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Article(id=1322934620320313362, tenantId=1146029695717560320, journalId=1319373162502258839, issueId=1322934619896688658, articleNumber=null, orderNo=null, doi=10.16016/j.2097-0927.202501018, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1735660800000, receivedDateStr=2025-01-01, revisedDate=1737302400000, revisedDateStr=2025-01-20, acceptedDate=null, acceptedDateStr=null, onlineDate=1793341792421, onlineDateStr=2026-10-30, pubDate=1740672000000, pubDateStr=2025-02-28, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1793341792421, onlineIssueDateStr=2026-10-30, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1793341792421, creator=13701087609, updateTime=1793341792421, updator=13701087609, issue=Issue{id=1322934619896688658, tenantId=1146029695717560320, journalId=1319373162502258839, year='2025', volume='47', issue='4', pageStart='285', pageEnd='384', issueExtLink='null', onlineDate='null', pubDate='1740672000000', pubDateStr='2025-02-28', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1793341792320, creator='13701087609', updateTime=1793341792320, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext=null, issueFiles=null, downloadFileDto=null}, startPage=285, endPage=289, ext={EN=ArticleExt(id=1322934620517445651, articleId=1322934620320313362, tenantId=1146029695717560320, journalId=1319373162502258839, language=EN, title=Circadian Metabolism: What's new in circadian regulation of metabolism, columnId=1320434417353470541, journalTitle=Journal of Army Medical University, columnName=Expert Review, runingTitle=null, highlight=null, articleAbstract=

Metabolic diseases, including obesity and diabetes, etc., are a spectrum of life-threatening diseases, in which metabolic dysfunction is a hallmark. It is in urgent need of new insights for the prevention and treatment of the diseases. Circadian rhythm is fundamental in health and wellbeing. Research on circadian rhythm has shed light on metabolic homeostasis, thereby giving birth to a new direction, named circadian metabolism. Here, we summarize recent advances in circadian metabolism regarding organelle regulation, inter- and intra-organ crosstalk of circadian clocks, as well as molecular mechanisms of time-restricted eating, and propose circadian communication across the hierarchy of central and peripheral clocks as a key concept for understanding the temporal regulation of metabolism. The underlying mechanisms on circadian communication should shed light on precision medicine in a temporal resolution and provide guidelines on when to eat and when to exercise for the prevention of metabolic diseases.

, authors=Mindian LI1, Jianxin ZHANG1, 2, authorsList=Mindian LI, Jianxin ZHANG, authorCompany=null, correspAuthors=Mindian LI, authorNote=null, correspAuthorsNote=null, copyrightStatement=© 2025 Journal of Army Medical University, copyrightOwner=Editorial Office of Journal of Army Medical University, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=null), CN=ArticleExt(id=1322934621234671639, articleId=1322934620320313362, tenantId=1146029695717560320, journalId=1319373162502258839, language=CN, title=代谢生物钟：昼夜节律调控代谢研究的最新进展, columnId=1320434417496076879, journalTitle=陆军军医大学学报, columnName=专家述评, runingTitle=null, highlight=null, articleAbstract=

‍代谢性疾病是以糖脂代谢紊乱为病理生理学特征的慢性疾病，包括肥胖、糖尿病等，对人体健康构成严重威胁，亟需理论创新以指导防治窗口前移。昼夜节律作为健康机体的关键生理特性，从其调控角度出发防治代谢性疾病，已成为代谢生理学的新方向——代谢生物钟。近年来，国内外学者推动昼夜节律研究，在细胞内机制、跨器官机制及生活方式干预等方面取得显著进展。本文聚焦于细胞器调节、器官间与细胞间生物钟通讯以及限时进食的分子机制，阐述代谢生物钟领域的最新成果，并提出中枢神经与外周器官、外周器官间生物钟的相互作用是解析生物钟调控代谢稳态机制的核心科学问题。攻克这一难题，将为实现代谢稳态的精准时间干预提供关键靶点，为指导合理进食与运动以防治代谢性疾病提供全新理论依据。

, authors=李旻典1, 张舰心1, 2, authorsList=李旻典, 张舰心, authorCompany=null, correspAuthors=李旻典, authorNote=null, correspAuthorsNote=
LI Mindian，E-mail：
李旻典，博士，研究员，博士研究生导师，国家海外高层次引进青年人才，国家科技重大专项青年首席科学家，重庆市学术技术带头人。担任陆军军医大学第一附属医院心血管内科副主任，老年心脑血管病教育部重点实验室主任助理，创伤与化学中毒全国重点实验室PI。兼任中国细胞生物学学会生物节律分会秘书长、中国生物化学与分子生物学会代谢专业分会副秘书长、中国生物物理学会代谢生物学分会青年委员。获得基金委重大研究计划培育项目、面上项目、重庆市杰出青年科学基金等项目资助。从事生理学研究，聚焦生物钟与代谢，旨在精准调节人体生物钟实现主动健康，在Cell Metabolism、Nature Metabolism等期刊发表学术论文30余篇。E-mail：
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2联勤保障部队第九六〇医院心血管内科，山东 济南, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null)}, companyList=[AuthorCompany(id=1322934621448581145, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, xref=1, ext=[AuthorCompanyExt(id=1322934621461164058, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, companyId=1322934621448581145, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1Department of Cardiovascular Diseases，Center for Circadian Metabolism and Cardiovascular Disease，Key Laboratory of Geriatric Cardiovascular and Cerebrovascular Diseases of Ministry of Education，Southwest Hospital，Army Medical University （Third Military Medical University），Chongqing), AuthorCompanyExt(id=1322934621469552667, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, companyId=1322934621448581145, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1陆军军医大学第一附属医院心血管内科，代谢生物钟与心血管疾病研究中心，老年心脑血管病教育部重点实验室，重庆)]), AuthorCompany(id=1322934621540855836, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, xref=2, ext=[AuthorCompanyExt(id=1322934621549244445, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, companyId=1322934621540855836, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2Department of Cardiovascular Diseases，The 960th Hospital of the PLA Joint Service Support Force，Jinan，Shandong，China), AuthorCompanyExt(id=1322934621557633054, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, companyId=1322934621540855836, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2联勤保障部队第九六〇医院心血管内科，山东 济南)])])], keywords=[Keyword(id=1322934623776419883, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, language=EN, orderNo=1, keyword=circadian rhythm), Keyword(id=1322934623843528748, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, language=EN, orderNo=2, keyword=circadian clock), Keyword(id=1322934623906443309, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, language=EN, orderNo=3, keyword=metabolism), Keyword(id=1322934623981940782, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, language=EN, orderNo=4, keyword=metabolic diseases), Keyword(id=1322934624057438256, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, language=CN, orderNo=1, keyword=昼夜节律), Keyword(id=1322934624158101553, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, language=CN, orderNo=2, keyword=生物钟), Keyword(id=1322934624229404722, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, language=CN, orderNo=3, keyword=糖脂代谢), Keyword(id=1322934624300707891, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, language=CN, orderNo=4, keyword=代谢性疾病)], refs=[Reference(id=1322934624703361079, tenantId=1146029695717560320, journalId=1319373162502258839, articleId=1322934620320313362, doi=10.1126/science.abd0951, pmid=null, pmcid=null, year=2021, volume=371, issue=6530, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=1, rfOrder=0, authorNames=KORONOWSKI K B, SASSONE-CORSI P, journalName=Science, refType=null, unstructuredReference=KORONOWSKI K B，SASSONE-CORSI P. 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A：细胞生物钟调控新模式；B：器官间生物钟相互作用；C：细胞间生物钟相互作用；D：营养节律调控代谢

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代谢生物钟：昼夜节律调控代谢研究的最新进展
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李旻典 1 , 张舰心 1, 2
陆军军医大学学报 | 专家述评 2025,47(4): 285-289
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陆军军医大学学报 |专家述评 2025 , 47 (4) : 285 -289
代谢生物钟：昼夜节律调控代谢研究的最新进展
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李旻典1 , 张舰心1, 2
作者信息
  • 1陆军军医大学第一附属医院心血管内科，代谢生物钟与心血管疾病研究中心，老年心脑血管病教育部重点实验室，重庆
  • 2联勤保障部队第九六〇医院心血管内科，山东 济南
通讯作者:
LI Mindian，E-mail：
李旻典，博士，研究员，博士研究生导师，国家海外高层次引进青年人才，国家科技重大专项青年首席科学家，重庆市学术技术带头人。担任陆军军医大学第一附属医院心血管内科副主任，老年心脑血管病教育部重点实验室主任助理，创伤与化学中毒全国重点实验室PI。兼任中国细胞生物学学会生物节律分会秘书长、中国生物化学与分子生物学会代谢专业分会副秘书长、中国生物物理学会代谢生物学分会青年委员。获得基金委重大研究计划培育项目、面上项目、重庆市杰出青年科学基金等项目资助。从事生理学研究，聚焦生物钟与代谢，旨在精准调节人体生物钟实现主动健康，在Cell Metabolism、Nature Metabolism等期刊发表学术论文30余篇。E-mail：
Circadian Metabolism: What's new in circadian regulation of metabolism
Mindian LI1 , Jianxin ZHANG1, 2
Affiliations
  • 1Department of Cardiovascular Diseases，Center for Circadian Metabolism and Cardiovascular Disease，Key Laboratory of Geriatric Cardiovascular and Cerebrovascular Diseases of Ministry of Education，Southwest Hospital，Army Medical University （Third Military Medical University），Chongqing
  • 2Department of Cardiovascular Diseases，The 960th Hospital of the PLA Joint Service Support Force，Jinan，Shandong，China
出版时间: 2025-02-28 doi: 10.16016/j.2097-0927.202501018
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摘要
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‍代谢性疾病是以糖脂代谢紊乱为病理生理学特征的慢性疾病，包括肥胖、糖尿病等，对人体健康构成严重威胁，亟需理论创新以指导防治窗口前移。昼夜节律作为健康机体的关键生理特性，从其调控角度出发防治代谢性疾病，已成为代谢生理学的新方向——代谢生物钟。近年来，国内外学者推动昼夜节律研究，在细胞内机制、跨器官机制及生活方式干预等方面取得显著进展。本文聚焦于细胞器调节、器官间与细胞间生物钟通讯以及限时进食的分子机制，阐述代谢生物钟领域的最新成果，并提出中枢神经与外周器官、外周器官间生物钟的相互作用是解析生物钟调控代谢稳态机制的核心科学问题。攻克这一难题，将为实现代谢稳态的精准时间干预提供关键靶点，为指导合理进食与运动以防治代谢性疾病提供全新理论依据。

昼夜节律  /  生物钟  /  糖脂代谢  /  代谢性疾病

Metabolic diseases, including obesity and diabetes, etc., are a spectrum of life-threatening diseases, in which metabolic dysfunction is a hallmark. It is in urgent need of new insights for the prevention and treatment of the diseases. Circadian rhythm is fundamental in health and wellbeing. Research on circadian rhythm has shed light on metabolic homeostasis, thereby giving birth to a new direction, named circadian metabolism. Here, we summarize recent advances in circadian metabolism regarding organelle regulation, inter- and intra-organ crosstalk of circadian clocks, as well as molecular mechanisms of time-restricted eating, and propose circadian communication across the hierarchy of central and peripheral clocks as a key concept for understanding the temporal regulation of metabolism. The underlying mechanisms on circadian communication should shed light on precision medicine in a temporal resolution and provide guidelines on when to eat and when to exercise for the prevention of metabolic diseases.

circadian rhythm  /  circadian clock  /  metabolism  /  metabolic diseases
李旻典, 张舰心. 代谢生物钟：昼夜节律调控代谢研究的最新进展. 陆军军医大学学报, 2025 , 47 (4) : 285 -289 . DOI: 10.16016/j.2097-0927.202501018
Mindian LI, Jianxin ZHANG. Circadian Metabolism: What's new in circadian regulation of metabolism[J]. Journal of Army Medical University, 2025 , 47 (4) : 285 -289 . DOI: 10.16016/j.2097-0927.202501018
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代谢性疾病是以糖脂代谢紊乱为特征的慢性疾病，包括肥胖、糖尿病、代谢性脂肪肝病、高脂血症等，对人体健康造成重大危害。节食、运动等生活方式干预，以及结合药物治疗、减重与代谢手术，在防治代谢性疾病方面发挥重要作用。然而，代谢性疾病的总体防控压力仍然很大；领域存在知晓率低、诊断率低、控制率低的难题，亟需防治窗口前移。昼夜节律是健康机体的重要生理特征，解析昼夜节律分子机制的研究成果获得2017年诺贝尔医学奖（https://www.nobelprize.org/prizes/medicine/2017/press-release/）；从昼夜节律调控的角度防治代谢性疾病需要时间生物学与临床医学跨学科合作解决，由此形成了代谢生理学研究的一个交叉方向——代谢生物钟（circadian metabolism）。代谢生物钟的重要科学问题包括：①机体代谢与外环境（光照、食物等）的昼夜节律相互作用的机制及生理意义；②中枢与外周器官生物钟如何相互作用形成机体代谢昼夜节律；③干预生物钟实现调控代谢稳态的作用机制等。
在绝大部分细胞，由脑和肌肉芳香烃受体核转运样1蛋白（brain and muscle arnt-like 1，BMAL1）和昼夜运动节律输出循环突变蛋白（circadian locomoter output cycles kaput，CLOCK）构成的转录因子复合物驱动下游负反馈基因PER（周期蛋白，period）和CRY（隐花色素，cryptochrome）的表达，形成24 h的周期性自发性昼夜节律，也就是核心生物钟；同时核受体亚家族1组D成员1［nuclear receptor subfamily 1，group D member 1，REV-ERBα（又称NR1D1）］、白细胞介素3调节的核因子（nuclear factor，interleukin 3 regulated，NFIL3）等转录因子基因的表达也受BMAL1和CLOCK的周期性调控，驱动下游代谢、细胞生理的相关基因表达，与核心生物钟一起形成细胞生命活动的昼夜节律［1］。细胞生物钟之间存在相互作用，接收环境神经、体液信号和来自其他生物钟的信号与机体昼夜节律同步；位于下丘脑视交叉上核（suprachiasmatic nucleus，SCN）的中枢生物钟与位于其他神经核团、外周器官组织和细胞的外周生物钟形成层级结构，整合光照和营养信号的昼夜变化，形成机体24 h的昼夜节律［2-3］。流行病学和小样本观察性临床研究提示生物钟紊乱增加代谢性疾病的发病风险；基于小鼠的遗传研究发现生物钟基因突变导致肥胖、糖尿病、脂肪肝等代谢性疾病［2］。近年来昼夜节律研究在细胞内机制、跨器官机制和生活方式干预等方面取得重要进展；揭示了调节生物钟的细胞器、发现器官间和细胞间生物钟通讯机制、深入研究干预器官昼夜节律的饮食方式改善代谢稳态的效应，为基于生物钟理论防治代谢性疾病提供新方向。
1　细胞通过纤毛动态、基因表达的相变输出昼夜节律
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细胞器作为细胞行为、生理和代谢的重要结构，是生物钟发挥功能的重要结构基础。线粒体形态和溶酶体活性都受到生物钟的调节，呈现昼夜节律。纤毛是一种细胞表面突起的由微管构成轴丝的毛发状细胞器；不运动的纤毛称为初级纤毛，分布着分节极性蛋白Hedgehog信号通路，调节细胞生长、增殖分化等过程。研究［4］揭示，SCN的神经调节素S（neuromedin S）神经元存在初级纤毛形态和功能的昼夜节律，这些SCN初级纤毛具有开灯时（对应生理傍晚）数量多、长度长的昼夜节律，维持小鼠生物钟稳定性；机制上，SCN初级纤毛昼夜交替地伸缩，从而周期性激活Hedgehog通路，调节SCN核心生物钟基因和神经递质释放的昼夜节律［4］（图1A）。通过遗传或药物手段干预纤毛形成或Hedgehog通路，导致小鼠SCN神经元的动作电位发放与恢复无法形成昼夜周期，跑轮行为的昼夜节律更容易适应光照周期的改变［4］。
相变或相分离是细胞生物学研究的热点和前沿，在细胞生物学过程发挥独特的无膜细胞器功能，形成细胞内蛋白质机器的区块化，保证专一执行特定功能。研究发现小鼠肝脏细胞核内REV-ERBα的固有无序结构域参与凝聚体组装，凝聚体是由液体相分离形成的液滴，REV-ERBα形成的凝聚体招募其他含无序结构域蛋白如转录共抑制因子NCOR1，维持转录抑制所需的染色质高级结构，从而抑制肝脏生物钟基因和代谢基因表达［5］。RNA结合蛋白ATXN2及其同源蛋白ATXN2L在细胞内发生相分离，其形成的液滴随昼夜节律振荡，在振荡高峰期募集核糖体和特异RNA，促进关键生物钟基因PER2的转录本翻译为蛋白质，精确调控细胞和行为的昼夜节律周期［6］（图1A）。
2　中枢神经与外周生物钟相互作用维持机体代谢稳态
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下丘脑-垂体轴介导的激素分泌与肝脏糖异生反应在黎明达到高峰，形成内分泌与代谢的“黎明现象”。O连接N乙酰氨基葡萄糖转移酶（O-GlcNAc转移酶，O-GlcNAc transferase，OGT）感知葡萄糖波动，调节肝细胞生物钟和胰岛素受体信号通路；干预肝细胞OGT影响血糖和胰岛素敏感性的昼夜波动［7］。研究发现，SCN的γ-氨基丁酸（gamma-aminobutyric acid，GABA）神经元生物钟向肝脏传递昼夜节律信号，调节胰岛素信号通路活性和肝脏糖异生反应的昼夜节律；在下丘脑GABA神经元敲除REV-ERBα及其同源基因REV-ERBβ抑制肝脏胰岛素受体通路活性，影响血糖的昼夜波动［8］。因此，中枢生物钟向肝脏生物钟传递时间信号，调节机体糖代谢的稳态，机制上需要肝脏OGT介导的营养感知机制和胰岛素信号通路。
最新研究［9］发现，机体存在从肝脏生物钟到中枢生物钟的反馈作用。肝脏生物钟经肝脏迷走传入神经反馈到中枢生物钟，影响摄食量的昼夜节律；靶向生物钟基因破坏肝脏生物钟引起摄食量高峰时间从生理活跃期向静息期迁移，造成生理静息期摄食量增加；进一步手术切除肝脏迷走神经可以纠正节律紊乱小鼠的摄食量模式。
中枢生物钟除了与肝脏生物钟相互作用，还可以与骨骼肌生物钟、表皮生物钟相互作用（图1B）。中枢生物钟与骨骼肌生物钟的相互作用调节骨骼肌线粒体代谢稳态，延缓肌肉纤维在衰老过程中的退行性变化［10］。中枢生物钟需要表皮生物钟调节细胞周期，实现皮肤细胞的更替和稳态［11］。外周生物钟之间也存在相互作用。肝脏生物钟可以影响肺脏转录组的昼夜节律，与缺氧应激通路协同影响肺脏的生理稳态；肝脏缺乏BMAL1和缺氧诱导因子1α（hypoxia inducible factor 1 subunit alpha，HIF1α）的小鼠出现低氧血症，表现出肝脏功能障碍、肺血管舒张和低氧血症等肝肺综合征特征［12-13］。
3　实质细胞与微环境生物钟互作维持器官代谢稳态
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同一器官的不同细胞存在生物钟相互作用。肝细胞生物钟的紊乱会影响肝脏巨噬细胞和内皮细胞的昼夜节律［14］；肝细胞特异性的敲除REV-ERB基因造成肝脏巨噬细胞脂代谢昼夜节律的改变。机制上，肝细胞的胆固醇调节元件结合蛋白（sterol-regulatory element binding protein，SREBP）转录因子介导这一从肝细胞生物钟到巨噬细胞的昼夜时间信号通讯轴［15］（图1C）。糖脂代谢紊乱引起慢性低度的代谢性炎症，主要发生在脂肪组织；对于衰老和肥胖小鼠，丙种球蛋白在脂肪组织的异位堆积是代谢性炎症的关键病理生理学特征［16］。研究发现生物钟基因在脂肪组织的一种固有样淋巴细胞高表达；这类T细胞受体由γ链和δ链组成的T细胞合成和分泌IL-17细胞因子的功能具有昼夜节律，调节脂肪细胞脂肪代谢，维持体温和能量代谢的昼夜节律［17］。这些最新研究进展提示，器官内不同细胞的生物钟可能存在层级结构，跨层级的生物钟相互作用形成器官统一的昼夜节律，维持器官细胞和代谢的稳态［1，3，14］。
4　调整进食时间可增强代谢与体能、预防代谢性疾病
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相比于光照对于中枢生物钟的调节作用，调整进食时间引起的营养代谢节律对于肝脏、脂肪组织等器官生物钟的调节作用超过中枢生物钟的影响［1-2］。限时进食通过将进食时间限制在活跃期（对应生理白天）或静息期（对应生理夜晚）可以在7 d内快速调整肝脏生物钟的高峰时间，而不影响中枢生物钟［3］。因此，限时进食的生理学成为研究生物钟调控代谢稳态的重要范式（图1D）。
位于肝脏、脂肪组织、骨骼肌等代谢器官组织的生物钟维持糖脂代谢稳态，这些器官的生物钟紊乱加剧脂肪肝、肥胖、胰岛素抵抗等代谢紊乱［2］。利用肝脏、脂肪等外周生物钟快速应答进食时间改变的特点，不同限制进食时间的生活方式干预方案（即限时进食），如16∶8间歇性禁食、斋月禁食，在预防代谢性疾病具有安全、易行、有效的潜在优势，日益受到医学研究和社会重视［18-19］。9～10 h的活跃期限时进食（类似16∶8间歇性禁食）大幅降低肥胖小鼠的体质量、体脂，改善胰岛素抵抗、代谢性炎症和动脉粥样硬化［20-21］。然而，16∶8间歇性禁食可诱导毛囊干细胞凋亡，从而抑制毛囊的再生能力，造成脱发［22］。16∶8间歇性禁食联合节食在1年左右的观察期内对人体安全，有效降低体脂量［23］。从生物钟调节的角度看，12 h限时进食已经足够引起肝脏和脂肪生物钟的相位和振幅改变，协调外周生物钟与中枢生物钟的运行；然而低于12 h的进食时间会引起能量摄入的降低。因此，9～10 h的活跃期限时进食并不是理论意义上的仅调节生物钟、不影响能量代谢稳态的干预方案，准确的说，这类应用于小鼠的限时进食方案应该被称为“限时节食”（timed calorie restriction）。
与活跃期限时进食的进食时间相反，静息期限时进食（类似斋月禁食）在协调运动相关的生物钟与生理功能上有独特的作用机制。研究发现静息期限时进食增强小鼠运动耐力，通过激活骨骼肌生物钟介导的脂滴表面蛋白5（perilipin 5，PLIN5）的昼夜节律，促进骨骼肌纤维从糖酵解型向线粒体代谢型的转换，提高脂肪酸代谢活性［24］。相比于活跃期限时进食，除非在应急动员或特定习俗情景下，静息期限时进食的普及性不高。然而，基于动物模型，可以通过解析静息期限时进食改善体能耐力的分子和细胞生理机制，找到生物标记物或可干预靶标实现体能增强。
5　小结与展望
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基于这些最新研究，生物钟的理论面临巨大的创新机遇。首先，越来越多的证据支持外周生物钟存在与中枢生物钟的相互作用；不同器官、组织，甚至同一器官的不同细胞存在生物钟相互作用。因此，机体昼夜节律的形成机制不能套用“一个位于中枢SCN的主生物钟，调节除此以外的外周生物钟”的主仆层级结构描述；于是国际上出现了网络结构假说、外周生物钟层级结构假说等竞争性假说［1，3］。
其次，中枢神经与外周生物钟相互作用、外周生物钟之间相互作用的分子机制是指导药物研发和转化应用的关键科学问题。神经调节、体液调节和生物物理因素（体温、机械应力、流体剪切力）的单一要素或组合可能为调节生物钟系统、实现代谢稳态的精准时间干预提供重要靶标，为指导在什么时间进食和运动防治代谢性疾病提供新理论。然而这些正是目前知识的盲点，需要深入系统研究。
最后，小鼠与人类在昼夜行为模式和能量代谢率方面存在显著差异。尽管昼夜节律对所有生物体，包括小鼠和人类，都具有重要的适应性意义，但小鼠的研究结果需要通过基于人群队列的昼夜多组学数据进行验证。在这方面，国际同行已经取得了领先，提供了公共数据库，但来自中国人群的循证证据至今缺乏并至关重要。
面对这些创新机遇和挑战，代谢生物钟研究将利用人工智能辅助的时空多组学技术、高内涵机制研究体系等先进平台，实现理论创新，从时间视角为防治代谢性疾病贡献“中国方案”。

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2025年第47卷第4期
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doi: 10.16016/j.2097-0927.202501018
  • 接收时间：2025-01-01
  • 首发时间：2026-10-30
  • 出版时间：2025-02-28
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  • 收稿日期：2025-01-01
  • 修回日期：2025-01-20
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    1陆军军医大学第一附属医院心血管内科，代谢生物钟与心血管疾病研究中心，老年心脑血管病教育部重点实验室，重庆
    2联勤保障部队第九六〇医院心血管内科，山东 济南

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李旻典，博士，研究员，博士研究生导师，国家海外高层次引进青年人才，国家科技重大专项青年首席科学家，重庆市学术技术带头人。担任陆军军医大学第一附属医院心血管内科副主任，老年心脑血管病教育部重点实验室主任助理，创伤与化学中毒全国重点实验室PI。兼任中国细胞生物学学会生物节律分会秘书长、中国生物化学与分子生物学会代谢专业分会副秘书长、中国生物物理学会代谢生物学分会青年委员。获得基金委重大研究计划培育项目、面上项目、重庆市杰出青年科学基金等项目资助。从事生理学研究，聚焦生物钟与代谢，旨在精准调节人体生物钟实现主动健康，在Cell Metabolism、Nature Metabolism等期刊发表学术论文30余篇。E-mail：
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2种不同金属材料的力学参数
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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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