运动因子对心血管系统保护作用的研究进展

来源期刊：广州医药 | 870-879 发布时间：2025-07-20 收稿时间：2025/8/6 16:51:06 阅读量：55

作者：: 欧阳嘉富,

袁秋菊,

江芷珊,

范玉冰,

冯彦翔,

孙江鸥,

刘东辉,

关键词：: 运动运动因子心血管疾病代谢; exercise exerkines cardiovascular disease metabolism

DOI：: 10. 20223 / j. cnki. 1000-8535. 2025. 07. 002

收稿时间：: 2024-11-22
修订日期：
接收日期：
引用总数：: 0

运动可以调节机体代谢，预防和治疗由糖脂代谢紊乱所引发的心血管疾病。运动因子是在运动过程中，由肌肉、脂肪以及肝脏等多个组织合成和分泌的一系列生物活性物质，包括蛋白质和多肽类分子、小分子代谢物以及核酸等。诸多研究证实，运动因子是运动调节机体代谢的重要因素之一，也是机体从运动中获益的关键分子机制。近年来，随着蛋白质组学、代谢组学以及高通量测序等相关技术的飞速发展，越来越多的运动因子被陆续发现和证实。这不仅拓宽了人们对机体从运动中获益相关机制的认知，还激发了人们对运动因子在健康领域应用前景的浓厚兴趣。文章系统地阐述了运动因子对机体心血管系统的影响，旨在揭示运动因子在促进心血管健康以及治疗心血管疾病等方面的积极效用。

刘东辉 广州市第一人民医院/华南理工大学附属第二医院老年医学科副主任/老年脏器康复中心主任。北京大学医学博士，主任医师，博士生导师，广州市卫生健康委优秀人才，第八届“羊城好医生”。担任中国健康管理协会生活方式医学分会理事，中国老年医学分会智慧医疗技术与管理分会委员，中国老年保健协会脏器康复专业委员会常务委员，中国心脏联盟广东省心脏康复中心联盟常务委员，中国民族卫生协会卫生健康技术推广专家委员会委员，广东省医学会老年医学分会常务委员，广东省医学会高血压分会委员，广州市医养结合指导中心专家组副主任，广州市医学会心血管病学分会常务委员等。担任Metabolism and Translational Medicine 编委，Health and Metabolism 青年编委。

主要研究方向：糖脂代谢、氧化应激、炎症与动脉粥样硬化和血管衰老等。擅长高血压、血脂紊乱、动脉粥样硬化、冠心病、心力衰竭及代谢性心血管疾病的临床诊治，尤其是各类慢性心血管疾病的“五大处方”心脏康复治疗和抗衰老的非药物治疗。

目前主持国家自然科学基金2项，省级课题5项。主持国家卫生健康技术推广项目“心肺运动试验在老年心血管疾病患者中的应用与推广”1项。以第1作者/通讯作者在Signal Transduct Target Ther等杂志发表SCI论文10余篇。参编中国首部《心血管运动医学指南》等指南2部。主编英文论著Statins - From Lipid-Lowering Benefits to Pleiotropic Effects 1部，参编英文论著Apolipoproteins，Triglycerides and Cholesterol 1部。获得计算机软件著作权1项。获得“福建医学科技奖三等奖”1项，获得“厦门市科学技术进步三等奖”1项。

规律运动对人体健康有诸多益处，尤其在预防和治疗肥胖症、2型糖尿病以及心血管疾病等方面发挥重要作用^［1］。研究表明，适度运动不仅能够降低2型糖尿病的发病风险，有助于2型糖尿病患者提高胰岛素敏感性和控制血糖水平，还可以降低心血管疾病以及心血管不良事件的发生率^［2］。对大鼠耐力训练后的全生物组分子图谱进行分析，结果发现耐力运动可以下调脂肪组织和肝脏中与肥胖症、2型糖尿病以及心血管疾病相关基因的表达^［3］。对小鼠运动前后的心脏、肝脏、脂肪组织以及血浆等多个组织进行代谢组学分析，发现运动对代谢的影响与运动持续时间密切相关^［4］。临床研究也发现，机体的昼夜节律、运动时间的选择以及运动后的饮食构成都是影响运动效果的因素^［5-6］。此外，运动对不同个体之间的效果也存在较大的差异^［7］。因此，为了将运动转化为可精准把控的治疗手段，需要详细阐明运动促进机体健康的分子机制。

运动因子（exerkines）是机体在运动过程中合成和分泌的一系列生物活性物质，其通过自分泌、旁分泌或者内分泌等方式作用于靶细胞来发挥生物学效应^［7］。最初，对运动因子的研究主要聚焦于骨骼肌来源的肌肉因子（myokines）^［7］。随着研究的不断深入以及高通量检测技术的应用，脂肪组织、肝脏和神经系统也逐渐被发现可以分泌运动因子，分别被命名为脂肪因子（adipokines）、肝因子（hepatokines）和神经因子（neurokines）^［8］（图1）。运动因子主要以细胞囊泡的形式被分泌到细胞外，其种类涵盖蛋白多肽类细胞因子、小分子代谢物、脂质分子以及核酸等一系列信号分子^［7］。运动因子所介导的生物学效应可被复制和调控，从而有可能通过类似药物的干预来模拟运动产生的有益效果，为运动不耐受患者以及残疾人群等无法进行运动的人群制订预防和治疗疾病的策略。目前，对动物模型和人体的研究表明，运动因子可对心血管系统发挥重要的保护作用^［9］。本综述将围绕不同运动因子对心血管系统的调控作用展开阐述。

1 运动因子对心血管系统的调控作用

1.1 白细胞介素- 6

白细胞介素-6（interleukin-6，IL-6）是较早进入研究者视野以及研究较为深入的运动因子。在静息状态下，IL-6储存于骨骼肌细胞的囊泡中；在运动过程中，IL-6以胞吐方式释放到细胞外，同时骨骼肌细胞可通过上调IL-6的mRNA表达对细胞内的IL-6进行补充^［10-11］。骨骼肌来源的IL-6释放受到运动强度的影响。与静息状态相比，低强度运动（约为最大摄氧量的50%）可使血浆中IL-6水平升高1.4倍；而进行相同时间的高强度运动（约为最大摄氧量的69%）则使血浆中IL-6水平提高2.7倍^［12］。

运动诱导骨骼肌分泌IL-6的分子机制尚未完全明确，可能存在以下三种分子机制^［13］。一是细胞产生的乳酸介导骨骼肌细胞分泌IL-6。在高强度运动期间，人体循环中IL-6水平与乳酸水平呈明显正相关；予小鼠肌内注射乳酸能够模拟运动诱导的IL-6分泌^［14］。这在一定程度上可以解释IL-6的分泌与运动强度之间的关系。然而，中等强度的运动在乳酸水平较低的情况下也能诱导大量的IL-6分泌，其具体原因尚不明确^［13］。二是IL-6 的分泌受到AMP依赖的蛋白激酶（AMP-activated protein kinase，AMPK）的调控。AMPK激动剂阿卡地新（acadesine，AICAR）能够诱导小鼠骨骼肌分泌IL-6，这表明AMPK介导的信号通路也是运动诱导骨骼肌分泌 IL-6 的重要机制^［11］。此外，骨骼肌分泌的IL-6还能以自分泌的形式作用于骨骼肌，持续激活AMPK并诱导分泌更多的IL-6，这种现象在一定程度上可以解释高强度运动期间IL-6水平急速上升的原因^{［11，15］}。三是与骨骼肌去极化有关。运动期间，去极化的骨骼肌分泌ATP，胞外的ATP作用于肌膜上的嘌呤受体，诱导肌浆网Ca²⁺释放，提高骨骼肌细胞内Ca²⁺水平。随后，核因子κB（nuclear factor-κB，NF-κB）以及激活蛋白-1（activator protein 1，AP-1）被Ca²⁺激活，进而促进骨骼肌IL-6的表达^［15］。

运动诱导释放的IL-6对调节机体代谢以及保护心血管系统发挥重要作用。骨骼肌不仅是IL-6 的重要来源，也是IL-6作用的关键靶器官之一。运动诱导的IL-6可提高骨骼肌细胞中葡萄糖转运蛋白4（glucose transporter 4，GLUT4）的表达，促进运动期间骨骼肌细胞对葡萄糖的摄取；同时IL-6还能增强骨骼肌的脂肪酸氧化^［1，16］。除了以自分泌形式调控骨骼肌代谢，IL-6还可调节心脏代谢，在预防和治疗心血管疾病中发挥积极效用。有研究报道，有氧运动可以降低肥胖患者的心外膜和腹腔内脏脂肪组织水平，改善肥胖导致的心室肥大；使用IL-6受体拮抗剂（托珠单抗）后，有氧运动减少心外膜和腹腔内脏脂肪组织的效果则被抵消^［17-18］。在小鼠心肌缺血/再灌注模型中，运动能够减少心肌梗死的面积以及心律失常的发生；而在IL-6基因敲除小鼠中，运动对小鼠心肌的保护作用则明显削弱^［19］。这表明，IL-6是维持机体健康的重要运动因子，具有改善糖脂代谢以及缓解心脏损伤的作用。

1.2 肌肉素

肌肉素（musclin）是骨骼肌分泌的多肽类分子。在小鼠模型中，运动通过激活骨骼肌中Ca²⁺/Akt信号通路，降低叉头框蛋白O1（forkhead box O1，FOXO1）的活性，进而上调肌肉素编码基因骨织素（osteocrin，OSTN）的表达，提高小鼠循环中肌肉素的水平^［20］。肌肉素可通过提高骨骼肌细胞中环磷酸鸟苷（cyclic guanosine 3′，5′-monophosphate，cGMP）的水平，增强过氧化物酶体增殖物激活受体γ辅激活因子1α（peroxisome proliferator-activated receptor-γcoactivator-1-α，PGC-1α）介导的线粒体生物发生，提高小鼠的有氧运动耐力^［20］。在OSTN基因敲除小鼠中，外源性补充与有氧运动时相同剂量的肌肉素，能够使其恢复有氧运动耐力^［20］。

运动诱导的肌肉素能够减轻心脏损伤。在小鼠心脏压力负荷模型中，转染腺病毒过表达肌肉素可以通过激活蛋白激酶A（protein kinase A，PKA）信号通路促进心肌收缩；通过激活蛋白激酶G（protein kinase G，PKG）信号通路抑制心肌纤维化^［21］。在小鼠心肌缺血/再灌注模型中，运动可以升高血浆中肌肉素的水平，其通过激活心肌细胞中PKG/PGC-1α信号通路，增强心肌线粒体生物发生，缓解心肌损伤；对Ostn基因敲除小鼠和缺乏运动小鼠注射与运动产生的相同剂量肌肉素，能够复制相似的保护作用^［22］。这些结果表明，肌肉素具有成为治疗心脏疾病药物的潜力。

肌肉素还可能与2型糖尿病以及肥胖症的发生有关。无论是肥胖人群还是肥胖小鼠，骨骼肌中肌肉素的表达均高于正常对照组^［23］。转染腺病毒过表达肌肉素能够升高高脂饮食小鼠的空腹血糖水平，加重胰岛素抵抗；使用肌肉素中和抗体则显著降低高脂饮食小鼠的空腹血糖，改善胰岛素抵抗，并减少内脏脂肪组织重量^［23］。临床研究也证实，人体循环中肌肉素水平与内脏脂肪质量呈正相关，与高胰岛素血症和胰岛素抵抗也密切相关^［24］。因此，肌肉素有可能抑制机体的分解代谢，不利于改善糖脂代谢紊乱。

1.3 鸢尾素

鸢尾素（irisin）是由运动诱导分泌的一种蛋白类肌肉因子，来源于前体蛋白纤连蛋白Ⅲ型结构域含蛋白5（fibronectin type III domain-containing protein 5，FNDC5）^［25］。FNDC5是一种细胞膜蛋白，运动通过上调骨骼肌细胞中PGC-1α的表达来提高FNDC5的表达水平，并促进FNDC5在细胞外的N端部分被酶解，产生鸢尾素并释放到血液中^［26］。

研究发现，运动诱导的鸢尾素对糖脂代谢具有重要调节作用。肥胖小鼠进行有氧运动或注射外源性鸢尾素，能够通过上调白色脂肪组织中解偶联蛋白-1（uncoupling protein-1，UCP-1）、PGC-1α 以及其他产热基因的表达，促使白色脂肪组织转化为棕色脂肪组织；棕色脂肪组织可吸收并利用过量的能量物质（葡萄糖或脂肪酸）产生热量，从而提高能量消耗、降低体质量以及改善胰岛素抵抗^［27-28］。此外，鸢尾素还可促进脂肪前体细胞分化。对小鼠进行脂肪组织单细胞RNA测序，发现鸢尾素能够刺激脂肪前体细胞CD81与整合素α V/β 1和α V/ β5

形成复合物并激活粘着

斑激酶（focal adhesion kinase，FAK）信号通路，促进脂肪前体细胞增殖以及分化为米色脂肪；敲除CD81则诱导小鼠肥胖和胰岛素抵抗^［30］。临床研究也发现，肥胖儿童及2型糖尿病患者的血浆鸢尾素水平均低于正常水平，这表明鸢尾素水平下降与代谢紊乱有一定相关性^［31-32］。由此可见，鸢尾素具有促进糖脂代谢以及缓解胰岛素抵抗的作用，有助于改善肥胖和维持血糖平衡。

运动诱导的鸢尾素对心血管系统也有直接保护作用。抗阻运动能够增加小鼠心肌细胞中鸢尾素的表达，激活Parkin-LC3/p62信号通路，引发线粒体自噬并降低心肌细胞氧化应激，减轻小鼠的心肌缺血性损伤^［33］。此外，鸢尾素可以通过增强自噬来提高清除活性氧的能力，拮抗阿霉素诱导的内皮-间质转化，保护血管内皮细胞^［34］。有氧运动能够通过增加肥胖患者循环中鸢尾素水平，降低动脉僵硬度^［35］。

1.4 肌联素

肌联素（myonectin）也被称为补体C1q/肿瘤坏死因子相关蛋白15，运动能够增加骨骼肌中肌联素表达水平以及升高循环中肌联素水平^［36］。肌联素可通过作用于肝细胞和脂肪细胞调节机体的脂质代谢。肌联素能够上调肝细胞和脂肪细胞中CD36、脂肪酸转运蛋白1（fatty acid transport protein 1，FATP1）、脂肪酸结合蛋白1（fatty acid binding protein 1，FABP1）以及脂肪酸结合蛋白4（fatty acid binding protein 4，FABP4）的表达，提高细胞对脂肪酸的摄取能力，降低游离脂肪酸的水平^［37］。此外，肌联素还可以直接减轻心脏损伤。运动能够提高小鼠血浆中肌联素的水平，激活心肌细胞中鞘氨醇-1-磷酸（sphingosine-1-phosphate，S1P）信号通路，减少心肌缺血/再灌注介导的细胞凋亡，缩小心肌梗死面积；下调肌联素表达，运动对心脏的保护作用则明显减弱^［38］。

此外，肌联素水平除了受运动调控，还受机体营养状况的影响。在禁食期间，小鼠体内肌联素的水平下降，重新喂食后，骨骼肌中肌联素mRNA水平以及循环中肌联素水平会迅速升高^［36］。临床研究也发现，与健康人相比，2型糖尿病患者血浆中肌联素水平明显升高，并与甘油三酯、空腹血糖以及葡萄糖耐量试验后2 h血糖水平呈正相关^［39］。因此，肌联素也是一种营养感知性肌肉因子，调节机体对葡萄糖和脂质等营养物质的反应，有可能会成为诊断胰岛素抵抗和2型糖尿病的依据之一^［39-40］。

1.5 脂联素

脂联素（adiponectin）是一种主要由脂肪组织分泌的糖蛋白，骨髓细胞、骨骼肌细胞及心肌细胞也具有分泌脂联素的能力^［41］。人体脂联素包含244个氨基酸残基，而小鼠脂联素则由247个氨基酸残基组成^［42］。脂联素单体（相对分子量30 kDa）仅存在于脂肪细胞中，但常以多聚体的形式分泌到血液循环中，其多聚体形式包括三聚体、六聚体以及高分子量多聚体^［41］。脂联素受体包括脂联素受体1（adiponectin receptor 1，AdipoR1）、脂联素受体2（adiponectin receptors 2，AdipoR2）和T-钙黏素（t-cadherin）^［42-43］。

目前，关于运动对脂联素分泌的影响尚未有定论。高强度和中强度的运动对中心型肥胖患者循环中脂联素的水平和组成具有不同影响^［44］。高强度运动会降低中心型肥胖患者血浆中总脂联素的水平，而中强度运动则对总脂联素的水平无明显影响。进一步研究发现，高强度运动不影响高分子量脂联素在循环中的水平，但三聚体和六聚体的水平下降，这表明是循环中低分子量形式的脂联素水平下降导致了总脂联素水平降低^［44］。另一项研究却得出不同结论，进行20 min运动后，机体血浆中脂联素的水平与运动前并无显著差异；但在休息30 min后，血浆中脂联素水平才有所升高^［45］。因此，运动对脂联素分泌的影响可能与运动形式以及运动强度有关。

脂联素对机体的心血管系统具有重要影响。在小鼠久坐模型中，有氧运动可增加野生型小鼠的左心室射血分数和冠状动脉血流，然而对脂联素基因敲除小鼠的心脏功能则并无改善作用^［46］。此外，脂联素缺失还能增加ApoE^-/-小鼠血浆中干扰素诱导蛋白10（interferon-inducible protein-10，IP-10）的水平，诱导T细胞募集到动脉粥样斑块中，加速动脉粥样硬化的进程^［47］。这表明，脂联素具有成为治疗心血管疾病药物靶点的潜力。

1.6 成纤维细胞生长因子 21

成纤维细胞生长因子 21（fibroblast growth factor 21，FGF21）主要由肝脏分泌，其通过与由成纤维细胞生长因子受体1（fibroblast growth factor receptor 1，FGFR1）和β-Klotho（KLB）组成的受体复合物结合而发挥作用^［48］。经过抗阻训练，小鼠脂肪组织中FGF21水平明显升高^［49］。然而，外周循环中FGF21水平则不同。急性运动能够提高人体和小鼠循环中FGF21的水平；然而，长期运动（≥4周）则降低血浆中FGF21的水平，这可能与脂肪组织、肝脏和骨骼肌中FGF21受体复合物的表达水平升高以及机体对FGF21的反应性增强有关^［50-51］。

FGF21能够调节糖脂代谢及维持能量稳态。有研究发现，在肥胖小鼠的脂肪组织中，FGF21受体表达下降，出现FGF21抵抗，导致血糖升高和胰岛素抵抗；运动通过激活肥胖小鼠脂肪组织中过氧化物酶体增殖物激活受体γ（peroxisome proliferator-activated receptor γ，PPARγ）上调FGFR1和KLB表达，改善FGF21抵抗，缓解高血糖、胰岛素抵抗等代谢紊乱；脂肪组织中KLB基因敲除后，运动改善肥胖小鼠糖脂代谢紊乱的保护作用则消失^［52］。在FGF21基因敲除小鼠中，运动无法改善胰岛素抵抗及肝脏甘油三酯积累^［53］。这表明，运动通过上调FGF21受体复合物的表达，提高细胞对FGF21的反应性，调节糖脂代谢紊乱，对肥胖症以及2型糖尿病具有一定的治疗效果。

此外，运动能够上调小鼠心肌细胞中KLB的表达，增强心肌细胞对FGF21的反应性，这表明心脏也是FGF21发挥作用的重要靶器官^［54］。FGF21通过激活小鼠心肌细胞中AMPK/FOXO3信号通路，增强线粒体去乙酰化酶沉默信息调节蛋白3（sirtuin 3，SIRT3）表达，促进心肌细胞中线粒体功能恢复，缓解糖尿病导致的心肌病^［54］。有氧运动可以通过FGF21介导的信号通路促进小鼠心肌梗死灶中血管新生，抑制心肌缺血诱导的细胞凋亡、氧化应激和心肌纤维化^［55-56］。因此，运动通过增加心脏FGF21受体的表达，提高心脏对FGF21的反应性，也是运动发挥心血管保护作用的重要环节之一。

1.7 脑源性神经营养性因子

脑源性神经营养性因子（brain - derived neurotrophic factor，BDNF）主要来源于大脑皮质和海马体，并可被释放进入血液循环^［57］。BDNF通过与原肌球蛋白相关激酶受体B（tropomyosin-related kinase B receptor，Trk B）结合，触发下游信号通路并发挥广泛作用^［58］。运动能显著提高人体（包括老年人）循环中BDNF的水平^［59-60］。此外，长期规律运动还可以提高静息状态下人体血液中BDNF水平^［61］。

BDNF/Trk B信号通路在心血管系统中广泛表达，与心血管疾病的发生、发展密切相关。有研究报道，在大鼠心肌梗死模型中，运动能够提高左心室非梗死区心肌中BDNF水平，增加每搏输出量和心指数，抑制心肌梗死后射血分数下降以及左心室舒张期末压升高；Trk B抑制剂ANA-12可以阻断运动对心脏功能的改善作用^［62-63］。临床研究也发现，人体血浆中BDNF水平与动脉粥样硬化指数、C反应蛋白、氧化型低密度脂蛋白（oxidized low-density lipoprotein，ox-LDL）以及罹患心血管疾病的风险呈负相关^［64-65］。在心力衰竭患者中，血浆BDNF水平明显降低，并与患者的预后密切相关^［66］。因此，运动诱导的 BDNF 对心血管系统具有积极的保护作用。

1.8 血管内皮生长因子

血管内皮生长因子（vascular endothelial growth factor，VEGF）是一种重要的血管生成因子，在骨骼肌、周细胞和内皮细胞中均有表达^［67］。在运动过程中，心脏和大脑均有表达VEGF^［68-69］。运动能增加人体骨骼肌中VEGF表达并促进VEGF分泌，促进肌肉组织中毛细血管生成以及降低血压^［70］。在小鼠心肌梗死模型中，运动可以增加心脏中VEGF和Fms样酪氨酸激酶1（fms-like tyrosine kinase 1，Flt-1）表达，促进梗死区域的细胞增殖及血管新生，减少心肌梗死的面积^［71］。运动期间，VEGF的分泌可能受到乳酸的调控。高强度运动能够增强小鼠大脑VEGF表达以及大脑毛细血管密度；注射与运动期间相同剂量的乳酸能够模拟高强度运动介导的血管新生；基因敲除小鼠乳酸受体羟基羧酸受体1（hydroxycarboxylic acid receptor 1，HCAR1），则抑制VEGF的表达，运动诱导血管新生的效应也随之消失^［69］。因此，运动诱导的VEGF具有减轻心血管损伤的作用，但仍需更深入的研究来明确运动诱导VEGF表达的相关机制。

1.9 其他类型的运动因子

肠道菌群相关代谢产物也包含可以作用于心血管系统的运动因子。运动能够改变糖尿病前期患者的肠道菌群组成，促进肠道菌群合成短链脂肪酸以及分解支链氨基酸，改善患者的糖代谢和胰岛素抵抗；对肥胖小鼠进行肠道菌群移植也能产生类似的效果^［72］。在运动过程中，肠道菌群还可以合成一些小分子代谢物（如3-羟基苯乙酸和4-羟基苯甲酸），降低心肌损伤，恢复小鼠心肌梗死后的心脏功能^［73］。此外，microRNA也是运动发挥心血管保护效应的因素之一。高强度间隔训练及中高强度有氧运动均能提高循环中miR-126水平，miR-126能够促进血管新生，调控心肌细胞自噬，抑制细胞凋亡和炎症，对糖尿病患者的心血管系统发挥保护作用^［74］。运动可以诱导脂肪组织释放12，13-二羟基-9Z-十八烷酸（12，13-diHOME），12，13-diHOME能够促进脂肪组织和骨骼肌摄取脂肪酸，改善心脏舒张和收缩功能^［75］。另外，在运动过程中，骨骼肌释放的乳酸可上调脂肪组织中转化生长因子-β2表达，改善胰岛素抵抗，提高葡萄糖耐量^［76］。

2 总结与展望

运动能够促进机体多个组织及器官合成和分泌诸如IL-6、肌肉素、鸢尾素等运动因子，其通过以自分泌、旁分泌或内分泌的形式与多种组织或器官的靶细胞上相应受体结合，发挥改善糖脂代谢以及预防和治疗心血管疾病的保护作用。关于运动因子的研究，揭示了运动保护机体心血管系统的分子机制针对运动因子、运动因子模拟物以及运动因子受体激动剂的研究也成为关注热点，被寄望成为治疗心血管疾病的新型药物。然而，随着研究的持续深入，发现运动因子的分泌依赖于运动类型、运动强度以及运动持续时间。不同的运动类型、强度以及持续时间，在不同的研究之间存在诸多差异，这使得运动因子在运动中的变化缺乏一致的结论，直接将运动因子应用于临床治疗中还存在很多困惑。因此，不同的运动模式对各种运动因子的影响仍需要进一步明确，各种运动因子之间的调控网络以及不同运动因子对各个器官的影响也需要进一步深入探讨。另外，随着蛋白质组学、代谢组学、脂质组学等组学技术的进步，越来越多的运动因子被发现和验证，然而这些运动因子作用的部位、结合受体后引起的生物学作用及其分子机制、在体内发挥的整体性效应，尚缺乏相关的研究。总体而言，运动因子是一个极具应用价值的研究对象，有潜力成为临床治疗心血管疾病的新靶点。

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11、%E2%80%83%20LAURITZEN%E2%80%83H%E2%80%83P%EF%BC%8CBRANDAUER%E2%80%83J%EF%BC%8CSCHJERLING%E2%80%83%0AP%EF%BC%8Cet%E2%80%83al%EF%BC%8EContraction%E2%80%83%20and%E2%80%83AICAR%E2%80%83%20stimulate%E2%80%83%20IL-6%E2%80%83%0Avesicle%E2%80%83depletion%E2%80%83from%E2%80%83skeletal%E2%80%83muscle%E2%80%83fibers%E2%80%83in%E2%80%83vivo%0A%EF%BC%BBJ%EF%BC%BD%EF%BC%8EDiabetes%EF%BC%8C2013%EF%BC%8C62%EF%BC%889%EF%BC%89%EF%BC%9A3081-3092%EF%BC%8E

12、CULLEN%E2%80%83T%EF%BC%8CTHOMAS%E2%80%83A%E2%80%83W%EF%BC%8CWEBB%E2%80%83R%EF%BC%8Cet%E2%80%83al%EF%BC%8E%0AInterleukin-6%E2%80%83and%E2%80%83associated%E2%80%83cytokine%E2%80%83%20responses%E2%80%83to%E2%80%83an%E2%80%83%0Aacute%E2%80%83bout%E2%80%83of%E2%80%83high-intensity%E2%80%83interval%E2%80%83exercise%EF%BC%9AThe%E2%80%83%0Aeffect%E2%80%83of%E2%80%83exercise%E2%80%83intensity%E2%80%83and%E2%80%83volume%EF%BC%BBJ%EF%BC%BD%EF%BC%8EAppl%E2%80%83%0APhysiol%E2%80%83Nutr%E2%80%83Metab%EF%BC%8C2016%EF%BC%8C41%EF%BC%888%EF%BC%89%EF%BC%9A803-808%EF%BC%8E

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17、CHRISTENSEN%E2%80%83R%E2%80%83H%EF%BC%8CLEHRSKOV%E2%80%83L%E2%80%83L%EF%BC%8CWEDELL%02NEERGAARD%E2%80%83A%E2%80%83S%EF%BC%8Cet%E2%80%83al%EF%BC%8EAerobic%E2%80%83exercise%E2%80%83induces%E2%80%83%0Acardiac%E2%80%83fat%E2%80%83loss%E2%80%83and%E2%80%83alters%E2%80%83cardiac%E2%80%83muscle%E2%80%83mass%E2%80%83through%E2%80%83%0Aan%E2%80%83interleukin-6%E2%80%83receptor-dependent%E2%80%83mechanism%EF%BC%9A%0ACardiac%E2%80%83%20analysis%E2%80%83%20of%E2%80%83%20a%E2%80%83%20double-blind%E2%80%83%20randomized%E2%80%83%0Acontrolled%E2%80%83clinical%E2%80%83trial%E2%80%83in%E2%80%83abdominally%E2%80%83obese%E2%80%83humans%0A%EF%BC%BBJ%EF%BC%BD%EF%BC%8ECirculation%EF%BC%8C2019%EF%BC%8C140%EF%BC%8820%EF%BC%89%EF%BC%9A1684-1686%EF%BC%8E

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19、MCGINNIS%E2%80%83G%E2%80%83R%EF%BC%8CBALLMANN%E2%80%83C%EF%BC%8CPETERS%E2%80%83B%EF%BC%8Cet%E2%80%83%0Aal%EF%BC%8EInterleukin-6%E2%80%83mediates%E2%80%83exercise%E2%80%83%20preconditioning%E2%80%83%0Aagainst%E2%80%83myocardial%E2%80%83ischemia%E2%80%83reperfusion%E2%80%83injury%EF%BC%BBJ%EF%BC%BD%EF%BC%8E%0AAm%E2%80%83J%E2%80%83Physiol%E2%80%83Heart%E2%80%83Circ%E2%80%83Physiol%EF%BC%8C2015%EF%BC%8C308%EF%BC%8811%EF%BC%89%EF%BC%9A%0AH1423-H1433%EF%BC%8E

20、SUBBOTINA%E2%80%83E%EF%BC%8CSIERRA%E2%80%83A%EF%BC%8CZHU%E2%80%83Z%EF%BC%8Cet%E2%80%83al%EF%BC%8EMusclin%E2%80%83%0Ais%E2%80%83%20an%E2%80%83%20activity-stimulated%E2%80%83%20myokine%E2%80%83%20that%E2%80%83%20enhances%E2%80%83%0Aphysical%E2%80%83endurance%EF%BC%BBJ%EF%BC%BD%EF%BC%8EProc%E2%80%83Natl%E2%80%83Acad%E2%80%83Sci%E2%80%83U%E2%80%83S%E2%80%83A%EF%BC%8C%0A2015%EF%BC%8C112%EF%BC%8852%EF%BC%89%EF%BC%9A16042-16047%EF%BC%8E

21、SZAROSZYK%E2%80%83M%EF%BC%8CKATTIH%E2%80%83B%EF%BC%8CMARTIN-GARRIDO%E2%80%83%0AA%EF%BC%8Cet%E2%80%83al%EF%BC%8ESkeletal%E2%80%83muscle%E2%80%83%20derived%E2%80%83Musclin%E2%80%83%20protects%E2%80%83%0Athe%E2%80%83heart%E2%80%83during%E2%80%83pathological%E2%80%83overload%EF%BC%BBJ%EF%BC%BD%EF%BC%8ENat%E2%80%83%0ACommun%EF%BC%8C2022%EF%BC%8C13%EF%BC%881%EF%BC%89%EF%BC%9A149%EF%BC%8E

22、HARRIS%E2%80%83M%E2%80%83P%EF%BC%8CZENG%E2%80%83S%EF%BC%8CZHU%E2%80%83Z%EF%BC%8Cet%E2%80%83al%EF%BC%8EMyokine%E2%80%83%0Amusclin%E2%80%83%20is%E2%80%83%20critical%E2%80%83%20for%E2%80%83%20exercise-induced%E2%80%83%20cardiac%E2%80%83%0Aconditioning%EF%BC%BBJ%EF%BC%BD%EF%BC%8EInt%E2%80%83J%E2%80%83Mol%E2%80%83Sci%EF%BC%8C2023%EF%BC%8C24%EF%BC%887%EF%BC%89%EF%BC%9A%0A6525%EF%BC%8E

23、JIN%E2%80%83L%EF%BC%8CHAN%E2%80%83S%EF%BC%8CLV%E2%80%83X%EF%BC%8Cet%E2%80%83al%EF%BC%8EThe%E2%80%83muscle-enriched%E2%80%83%0Amyokine%E2%80%83Musclin%E2%80%83impairs%E2%80%83beige%E2%80%83fat%E2%80%83thermogenesis%E2%80%83and%E2%80%83%0Asystemic%E2%80%83energy%E2%80%83homeostasis%E2%80%83via%E2%80%83Tfr1%2FPKA%E2%80%83signaling%E2%80%83in%E2%80%83%0Amale%E2%80%83mice%EF%BC%BBJ%EF%BC%BD%EF%BC%8ENat%E2%80%83Commun%EF%BC%8C2023%EF%BC%8C14%EF%BC%881%EF%BC%89%EF%BC%9A%0A4257%EF%BC%8E

24、SANCHEZ%E2%80%83Y%E2%80%83L%EF%BC%8CYEPES-CALDERON%E2%80%83M%EF%BC%8C%0AVALBUENA%E2%80%83L%EF%BC%8Cet%E2%80%83al%EF%BC%8EMusclin%E2%80%83is%E2%80%83%20related%E2%80%83to%E2%80%83insulin%E2%80%83%0Aresistance%E2%80%83and%E2%80%83body%E2%80%83composition%EF%BC%8Cbut%E2%80%83not%E2%80%83to%E2%80%83body%E2%80%83mass%E2%80%83%0Aindex%E2%80%83or%E2%80%83cardiorespiratory%E2%80%83capacity%E2%80%83in%E2%80%83adults%EF%BC%BBJ%EF%BC%BD%EF%BC%8E%0AEndocrinol%E2%80%83Metab%EF%BC%8C2021%EF%BC%8C36%EF%BC%885%EF%BC%89%EF%BC%9A1055-1068%EF%BC%8E

25、MAAK%E2%80%83S%EF%BC%8CNORHEIM%E2%80%83F%EF%BC%8CDREVON%E2%80%83C%E2%80%83A%EF%BC%8Cet%E2%80%83al%EF%BC%8E%0AProgress%E2%80%83and%E2%80%83challenges%E2%80%83in%E2%80%83the%E2%80%83biology%E2%80%83of%E2%80%83FNDC5%E2%80%83and%E2%80%83%0Airisin%EF%BC%BBJ%EF%BC%BD%EF%BC%8EEndocr%E2%80%83Rev%EF%BC%8C2021%EF%BC%8C42%EF%BC%884%EF%BC%89%EF%BC%9A436-%0A456%EF%BC%8E

26、SOUSA%E2%80%83R%E2%80%83A%E2%80%83L%EF%BC%8CIMPROTA-CARIA%E2%80%83A%E2%80%83C%EF%BC%8CSOUZA%E2%80%83B%E2%80%83%0AS%E2%80%83F%EF%BC%8EExercise-linked%E2%80%83irisin%EF%BC%9AConsequences%E2%80%83on%E2%80%83mental%E2%80%83and%E2%80%83cardiovascular%E2%80%83health%E2%80%83in%E2%80%83type%E2%80%832%E2%80%83diabetes%EF%BC%BBJ%EF%BC%BD%EF%BC%8EInt%E2%80%83%0AJ%E2%80%83Mol%E2%80%83Sci%EF%BC%8C2021%EF%BC%8C22%EF%BC%884%EF%BC%89%EF%BC%9A2199%EF%BC%8E

27、LIU%E2%80%83S%EF%BC%8CCUI%E2%80%83F%EF%BC%8CNING%E2%80%83K%EF%BC%8Cet%E2%80%83al%EF%BC%8ERole%E2%80%83%20of%E2%80%83irisin%E2%80%83in%E2%80%83%0Aphysiology%E2%80%83and%E2%80%83pathology%EF%BC%BBJ%EF%BC%BD%EF%BC%8EFront%E2%80%83Endocrinol%EF%BC%8C%0A2022%EF%BC%8813%EF%BC%89%EF%BC%9A962968%EF%BC%8E

28、%E2%80%83%20BOSTROM%E2%80%83P%EF%BC%8CWU%E2%80%83J%EF%BC%8CJEDRYCHOWSKI%E2%80%83M%E2%80%83P%EF%BC%8C%0Aet%E2%80%83al%EF%BC%8EA%E2%80%83%20PGC1-alpha-dependent%E2%80%83%20myokine%E2%80%83%20that%E2%80%83%0Adrives%E2%80%83%20brown-fat-like%E2%80%83%20development%E2%80%83%20of%E2%80%83%20white%E2%80%83%20fat%E2%80%83%0Aand%E2%80%83thermogenesis%EF%BC%BBJ%EF%BC%BD%EF%BC%8ENature%EF%BC%8C2012%EF%BC%8C481%0A%EF%BC%887382%EF%BC%89%EF%BC%9A463-468%EF%BC%8E

29、OGURI%E2%80%83Y%EF%BC%8CSHINODA%E2%80%83K%EF%BC%8CKIM%E2%80%83H%EF%BC%8Cet%E2%80%83al%EF%BC%8ECD81%E2%80%83%0Acontrols%E2%80%83%20beige%E2%80%83fat%E2%80%83%20progenitor%E2%80%83cell%E2%80%83growth%E2%80%83and%E2%80%83energy%E2%80%83%0Abalance%E2%80%83via%E2%80%83FAK%E2%80%83signaling%EF%BC%BBJ%EF%BC%BD%EF%BC%8ECell%EF%BC%8C2020%EF%BC%8C182%0A%EF%BC%883%EF%BC%89%EF%BC%9A563-577%EF%BC%8Ee20%EF%BC%8E

30、EL-LEBEDY%E2%80%83D%E2%80%83H%EF%BC%8CIBRAHIM%E2%80%83A%E2%80%83A%EF%BC%8CASHMAWY%E2%80%83I%E2%80%83O%EF%BC%8E%0ANovel%E2%80%83adipokines%E2%80%83vaspin%E2%80%83and%E2%80%83irisin%E2%80%83as%E2%80%83%20risk%E2%80%83biomarkers%E2%80%83%0Afor%E2%80%83cardiovascular%E2%80%83diseases%E2%80%83in%E2%80%83type%E2%80%832%E2%80%83diabetes%E2%80%83mellitus%0A%EF%BC%BBJ%EF%BC%BD%EF%BC%8EDiabetes%E2%80%83Metab%E2%80%83Syndr%EF%BC%8C2018%EF%BC%8C12%EF%BC%885%EF%BC%89%EF%BC%9A%0A643-648%EF%BC%8E

31、YIN%E2%80%83C%EF%BC%8CHU%E2%80%83W%EF%BC%8CWANG%E2%80%83M%EF%BC%8Cet%E2%80%83al%EF%BC%8EI%20risin%E2%80%83%20as%E2%80%83%20a%E2%80%83%0Amediator%E2%80%83between%E2%80%83obesity%E2%80%83and%E2%80%83vascular%E2%80%83inflammation%E2%80%83in%E2%80%83%0AChinese%E2%80%83children%E2%80%83and%E2%80%83adolescents%EF%BC%BBJ%EF%BC%BD%EF%BC%8ENutr%E2%80%83Metab%E2%80%83%0ACardiovasc%E2%80%83Dis%EF%BC%8C2020%EF%BC%8C30%EF%BC%882%EF%BC%89%EF%BC%9A320-329%EF%BC%8E

32、LI%E2%80%83H%EF%BC%8CQIN%E2%80%83S%EF%BC%8CLIANG%E2%80%83Q%EF%BC%8Cet%E2%80%83al%EF%BC%8EExercise%E2%80%83training%E2%80%83%0Aenhances%E2%80%83myocardial%E2%80%83mitophagy%E2%80%83and%E2%80%83improves%E2%80%83cardiac%E2%80%83%0Afunction%E2%80%83via%E2%80%83irisin%2FFNDC5-PINK1%2FParkin%E2%80%83%20pathway%E2%80%83in%E2%80%83%0AMI%E2%80%83mice%EF%BC%BBJ%EF%BC%BD%EF%BC%8EBiomedicines%EF%BC%8C2021%EF%BC%8C9%EF%BC%886%EF%BC%89%EF%BC%9A701%EF%BC%8E

33、PAN%E2%80%83J%E2%80%83A%EF%BC%8CZHANG%E2%80%83H%EF%BC%8CLIN%E2%80%83H%EF%BC%8Cet%E2%80%83al%EF%BC%8EI%20ri%20si%20n%E2%80%83%0Aameliorates%E2%80%83doxorubicin-induced%E2%80%83cardiac%E2%80%83perivascular%E2%80%83%0Afibrosis%E2%80%83through%E2%80%83inhibiting%E2%80%83endothelial-to-mesenchymal%E2%80%83%0Atransition%E2%80%83%20by%E2%80%83%20regulating%E2%80%83%20ROS%E2%80%83%20accumulation%E2%80%83%20and%E2%80%83%0Aautophagy%E2%80%83disorder%E2%80%83in%E2%80%83endothelial%E2%80%83cells%EF%BC%BBJ%EF%BC%BD%EF%BC%8ERedox%E2%80%83%0ABiol%EF%BC%8C2021%EF%BC%8846%EF%BC%89%EF%BC%9A102120%EF%BC%8E

34、%E2%80%83INOUE%E2%80%83K%EF%BC%8CFUJIE%E2%80%83S%EF%BC%8CHASEGAWA%E2%80%83N%EF%BC%8Cet%E2%80%83al%EF%BC%8E%0AAerobic%E2%80%83exercise%E2%80%83training-induced%E2%80%83irisin%E2%80%83%20secretion%E2%80%83is%E2%80%83%0Aassociated%E2%80%83with%E2%80%83the%E2%80%83%20reduction%E2%80%83of%E2%80%83arterial%E2%80%83%20stiffness%E2%80%83via%E2%80%83%0Anitric%E2%80%83oxide%E2%80%83production%E2%80%83in%E2%80%83adults%E2%80%83with%E2%80%83obesity%EF%BC%BBJ%EF%BC%BD%EF%BC%8E%0AAppl%E2%80%83Physiol%E2%80%83Nutr%E2%80%83Metab%EF%BC%8C2020%EF%BC%8C45%EF%BC%887%EF%BC%89%EF%BC%9A715-722%EF%BC%8E

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39、%E2%80%83%20DAS%E2%80%83D%E2%80%83K%EF%BC%8CGRAHAM%E2%80%83Z%E2%80%83A%EF%BC%8CCARDOZO%E2%80%83C%E2%80%83P%EF%BC%8EMyokines%E2%80%83%0Ain%E2%80%83skeletal%E2%80%83muscle%E2%80%83physiology%E2%80%83and%E2%80%83metabolism%EF%BC%9A%0ARecent%E2%80%83advances%E2%80%83and%E2%80%83future%E2%80%83perspectives%EF%BC%BBJ%EF%BC%BD%EF%BC%8EActa%E2%80%83%0APhysiol%EF%BC%8C2020%EF%BC%8C228%EF%BC%882%EF%BC%89%EF%BC%9Ae13367%EF%BC%8E

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41、%E2%80%83%20MAEDA%E2%80%83N%EF%BC%8CFUNAHASHI%E2%80%83T%EF%BC%8CMATSUZAWA%E2%80%83Y%EF%BC%8C%0Aet%E2%80%83al%EF%BC%8EAdiponectin%EF%BC%8Ca%E2%80%83%20unique%E2%80%83%20adipocyte-derived%E2%80%83%0Afactor%E2%80%83beyond%E2%80%83hormones%EF%BC%BBJ%EF%BC%BD%EF%BC%8EAtherosclerosis%EF%BC%8C2020%0A%EF%BC%88292%EF%BC%89%EF%BC%9A1-9%EF%BC%8E

42、%E2%80%83%20SCHEJA%E2%80%83L%EF%BC%8CHEEREN%E2%80%83J%EF%BC%8EThe%E2%80%83endocrine%E2%80%83function%E2%80%83of%E2%80%83%0Aadipose%E2%80%83tissues%E2%80%83in%E2%80%83health%E2%80%83and%E2%80%83cardiometabolic%E2%80%83disease%0A%EF%BC%BBJ%EF%BC%BD%EF%BC%8ENat%E2%80%83Rev%E2%80%83Endocrinol%EF%BC%8C2019%EF%BC%8C15%EF%BC%889%EF%BC%89%EF%BC%9A507-%0A524%EF%BC%8E

43、%E2%80%83%20OBATA%E2%80%83Y%EF%BC%8CKITA%E2%80%83S%EF%BC%8CKOYAMA%E2%80%83Y%EF%BC%8Cet%E2%80%83al%EF%BC%8EAdiponectin%2F%0AT-cadherin%E2%80%83system%E2%80%83enhances%E2%80%83exosome%E2%80%83biogenesis%E2%80%83%20and%E2%80%83%0Adecreases%E2%80%83cellular%E2%80%83ceramides%E2%80%83by%E2%80%83exosomal%E2%80%83release%EF%BC%BBJ%EF%BC%BD%EF%BC%8E%0AJCI%E2%80%83Insight%EF%BC%8C2018%EF%BC%8C3%EF%BC%888%EF%BC%89%EF%BC%9A99680.

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45、%E2%80%83%20JURIMAE%E2%80%83J%EF%BC%8CPURGE%E2%80%83P%EF%BC%8CJURIMAE%E2%80%83T%EF%BC%8EAdiponectin%E2%80%83%0Ais%E2%80%83altered%E2%80%83after%E2%80%83maximal%E2%80%83exercise%E2%80%83in%E2%80%83highly%E2%80%83trained%E2%80%83male%E2%80%83%0Arowers%EF%BC%BBJ%EF%BC%BD%EF%BC%8EEur%E2%80%83J%E2%80%83Appl%E2%80%83Physiol%EF%BC%8C2005%EF%BC%8C93%EF%BC%884%EF%BC%89%EF%BC%9A%0A502-505%EF%BC%8E

46、%E2%80%83%20CALDWELL%E2%80%83J%E2%80%83T%EF%BC%8CJONES%E2%80%83K%E2%80%83M%E2%80%83D%EF%BC%8CPARK%E2%80%83H%EF%BC%8Cet%E2%80%83al%EF%BC%8E%0AAerobic%E2%80%83exercise%E2%80%83training%E2%80%83%20reduces%E2%80%83cardiac%E2%80%83function%E2%80%83and%E2%80%83%0Acoronary%E2%80%83flow-induced%E2%80%83%20vasodilation%E2%80%83in%E2%80%83mice%E2%80%83lacking%E2%80%83%0Aadiponectin%EF%BC%BBJ%EF%BC%BD%EF%BC%8EAm%E2%80%83J%E2%80%83Physiol%E2%80%83Heart%E2%80%83Circ%E2%80%83Physiol%EF%BC%8C%0A2021%EF%BC%8C321%EF%BC%881%EF%BC%89%EF%BC%9AH1-H14%EF%BC%8E

47、OKAMOTO%E2%80%83Y%EF%BC%8CFOLCO%E2%80%83E%E2%80%83J%EF%BC%8CMINAMI%E2%80%83M%EF%BC%8Cet%E2%80%83al%EF%BC%8E%0AAdiponectin%E2%80%83inhibits%E2%80%83the%E2%80%83%20production%E2%80%83of%E2%80%83CXC%E2%80%83%20receptor%E2%80%83%0A3%E2%80%83%20chemokine%E2%80%83ligands%E2%80%83in%E2%80%83%20macrophages%E2%80%83%20and%E2%80%83%20reduces%E2%80%83%0AT-lymphocyte%E2%80%83recruitment%E2%80%83in%E2%80%83atherogenesis%EF%BC%BBJ%EF%BC%BD%EF%BC%8ECirc%E2%80%83%0ARes%EF%BC%8C2008%EF%BC%8C102%EF%BC%882%EF%BC%89%EF%BC%9A218-225%EF%BC%8E

48、ZHANG%E2%80%83T%EF%BC%8CJIANG%E2%80%83D%EF%BC%8CZHANG%E2%80%83X%EF%BC%8Cet%E2%80%83al%EF%BC%8EThe%E2%80%83%20role%E2%80%83%0Aof%E2%80%83nonmyocardial%E2%80%83cells%E2%80%83in%E2%80%83the%E2%80%83development%E2%80%83of%E2%80%83diabetic%E2%80%83%0Acardiomyopathy%E2%80%83and%E2%80%83the%E2%80%83protective%E2%80%83effects%E2%80%83of%E2%80%83FGF21%EF%BC%9Aa%E2%80%83current%E2%80%83understanding%EF%BC%8ECell%E2%80%83Commun%E2%80%83Signal%EF%BC%8E2024%EF%BC%8C%0A22%EF%BC%881%EF%BC%89%EF%BC%9A446%EF%BC%8E

49、PONTES%E2%80%83L%E2%80%83P%E2%80%83P%EF%BC%8CALVES%E2%80%83NAKAKURA%E2%80%83F%E2%80%83C%EF%BC%8CNETO%E2%80%83%0AN%E2%80%83I%E2%80%83P%EF%BC%8Cet%E2%80%83al%EF%BC%8EResistance%E2%80%83and%E2%80%83aerobic%E2%80%83training%E2%80%83were%E2%80%83%0Aeffective%E2%80%83in%E2%80%83activating%E2%80%83different%E2%80%83markers%E2%80%83of%E2%80%83the%E2%80%83browning%E2%80%83%0Aprocess%E2%80%83in%E2%80%83obesity%EF%BC%BBJ%EF%BC%BD%EF%BC%8EInt%E2%80%83J%E2%80%83Mol%E2%80%83Sci%EF%BC%8C2023%EF%BC%8C25%0A%EF%BC%881%EF%BC%89%EF%BC%9A275%EF%BC%8E

50、%E2%80%83PORFLITT-RODRIGUEZ%E2%80%83M%20%EF%BC%8C%20GUZMAN%02ARRIAGADA%E2%80%83V%EF%BC%8CSANDOVAL-VALDERRAMA%E2%80%83R%EF%BC%8C%0Aet%E2%80%83al%EF%BC%8EEffects%E2%80%83of%E2%80%83aerobic%E2%80%83exercise%E2%80%83on%E2%80%83fibroblast%E2%80%83growth%E2%80%83%0Afactor%E2%80%8321%E2%80%83in%E2%80%83overweight%E2%80%83and%E2%80%83obesity%EF%BC%8EA%E2%80%83%20systematic%E2%80%83%0Areview%EF%BC%BBJ%EF%BC%BD%EF%BC%8EMetabolism%EF%BC%8C2022%EF%BC%88129%EF%BC%89%EF%BC%9A155137%EF%BC%8E

51、KIM%E2%80%83K%E2%80%83H%EF%BC%8CKIM%E2%80%83S%E2%80%83H%EF%BC%8CMIN%E2%80%83Y%E2%80%83K%EF%BC%8Cet%E2%80%83al%EF%BC%8EAcute%E2%80%83%0Aexercise%E2%80%83induces%E2%80%83%20FGF21%E2%80%83%20expression%E2%80%83in%E2%80%83mice%E2%80%83%20and%E2%80%83in%E2%80%83%0Ahealthy%E2%80%83humans%EF%BC%BBJ%EF%BC%BD%EF%BC%8EPLoS%E2%80%83One%EF%BC%8C2013%EF%BC%8C8%EF%BC%885%EF%BC%89%EF%BC%9A%0Ae63517%EF%BC%8E

52、GENG%E2%80%83L%EF%BC%8CLIAO%E2%80%83B%EF%BC%8CJIN%E2%80%83L%EF%BC%8Cet%E2%80%83al%EF%BC%8EExercise%E2%80%83alleviates%E2%80%83%0Aobesity-induced%E2%80%83metabolic%E2%80%83dysfunction%E2%80%83via%E2%80%83enhancing%E2%80%83%0AFGF21%E2%80%83sensitivity%E2%80%83in%E2%80%83adipose%E2%80%83tissues%EF%BC%BBJ%EF%BC%BD%EF%BC%8ECell%E2%80%83Rep%EF%BC%8C%0A2019%EF%BC%8C26%EF%BC%8810%EF%BC%89%EF%BC%9A2738-2752%EF%BC%8Ee4%EF%BC%8E

53、LOYD%E2%80%83C%EF%BC%8CMAGRISSO%E2%80%83I%E2%80%83J%EF%BC%8CHAAS%E2%80%83M%EF%BC%8Cet%E2%80%83al%EF%BC%8E%0AFibroblast%E2%80%83growth%E2%80%83factor%E2%80%83%2021%E2%80%83is%E2%80%83%20required%E2%80%83for%E2%80%83%20beneficial%E2%80%83%0Aeffects%E2%80%83of%E2%80%83exercise%E2%80%83during%E2%80%83chronic%E2%80%83high-fat%E2%80%83feeding%EF%BC%BBJ%EF%BC%BD%EF%BC%8E%0AJ%E2%80%83Appl%E2%80%83Physiol%EF%BC%8C2016%EF%BC%8C121%EF%BC%883%EF%BC%89%EF%BC%9A687-698%EF%BC%8E

54、%E2%80%83%20JIN%E2%80%83L%EF%BC%8CGENG%E2%80%83L%EF%BC%8CYING%E2%80%83L%EF%BC%8Cet%E2%80%83al%EF%BC%8EFGF21-sirtuin%E2%80%833%E2%80%83%0Aaxis%E2%80%83confers%E2%80%83the%E2%80%83%20protective%E2%80%83effects%E2%80%83of%E2%80%83exercise%E2%80%83against%E2%80%83%0Adiabetic%E2%80%83cardiomyopathy%E2%80%83%20by%E2%80%83governing%E2%80%83mitochondrial%E2%80%83%0Aintegrity%EF%BC%BBJ%EF%BC%BD%EF%BC%8ECirculation%EF%BC%8C2022%EF%BC%8C146%EF%BC%8820%EF%BC%89%EF%BC%9A%0A1537-1557%EF%BC%8E

55、BO%E2%80%83W%20%EF%BC%8C%20MA%E2%80%83Y%20%EF%BC%8C%20FENG%E2%80%83L%20%EF%BC%8C%20et%E2%80%83al%20%EF%BC%8E%20F%20G%20F%202%201%E2%80%83%0Apromotes%E2%80%83myocardial%E2%80%83angiogenesis%E2%80%83and%E2%80%83mediates%E2%80%83the%E2%80%83%0Acardioprotective%E2%80%83%20effects%E2%80%83%20of%E2%80%83%20exercise%E2%80%83in%E2%80%83%20myocardial%E2%80%83%0Ainfarction%E2%80%83mice%EF%BC%BBJ%EF%BC%BD%EF%BC%8EJ%E2%80%83Appl%E2%80%83Physiol%EF%BC%8C2023%EF%BC%8C135%0A%EF%BC%883%EF%BC%89%EF%BC%9A696-705%EF%BC%8E

56、MA%E2%80%83Y%EF%BC%8CKUANG%E2%80%83Y%EF%BC%8CBO%E2%80%83W%EF%BC%8Cet%E2%80%83al%EF%BC%8EExercise%E2%80%83training%E2%80%83%0Aalleviates%E2%80%83cardiac%E2%80%83fibrosis%E2%80%83through%E2%80%83increasing%E2%80%83fibroblast%E2%80%83%0Agrowth%E2%80%83factor%E2%80%8321%E2%80%83and%E2%80%83regulating%E2%80%83TGF-%CE%B21-Smad2%2F3-%0AMMP2%2F9%E2%80%83signaling%E2%80%83in%E2%80%83mice%E2%80%83with%E2%80%83myocardial%E2%80%83infarction%0A%EF%BC%BBJ%EF%BC%BD%EF%BC%8EInt%E2%80%83J%E2%80%83Mol%E2%80%83Sci%EF%BC%8C2021%EF%BC%8C22%EF%BC%8822%EF%BC%89%EF%BC%9A12341%EF%BC%8E

57、RASMUSSEN%E2%80%83P%EF%BC%8CBRASSARD%E2%80%83P%EF%BC%8CADSER%E2%80%83H%EF%BC%8Cet%E2%80%83al%EF%BC%8E%0AEvidence%E2%80%83for%E2%80%83a%E2%80%83%20release%E2%80%83of%E2%80%83brain-derived%E2%80%83neurotrophic%E2%80%83%0Afactor%E2%80%83from%E2%80%83the%E2%80%83brain%E2%80%83during%E2%80%83exercise%EF%BC%BBJ%EF%BC%BD%EF%BC%8EExp%E2%80%83%0APhysiol%EF%BC%8C2009%EF%BC%8C94%EF%BC%8810%EF%BC%89%EF%BC%9A1062-1069%EF%BC%8E

58、HANG%E2%80%83P%E2%80%83Z%EF%BC%8CZHU%E2%80%83H%EF%BC%8CLI%E2%80%83P%E2%80%83F%EF%BC%8Cet%E2%80%83al%EF%BC%8EThe%E2%80%83emerging%E2%80%83%0Arole%E2%80%83of%E2%80%83BDNF%2FTrkB%E2%80%83signaling%E2%80%83in%E2%80%83cardiovascular%E2%80%83diseases%EF%BC%BBJ%EF%BC%BD%EF%BC%8ELife%EF%BC%8C2021%EF%BC%8C11%EF%BC%881%EF%BC%89%EF%BC%9A70%EF%BC%8E

59、SAUCEDO%E2%80%83MARQUEZ%E2%80%83C%E2%80%83M%EF%BC%8CVANAUDENAERDE%E2%80%83%0AB%EF%BC%8CTROOSTERS%E2%80%83T%EF%BC%8Cet%E2%80%83al%EF%BC%8EHigh-intensity%E2%80%83interval%E2%80%83%0Atraining%E2%80%83evokes%E2%80%83larger%E2%80%83%20serum%E2%80%83BDNF%E2%80%83levels%E2%80%83compared%E2%80%83%0Awith%E2%80%83intense%E2%80%83continuous%E2%80%83exercise%EF%BC%BBJ%EF%BC%BD%EF%BC%8EJ%E2%80%83%20Appl%E2%80%83%0APhysiol%EF%BC%8C2015%EF%BC%8C119%EF%BC%8812%EF%BC%89%EF%BC%9A1363-1373%EF%BC%8E

60、CANTON-MARTINEZ%E2%80%83E%EF%BC%8CRENTERIA%E2%80%83I%EF%BC%8CGARCIA%02SUAREZ%E2%80%83P%E2%80%83C%EF%BC%8Cet%E2%80%83al%EF%BC%8EConcurrent%E2%80%83training%E2%80%83increases%E2%80%83%0Aserum%E2%80%83brain-derived%E2%80%83neurotrophic%E2%80%83factor%E2%80%83in%E2%80%83older%E2%80%83adults%E2%80%83%0Aregardless%E2%80%83of%E2%80%83the%E2%80%83exercise%E2%80%83frequency%EF%BC%BBJ%EF%BC%BD%EF%BC%8EFront%E2%80%83Aging%E2%80%83%0ANeurosci%EF%BC%8C2022%EF%BC%8814%EF%BC%89%EF%BC%9A791698%EF%BC%8E

61、%E2%80%83MURAWSKA-CIALOWICZ%E2%80%83E%EF%BC%8CWOJNA%E2%80%83J%EF%BC%8C%0AZUWALA-JAGIELLO%E2%80%83J%EF%BC%8ECrossfit%E2%80%83training%E2%80%83%20changes%E2%80%83%0Abrain-derived%E2%80%83neurotrophic%E2%80%83factor%E2%80%83and%E2%80%83irisin%E2%80%83levels%E2%80%83at%E2%80%83%0Arest%EF%BC%8Cafter%E2%80%83wingate%E2%80%83and%E2%80%83progressive%E2%80%83tests%EF%BC%8Cand%E2%80%83improves%E2%80%83%0Aaerobic%E2%80%83%20capacity%E2%80%83%20and%E2%80%83%20body%E2%80%83%20composition%E2%80%83%20of%E2%80%83%20young%E2%80%83%0Aphysically%E2%80%83active%E2%80%83men%E2%80%83and%E2%80%83women%EF%BC%BBJ%EF%BC%BD%EF%BC%8EJ%E2%80%83%20Physiol%E2%80%83%0APharmacol%EF%BC%8C2015%EF%BC%8C66%EF%BC%886%EF%BC%89%EF%BC%9A811-821%EF%BC%8E

62、%E2%80%83%20LEE%E2%80%83H%E2%80%83W%EF%BC%8CAHMAD%E2%80%83M%EF%BC%8CWANG%E2%80%83H%E2%80%83W%EF%BC%8Cet%E2%80%83al%EF%BC%8EEffects%E2%80%83%0Aof%E2%80%83exercise%E2%80%83training%E2%80%83on%E2%80%83brain-derived%E2%80%83neurotrophic%E2%80%83factor%E2%80%83%0Ain%E2%80%83%20skeletal%E2%80%83muscle%E2%80%83and%E2%80%83%20heart%E2%80%83of%E2%80%83%20rats%E2%80%83%20post%E2%80%83myocardial%E2%80%83%0Ainfarction%EF%BC%BBJ%EF%BC%BD%EF%BC%8EExp%E2%80%83Physiol%EF%BC%8C2017%EF%BC%8C102%EF%BC%883%EF%BC%89%EF%BC%9A%0A314-328%EF%BC%8E

63、LEE%E2%80%83H%E2%80%83W%EF%BC%8CAHMAD%E2%80%83M%EF%BC%8CWELDRICK%E2%80%83J%E2%80%83J%EF%BC%8Cet%E2%80%83al%EF%BC%8EEffects%E2%80%83%0Aof%E2%80%83%20exercise%E2%80%83training%E2%80%83%20and%E2%80%83%20TrkB%E2%80%83%20blockade%E2%80%83%20on%E2%80%83%20cardiac%E2%80%83%0Afunction%E2%80%83%20and%E2%80%83BDNF-TrkB%E2%80%83%20signaling%E2%80%83%20postmyocardial%E2%80%83%0Ainfarction%E2%80%83in%E2%80%83rats%EF%BC%BBJ%EF%BC%BD%EF%BC%8EAm%E2%80%83%20J%E2%80%83%20Physiol%E2%80%83%20Heart%E2%80%83%20Circ%E2%80%83%0APhysiol%EF%BC%8C2018%EF%BC%8C315%EF%BC%886%EF%BC%89%EF%BC%9AH1821-H1834%EF%BC%8E

64、%E2%80%83ZZEMBRON-LACNY%E2%80%83A%20%EF%BC%8C%20DZIUBEK%E2%80%83W%20%EF%BC%8C%0ARYNKIEWICZ%E2%80%83M%EF%BC%8Cet%E2%80%83al%EF%BC%8EPeripheral%E2%80%83%20brain-derived%E2%80%83%0Aneurotrophic%E2%80%83factor%E2%80%83is%E2%80%83%20related%E2%80%83to%E2%80%83%20cardiovascular%E2%80%83%20risk%E2%80%83%0Afactors%E2%80%83in%E2%80%83active%E2%80%83and%E2%80%83inactive%E2%80%83elderly%E2%80%83men%EF%BC%BBJ%EF%BC%BD%EF%BC%8EBraz%E2%80%83J%E2%80%83%0AMed%E2%80%83Biol%E2%80%83Res%EF%BC%8C2016%EF%BC%8C49%EF%BC%887%EF%BC%89%EF%BC%9Ae5253%EF%BC%8E

65、KAESS%E2%80%83B%E2%80%83M%EF%BC%8CPREIS%E2%80%83S%E2%80%83R%EF%BC%8CLIEB%E2%80%83W%EF%BC%8Cet%E2%80%83al%EF%BC%8E%0ACi%20rculating%E2%80%83%20b%20rain-de%20rived%E2%80%83%20neu%20rot%20rophic%E2%80%83%20facto%20r%E2%80%83%0Aconcentrations%E2%80%83and%E2%80%83the%E2%80%83risk%E2%80%83of%E2%80%83cardiovascular%E2%80%83disease%E2%80%83in%E2%80%83%0Athe%E2%80%83community%EF%BC%BBJ%EF%BC%BD%EF%BC%8EJ%E2%80%83Am%E2%80%83Heart%E2%80%83Assoc%EF%BC%8C2015%EF%BC%8C4%0A%EF%BC%883%EF%BC%89%EF%BC%9Ae001544%EF%BC%8E

66、TAKASHIO%E2%80%83S%EF%BC%8CSUGIYAMA%E2%80%83S%EF%BC%8CYAMAMURO%E2%80%83M%EF%BC%8Cet%E2%80%83%0Aal%EF%BC%8ESignificance%E2%80%83of%E2%80%83low%E2%80%83plasma%E2%80%83levels%E2%80%83of%E2%80%83brain-derived%E2%80%83%0Aneurotrophic%E2%80%83factor%E2%80%83in%E2%80%83patients%E2%80%83with%E2%80%83heart%E2%80%83failure%EF%BC%BBJ%EF%BC%BD%EF%BC%8E%0AAm%E2%80%83J%E2%80%83Cardiol%EF%BC%8C2015%EF%BC%8C116%EF%BC%882%EF%BC%89%EF%BC%9A243-249%EF%BC%8E

67、ROSS%E2%80%83M%EF%BC%8CKARGL%E2%80%83C%E2%80%83K%EF%BC%8CFERGUSON%E2%80%83R%EF%BC%8Cet%E2%80%83al%EF%BC%8E%0AExercise-induced%E2%80%83skeletal%E2%80%83muscle%E2%80%83angiogenesis%EF%BC%9A%0AImpact%E2%80%83of%E2%80%83age%EF%BC%8Csex%EF%BC%8Cangiocrines%E2%80%83and%E2%80%83cellular%E2%80%83mediators%0A%EF%BC%BBJ%EF%BC%BD%EF%BC%8EEur%E2%80%83J%E2%80%83Appl%E2%80%83Physiol%EF%BC%8C2023%EF%BC%8C123%EF%BC%887%EF%BC%89%EF%BC%9A1415-1432%EF%BC%8E

68、YAZDANI%E2%80%83F%EF%BC%8CSHAHIDI%E2%80%83F%EF%BC%8CKARIMI%E2%80%83P%EF%BC%8EThe%E2%80%83effect%E2%80%83%0Aof%E2%80%83%208%E2%80%83%20weeks%E2%80%83%20of%E2%80%83%20high-intensity%E2%80%83interval%E2%80%83training%E2%80%83%20and%E2%80%83%0Amoderate-intensity%E2%80%83%20continuous%E2%80%83training%E2%80%83%20on%E2%80%83%20cardiac%E2%80%83%0Aangiogenesis%E2%80%83factor%E2%80%83in%E2%80%83diabetic%E2%80%83male%E2%80%83rats%EF%BC%BBJ%EF%BC%BD%EF%BC%8EJ%E2%80%83%0APhysiol%E2%80%83Biochem%EF%BC%8C2020%EF%BC%8C76%EF%BC%882%EF%BC%89%EF%BC%9A291-299%EF%BC%8E

69、MORLAND%E2%80%83C%EF%BC%8CANDERSSON%E2%80%83K%E2%80%83A%EF%BC%8CHAUGEN%E2%80%83%0AO%E2%80%83P%EF%BC%8Cet%E2%80%83al%EF%BC%8EExercise%E2%80%83induces%E2%80%83%20cerebral%E2%80%83VEGF%E2%80%83%20and%E2%80%83%0Aangiogenesis%E2%80%83via%E2%80%83the%E2%80%83lactate%E2%80%83receptor%E2%80%83HCAR1%EF%BC%BBJ%EF%BC%BD%EF%BC%8E%0ANat%E2%80%83Commun%EF%BC%8C2017%EF%BC%888%EF%BC%89%EF%BC%9A15557%EF%BC%8E

70、%E2%80%83%20HANSEN%E2%80%83A%E2%80%83H%EF%BC%8CNIELSEN%E2%80%83J%E2%80%83J%EF%BC%8CSALTIN%E2%80%83B%EF%BC%8Cet%E2%80%83al%EF%BC%8E%0AExercise%E2%80%83training%E2%80%83normalizes%E2%80%83skeletal%E2%80%83muscle%E2%80%83vascular%E2%80%83%0Aendothelial%E2%80%83%20growth%E2%80%83%20factor%E2%80%83%20levels%E2%80%83%20in%E2%80%83%20patients%E2%80%83%20with%E2%80%83%0Aessential%E2%80%83hypertension%EF%BC%BBJ%EF%BC%BD%EF%BC%8EJ%E2%80%83Hypertens%EF%BC%8C2010%EF%BC%8C28%0A%EF%BC%886%EF%BC%89%EF%BC%9A1176-1185%EF%BC%8E

71、%E2%80%83%20WU%E2%80%83G%EF%BC%8CRANA%E2%80%83J%E2%80%83S%EF%BC%8CWYKRZYKOWSKA%E2%80%83J%EF%BC%8Cet%E2%80%83al%EF%BC%8E%0AExercise-induced%E2%80%83expression%E2%80%83of%E2%80%83VEGF%E2%80%83and%E2%80%83salvation%E2%80%83of%E2%80%83%0Amyocardium%E2%80%83in%E2%80%83the%E2%80%83early%E2%80%83stage%E2%80%83of%E2%80%83myocardial%E2%80%83infarction%0A%EF%BC%BBJ%EF%BC%BD%EF%BC%8EAm%E2%80%83J%E2%80%83Physiol%E2%80%83Heart%E2%80%83Circ%E2%80%83Physiol%EF%BC%8C2009%EF%BC%8C296%0A%EF%BC%882%EF%BC%89%EF%BC%9AH389-H395%EF%BC%8E

72、LIU%E2%80%83Y%EF%BC%8CWANG%E2%80%83Y%EF%BC%8CNI%E2%80%83Y%EF%BC%8Cet%E2%80%83al%EF%BC%8EGut%E2%80%83%20microbiome%E2%80%83%0Afermentation%E2%80%83%20determines%E2%80%83the%E2%80%83%20efficacy%E2%80%83%20of%E2%80%83%20exercise%E2%80%83for%E2%80%83%0Adiabetes%E2%80%83prevention%EF%BC%BBJ%EF%BC%BD%EF%BC%8ECell%E2%80%83Metab%EF%BC%8C2020%EF%BC%8C31%0A%EF%BC%881%EF%BC%89%EF%BC%9A77-91%EF%BC%8Ee5%EF%BC%8E

73、%E2%80%83%20ZHOU%E2%80%83Q%EF%BC%8CDENG%E2%80%83J%EF%BC%8CPAN%E2%80%83X%EF%BC%8Cet%E2%80%83al%EF%BC%8EGut%E2%80%83microbiome%E2%80%83%0Amediates%E2%80%83%20the%E2%80%83%20protective%E2%80%83%20effects%E2%80%83%20of%E2%80%83%20exercise%E2%80%83%20after%E2%80%83%0Amyocardial%E2%80%83infarction%EF%BC%BBJ%EF%BC%BD%EF%BC%8EMicrobiome%EF%BC%8C2022%EF%BC%8C10%0A%EF%BC%881%EF%BC%89%EF%BC%9A82%EF%BC%8E

74、MA%E2%80%83Y%EF%BC%8CLIU%E2%80%83H%EF%BC%8CWANG%E2%80%83Y%EF%BC%8Cet%E2%80%83al%EF%BC%8ERoles%E2%80%83of%E2%80%83physical%E2%80%83%0Aexercise-induced%E2%80%83miR-126%E2%80%83in%E2%80%83cardiovascular%E2%80%83health%E2%80%83of%E2%80%83%0Atype%E2%80%832%E2%80%83diabetes%EF%BC%BBJ%EF%BC%BD%EF%BC%8EDiabetol%E2%80%83Metab%E2%80%83Syndr%EF%BC%8C2022%EF%BC%8C%0A14%EF%BC%881%EF%BC%89%EF%BC%9A169%EF%BC%8E

75、PINCKARD%E2%80%83K%E2%80%83M%EF%BC%8CSHETTIGAR%E2%80%83V%E2%80%83K%EF%BC%8CWRIGHT%E2%80%83K%E2%80%83%0AR%EF%BC%8Cet%E2%80%83al%EF%BC%8EA%E2%80%83novel%E2%80%83endocrine%E2%80%83role%E2%80%83for%E2%80%83the%E2%80%83BAT-released%E2%80%83%0Alipokine%E2%80%8312%EF%BC%8C13-diHOME%E2%80%83to%E2%80%83mediate%E2%80%83cardiac%E2%80%83function%0A%EF%BC%BBJ%EF%BC%BD%EF%BC%8ECirculation%EF%BC%8C2021%EF%BC%8C143%EF%BC%882%EF%BC%89%EF%BC%9A145-159%EF%BC%8E

76、TAKAHASHI%E2%80%83H%EF%BC%8CALVES%E2%80%83C%E2%80%83R%E2%80%83R%EF%BC%8CSTANFORD%E2%80%83K%E2%80%83I%EF%BC%8C%0Aet%E2%80%83al%EF%BC%8ETGF-%CE%B22%E2%80%83is%E2%80%83an%E2%80%83exercise-induced%E2%80%83adipokine%E2%80%83that%E2%80%83%0Aregulates%E2%80%83glucose%E2%80%83and%E2%80%83fatty%E2%80%83acid%E2%80%83metabolism%EF%BC%BBJ%EF%BC%BD%EF%BC%8ENat%E2%80%83%0AMetab%EF%BC%8C2019%EF%BC%8C1%EF%BC%882%EF%BC%89%EF%BC%9A291-303%EF%BC%8E

1、国家自然科学基金面上项目（32171174）()