其余组织液氮速冻后存于-80 ℃冰箱。组织经4%多聚甲醛固定24 h后，用PBS冲洗，脱水、包埋制作石蜡组织切片，进行H&E染色处理。对结肠组织切片进行阿利辛蓝(alcian)染色处理。在光学显微镜下观察小肠绒毛高度和隐窝深度以及结肠黏膜杯状细胞数量，并通过ImageJ软件进行定量。对肌肉和脂肪组织进行称重统计与形态对比。
1.5 小鼠结肠组织炎症因子测定
取适量结肠样本，Trlzol法提取组织RNA，测量提取的RNA浓度后将其反转录为cDNA，使用炎症因子引物进行定量PCR检测，检测样本中的白细胞介素-1β (interleukin-1β, IL-1β)、白细胞介素6 (interleukin 6, IL-6)、肿瘤坏死因子(tumor necrosis factor-α, TNF-α)和核因子-κB通路(nuclear factor kappa-light-chain-enhancer of activated B cells, NF-κB)，引物序列见表1。RT-qPCR反应体系按照SYBR Primerx Ex TaqTM II 说明书配制。将配制好的试剂经离心后加入384孔板，置于实时荧光定量PCR仪中检测。扩增程序：95 °C预变性30 s；95 °C变性5 s，60 °C退火延伸30 s，进行40个循环。采用2-ΔΔCt法计算基因的相对表达水平，以β-actin作为内参基因。
1.6 16S rRNA基因高通量测序分析
分离结肠并在两端结扎，剪取整段结肠置于无菌培养皿中。纵向剪开结肠，用无菌刮勺收集内容物，放入无菌离心管中。每笼随机选取1只小鼠即每组随机选取5只小鼠的结肠内容物送至苏州帕诺米克生物医药科技有限公司进行高通量测序分析，使用Soil DNA Kit 提取总基因组DNA样品，经质检后于-20 ℃保存。对肠道微生物16S rRNA基因V3-V4区域进行PCR扩增，引物为338F (5′-ACTCCTACGGGA GGCAGCA-3′)和806R (5′-GGACTACHVGGGT WTCTAAT-3′)，扩增产物纯化后进行定量。使用MiSeq Reagent Kit v3在Illumina MiSeq平台进行高通量测序。测序原始数据已上传至NCBI SRA数据库，编号为PRJNA1230388。对原始数据进行序列分析与扩增子序列变体(amplicon sequence variants, ASVs)对齐、分类等，使用Chao1指数和Shannon指数等α多样性指数来评估结肠微生物群的均匀性；非度量多维量表(nonmetric multidimensional scaling, NMDS)用于评估β多样性。线性判别分析(linear discriminant analysis, LDA)和效应大小测量使用LDA effect size (LEfSe)方法，以确定3组之间物种丰度谱的差异。
1.7 非靶向代谢组学分析
结肠内容物样品同1.6节，提取代谢物进行非靶向代谢组学检测(诺米代谢)。使用超高效液相系统和质谱检测器进行上机检测，在正离子和负离子模式下对所有样品进行分析。根据VIP>1和P<0.05 (Student’s t检验)确定显著性。使用MetaboAnalyst (v6.0)对差异代谢物进行通路分析。
1.8 统计分析
使用GraphPad Prism 9.0进行数据分析与绘制，结果以平均值±标准误(SEM)表示，各组之间的差异使用t检验或单因素方差分析，在P<0.05时，结果被认为具有统计学意义。
16S rRNA基因测序结果使用R软件(v2.15.3)进行α、β多样性指数组间差异分析。使用R软件(v2.15.3)对各分类水平下的组间差异物种进行
t检验。*表示P<0.05；**表示P<0.01；***表示P<0.001；****表示P<0.000 1；ns表示差异无统计学意义。差异肠道微生物和其差异代谢物的关联分析用R软件进行Spearman相关性分析，|r|>0.8且P<0.05认为有显著性意义，结果以热图展示。
2 结果与分析
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2.1 饲喂NMN和NR对小鼠生长性能的影响
饲喂NMN和NR对小鼠生长性能的影响结果如图1所示。结果显示，饲喂NMN和NR对小鼠体重增长无显著影响(图1A)。与对照组相比，NMN和NR组小鼠的采食量(图1B)、肌肉和脂肪质量(图1C)均无显著差异。对小鼠运动能力的分析表明，饲喂NMN和NR对小鼠跑步距离(图1D)和跑步时间(图1E)与对照组相比均无显著影响，而饲喂NR可显著增加小鼠的抓握能力(图1F)。
2.2 饲喂NMN和NR对小鼠肠道形态的影响
小鼠回肠组织H&E染色结果如图2A所示。测量小肠绒毛高度(图2C)和隐窝深度(图2D)，并计算肠绒毛高度与隐窝深度的比值(图2E)，结果显示3组之间无显著差异，表明饲喂NMN和NR对肠道吸收功能无显著影响。
对结肠组织进行阿利新蓝染色并进行杯状细胞计数统计，结果显示，与对照组相比添加NMN饲喂的小鼠结肠组织杯状细胞数量显著增多(图2B、2F)。由于消化道中的杯状细胞分泌黏蛋白，在消化道内与水、无机盐、抗菌肽等共同组成肠道黏液屏障，因此饲喂NMN可增强小鼠结肠黏膜屏障功能。
2.3 饲喂NMN和NR对小鼠肠道炎症指标的影响
利用RT-qPCR检测饲喂NMN和NR对肠道炎症因子基因表达水平的影响。结果显示，饲喂NMN和NR均可以显著下调肠道内促炎因子TNF-α、IL-6的表达(P<0.05) (图3)。上述结果表明，饲喂NMN和NR均可抑制促炎因子的表达，降低肠道炎症的发生，两者在抗炎功能方面无显著差异。
2.4 饲喂NMN和NR对小鼠肠道微生物多样性的影响
通过α多样性指标分析肠道微生物群多样性的变化。Chao1指数和observed_species指数反映了肠道微生物物种的丰富度，Shannon指数和Simpson指数则反映了肠道微生物群落的多样性。与对照组相比，饲喂NMN组小鼠肠道微生物的Chao1、observed_species、Shannon、Simpson指数均显著升高(图4A)，而饲喂NR组小鼠肠道微生物的Chao1、observed_species、Shannon指数显著升高(图4B)，表明饲喂NMN或NR均显著提高了肠道微生物的物种丰富度和群落多样性。进一步比较NMN组与NR组小鼠肠道微生物多样性发现，饲喂NMN对肠道微生物群的多样性影响更大(图4C)。
2.5 饲喂NMN和NR对肠道微生物物种组成结构的影响
β多样性用于反映样本间微生物群落的差异，结果显示对照组与NMN组和NR组间样本距离较远，表明样本群落的物种组成存在显著差异(图5A、5B)，说明饲喂NMN或NR均可调节肠道微生物群的组成。
在门水平上，相对丰度大于1%的有7种菌门，其中芽孢杆菌门(Bacillota)和拟杆菌门(Bacteroidota)是小鼠的主要优势菌群(图5C、5D)。由于芽孢杆菌门和拟杆菌门的比值 (Bacillota/Bacteroidota)可作为肠道菌群失调的相关标志物，因此对3组的芽孢杆菌门和拟杆菌门的物种组成进行了数量统计与比值计算。相比对照组，饲喂NMN或NR均降低了肠道芽孢杆菌门和拟杆菌门的比值，但差异并不显著(图5E、5F)，表明添加NMN或NR在不影响肠道菌群失调的条件下改变了肠道芽孢杆菌门和拟杆菌门的比例。
为了进一步比较肠道微生物群的具体差异，对肠道微生物在不同分类水平下构成进行了统计比较。在门水平上，饲喂NMN或NR均显著提高了拟杆菌门(Bacteroidota)、假单胞菌门(Pseudomonadota)的丰度，降低了放线菌门(Actinomycetota)的丰度(P<0.05)，其中饲喂NR组相较于饲喂NMN组，假单胞菌门的丰度也显著提高(图5G)。在属水平上，饲喂NMN显著升高了肠道文肯菌属(Rikenella)、阿德勒克洛伊次菌属(Adlercreutzia)的丰度(P<0.05)；饲喂NR显著升高了肠道文肯菌属(Rikenella)、阿德勒克洛伊次菌属(Adlercreutzia)、瘤胃球菌属(Ruminococcus)的丰度，降低了肠道双歧杆菌属(Bifidobacterium)的丰度(P<0.05)，且NMN组和NR组在阿德勒克洛伊次菌属和瘤胃球菌属的表达上也存在差异(图5H、5I、5J)。
2.6 饲喂NMN和NR对小鼠肠道代谢物的影响
肠内容物的非靶向代谢组测定结果如图6所示。通过正交-偏最小二乘判别分析(orthogonal projections to latent structures discriminant analysis, OPLS-DA)评分图分析，观察到对照组与NMN组和NR组之间存在明显的分离(图6A、6B)，表明各组之间肠道代谢特征存在显著差异。差异代谢物分析结果显示，对照组与NMN组和NR组均有较多差异代谢物。与对照组相比，饲喂NMN下调了胍基琥珀酸、西利替尼、苯乙酸酯，显著上调了脂肪酸类代谢物与核酸类代谢物，如油酸、脱氧胆酸钠、亚油酸、16-氧代棕榈酸酯、13,16,19-二十二碳三烯酸、(2E)-癸烯 酰-ACP、壬酸等(图6C)。与对照组相比，饲喂NR上调了脂肪酸、氨基酸及核苷类代谢物，如花生酸、2-脱氢泛酸、水杨酸、5′-脱氢腺苷、培哚普利、氟伏沙明、2′-脱氧鸟苷、L-天冬酰胺，同时也下调了对苯二酚、脱氢表雄酮、2-焦儿茶酸、L-苹果酸等(图6D)。京都基因与基因组百科全书(Kyoto encyclopedia of genes and genomes, KEGG)富集通路分析表明，添加NR可激活机体精氨酸生物合成通路(图6F)。综上所述，补充NMN和NR能够显著调节肠道代谢物的水平，从而改善机体的脂肪和蛋白质合成及其代谢过程。
2.7 肠道微生物与代谢物关联分析
对16S rRNA基因测序与代谢物分析结果进行关联分析，结果表明NMN组中有7种差异微生物与多种代谢物强相关，其中副拟杆菌属(Parabacteroides)与5′-脱氢腺苷(5′- dehydroadenosine)正相关，与苯乙酸酯(phenyl acetate)负相关(图7A)。NR组中有3种差异微生物与多种代谢物强相关，其中瘤胃球菌属(Ruminococcus)与水杨尿酸(salicyluric acid)、5′-脱氢腺苷(5′-dehydroadenosine)、培哚普利(perindopril)、2′-脱氧鸟苷(2′-deoxyguanosine)正相关，与瓜氨酸(citrulline)负相关(图7B)。这些结果表明，肠道内微生物种类和丰度的变化与其代谢物密切相关。
3 讨论与结论
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肠上皮细胞作为先天免疫系统的一部分，通过分泌黏液将肠腔内容物(尤其是细菌)与上皮细胞分离，避免其直接接触，是肠道屏障的重要组成部分[19]。结肠杯状细胞是特殊的肠上皮细胞，主要功能是通过分泌黏液和黏蛋白在宿主与肠道微生物群之间形成物理隔离，形成肠道屏障，防止细菌入侵和炎症的发生[20]。肠道屏障的破坏会导致细菌渗透到内黏液层，造成肠道感染和炎症性肠病[21]。杯状细胞分泌的黏液在维持大肠生理功能方面起着关键作用，且是预防结直肠疾病和实验性结肠炎的重要因素[22]。本研究的阿利新蓝染色结果显示，饲喂NMN的小鼠结肠杯状细胞数量显著高于对照组，杯状细胞增多，意味着饲喂NMN可以增强小鼠的结肠黏膜屏障，可能对结肠炎的预防和治疗有益。
肠道微生物群是一个由病毒、真菌和肠道细菌组成的复杂多样的群落，它们栖息在肠道中，参与食物消化和内部环境维持[23]。肠道微生物的组成和多样性与机体健康密切相关，肠道微生物群在调节机体免疫功能、抑制病原微生物入侵、参与机体代谢活动等方面发挥重要作用。肠道共生菌与免疫系统之间存在着复杂的微生物-微生物和微生物-宿主的相互作用网络，使得机体免疫力逐渐成熟，也将肠道微生物群的结构塑造得更加稳定[24]。研究表明更高的多样性可以使微生物系统更稳定，而且物种的丰富度增加会加强微生物群的功能。肠道微生物群在维持免疫和生理平衡方面起着重要作用，许多疾病(例如炎症性肠病)患者通常伴有肠道菌群紊乱，其主要特征就是肠道微生物群的丰度和多样性减少[25]。本研究对样品进行16S rRNA基因测序，α多样性分析结果显示，相较于对照组饲喂NMN和NR均会增加肠道微生物群的丰度和多样性(Chao1、Shannon等)，其中NMN处理组表现出更高的物种多样性，肠道微生物更丰富的物种组成有助于更好地增加肠道机能，维持肠道内部稳态；β多样性结果分析显示，对照组和NMN、NR组的样本间群落差异显著，表明两组间生物多样性差异明显，肠道微生物组成差异更大。综合上述结果可知，NMN和NR均会增强肠道微生物的丰度和多样性，调节肠道微生物的组成，进而增强肠道微生物群落的稳定性。
哺乳动物的肠道微生物群主要由芽孢杆菌门(Bacillota)和拟杆菌门(Bacteroidota)这2个优势菌门组成，占总菌群的90%以上[1]。通常用芽孢杆菌门与拟杆菌门的比率来表征这2个优势菌门之间的相对关系，该比率与多种胃肠道、中枢神经系统以及宿主能量代谢病的病理状况有关[26]。本研究结果显示，饲喂NMN或NR在未引起菌群失调的条件下可以下调芽孢杆菌门与拟杆菌门的比率(图5E、5F)。有研究指出，芽孢杆菌门相较于拟杆菌门更有效地从食物中获取能量，从而促进卡路里的吸收及随后的体重增加[27]，芽孢杆菌门与拟杆菌门比值的降低表明机体对炎症的抵抗能力增强，这一点在对结肠组织炎症因子表达的检测(图3)中得到了支持。对肠道微生物门水平进行可视化分析，结果显示饲喂NMN或NR后小鼠肠道拟杆菌门的微生物数量均显著高于对照组，拟杆菌门作为肠道的重要菌群，在机体营养吸收、免疫调节、维持稳态、预防骨质疏松、神经发育等方面起到不可或缺的作用[28]。
肠道微生物群在肠道内进行着复杂活跃的代谢活动，不仅为自身提供了生长和繁殖所需的能量和营养，也产生了大量代谢产物进入机体[29]。研究表明，人体血液中有约36%的小分子物质是由微生物代谢产生或修饰的[30]。对肠道代谢物进行差异性分析可以得出，饲喂NMN或NR均增加了有益代谢物的产生，如脂肪酸、有机酸类、氨基酸代谢物等。脂肪酸等脂质代谢物可以调控脂质代谢、促进细胞膜稳定，部分具有抗炎作用，具有降低结肠pH、抑制病原繁殖、抗糖尿病等作用[31]；胆汁酸类代谢物可以抑制有害菌生长、维持肠黏膜完整性等[29]；黄酮类化合物具有抗氧化、抗肿瘤、抗炎等作用。
通过整合分析发现，NMN/NR可能通过微生物-代谢轴来影响宿主健康。饲喂NMN可以增加肠道拟杆菌属的丰度，拟杆菌属(Bacteroides)的增殖与油酸等脂肪酸代谢物水平正相关，后者已知通过激活PPARγ促进杯状细胞分化[20]，这与本研究中杯状细胞数量增加(图2F)和炎症因子下降(图3)的表型一致。饲喂NR则增加肠道瘤胃球菌属(Ruminococcus)的丰度，瘤胃球菌与5′-脱氢腺苷等核苷代谢物相关，这些物质可通过抑制NF-κB通路[15]解释观察到的抗炎效应(图3)，增强系统免疫功能。
饲喂NMN和NR均通过改变肠道微生物群而影响代谢物的组成，尤其在脂肪酸、氨基酸及核苷类代谢物的富集上，进而参与机体的多项生理功能。饲喂NMN可通过副拟杆菌(Parabacteroides)降低乙酸苯酯(phenyl acetate)的表达水平。副拟杆菌与宿主健康密切相关，涉及代谢综合征、炎症性肠病以及肥胖症等多个方面[32]，乙酸苯酯作为芳香族化合物的代谢产物，属于苯丙氨酸代谢途径的衍生物，其水平升高常被视为代谢紊乱(如苯丙酮尿症)的标志[33]。因此NMN通过调节肠道微生物和代谢物的表达，有助于增强机体的稳态。饲喂NR组的分析结果表明，瘤胃球菌(Ruminococcus)可能通过促进某些代谢物的合成来影响宿主的代谢状态，尤其是与能量代谢、抗炎反应和细胞功能相关的代谢物，如水杨尿酸(salicylic acid)、5′-脱氢腺苷(5′-dehydroadenosine)、2′-脱氧鸟苷 (2′-deoxyguanosine)等。瘤胃球菌与各种肠内和肠外疾病有关，并在炎症性肠病和代谢疾病中发挥重要作用[34]。目前尚未明确瘤胃球菌在疾病发展中的具体作用，这一研究结果可能为瘤胃球菌在促进宿主健康方面的有益作用提供了初步证据。
NR是NAD+前体，提升NAD+水平可改善线粒体功能，增强肌肉能量代谢(对快速收缩肌纤维尤其重要)，从而增强肌肉耐力和力量[6]。在本研究中，饲喂NR组的抓握能力显著提升，与观测到的代谢物之间的变化存在潜在关联。与对照组相比，饲喂NR上调了脂肪酸代谢(花生酸)、氨基酸代谢物(L-天冬酰胺、2-脱氢泛酸)和水杨酸等代谢物，花生酸(二十烷酸)是细胞膜磷脂成分，其上调可能反映膜修复或能量储存需求增加，支持肌肉活动[35]；L-天冬酰胺参与三羧酸循环(tricarboxylic acid cycle, TCA cycle)中间代谢，可能优化能量供应；2-脱氢泛酸(维生素B5衍生物)是辅酶A前体，促进脂肪酸氧化和乙酰CoA生成；水杨酸的抗炎作用可能减轻运动诱导的肌肉损伤。饲喂NR下调了对苯二酚和苹果酸，对苯二酚是氧化应激标志物，下调可能反映NR的抗氧化作用，减少运动导致的活性氧积累；苹果酸是TCA循环中间体，下调可能表明TCA循环通量增加，苹果酸被快速消耗以支持ATP生成。对饲喂NR组的KEGG富集分析结果表明(图6F)，TCA循环、丙酮酸代谢、丙氨酸、天冬氨酸和谷氨酸代谢等核心代谢通路以及牛磺酸和亚牛磺酸代谢等氧化应激通路显著富集，且与代谢物变化高度关联。基于以上结论，本研究推测饲喂NR可能通过促进TCA循环、调控能量代谢(脂肪酸、氨基酸途径)以及维持氧化还原平衡(下调对苯二酚、上调水杨酸，协同减少氧化损伤)来优化神经肌肉功能，从而增强小鼠抓握能力。
对上述结果进行汇总与分析，并对NMN与NR的作用差异进行比较，可以得出以下结论：饲喂NMN在增加肠黏膜杯状细胞数量(图2)和提升肠道微生物多样性方面优于饲喂NR (图4)；而饲喂NR在增强小鼠抓握能力方面优于饲喂NMN组(图1)。这表明NMN通过Slc12a8介导的肠道直接吸收[36]，在肠道中高效转化为NAD+，激活SIRT1依赖的黏液分泌通路[7]，从而在改善肠道微生态和屏障功能方面优于NR。相比之下，NR需要依赖NRK1/2进行转化，其效应可能更多通过系统性NAD⁺水平的提升来实现[18]，NR的全身分布特性使其在肌肉功能调节方面表现更佳。结合代谢组学结果分析，上述差异可能与两者的代谢途径有关：NMN通过“拟杆菌-脂肪酸”轴增强局部免疫(图7A)，而NR则通过“瘤胃球菌-精氨酸”轴影响系统代谢(图7B)，这与两者的药代动力学特性一致。此外，NMN上调的代谢物(如油酸)与线粒体功能增强相关，而NR更倾向于调节氨基酸代谢(图6E)，提示两者靶向的细胞通路存在差异。未来需要通过同位素标记或组织特异性敲除模型进一步验证这些差异。若扩展到实际应用中，则需根据治疗目标选择合适的NAD⁺前体：对于肠道疾病，NMN可能是更好的选择；而对于代谢相关疾病，NR可能更具优势。
综上所述，在小鼠饲养过程中补充NMN或NR均可以调节肠道微生物群的组成结构，增加益生菌的丰度，减少与疾病相关的菌群丰度，其中NMN对肠道微生物的影响更为显著。同时，NMN和NR均可通过改变肠道微生物群来调节机体代谢，增加粪便中有益代谢物的浓度，并与代谢物相互作用发挥健康效应，尤其在抗炎和抗氧化方面表现出积极作用。此外，补充NR可以增强小鼠的抓握能力，而补充NMN则可以维持肠上皮的完整性，增加结肠杯状细胞的数量，从而加强肠道屏障功能。
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  • 中央高校基本科研业务费专项资金(Z1090223017)
  • 陕西省秦创原引用高层次创新创业人才项目(QCYRCXM-2022-212)
  • 陕西省畜禽育种“两链”融合重点项目(2022GD-TSLD-46)
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2025年第65卷第9期
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doi: 10.13343/j.cnki.wsxb.20250174
  • 接收时间：2025-03-04
  • 首发时间：2026-02-07
  • 出版时间：2025-09-04
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  • 收稿日期：2025-03-04
  • 录用日期：2025-04-08
基金
Fundamental Research Funds for the Central Universities(Z1090223017)
中央高校基本科研业务费专项资金(Z1090223017)
Qin Chuang Yuan Innovation and Entrepreneurship Talent Project(QCYRCXM-2022-212)
陕西省秦创原引用高层次创新创业人才项目(QCYRCXM-2022-212)
Shaanxi Livestock and Poultry Breeding Double-chain Fusion Key Project(2022GD-TSLD-46)
陕西省畜禽育种“两链”融合重点项目(2022GD-TSLD-46)
作者信息
    西北农林科技大学 动物科技学院，陕西省动物遗传育种与繁殖重点实验室，陕西 杨凌
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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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