Chinese Journal of Tissue Engineering Research ›› 2016, Vol. 20 ›› Issue (37): 5594-5601.doi: 10.3969/j.issn.2095-4344.2016.37.018
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Li Jing, Zhang Xiang, Wang Li-ping, Shi Lei, Mu Yu-zheng, Chang Qian-qian
Received:
2016-06-12
Online:
2016-09-09
Published:
2016-09-09
Contact:
Zhang Xiang, Master, Professor, Xinzhou Teachers University, Xinzhou 034000, Shanxi Province, China
About author:
Li Jing, Xinzhou Teachers University, Xinzhou 034000, Shanxi Province, China
Supported by:
the Shanxi Provincial Foundation for College Students Entrepreneurship Innovation Training in 2014; the Key Construction Disciplines Project of Xinzhou Teachers University in Shanxi Province, No. xk201403
CLC Number:
Li Jing, Zhang Xiang, Wang Li-ping, Shi Lei, Mu Yu-zheng, Chang Qian-qian. Effects of cytokines on skeletal muscle cells and pancreatic beta cells[J]. Chinese Journal of Tissue Engineering Research, 2016, 20(37): 5594-5601.
2.1 脂联素与骨骼肌细胞和胰岛β细胞 脂联素是最近发现的一种脂肪特异性蛋白质。脂联素对肌卫星细胞有激活作用,抑制成肌细胞的分化,促进骨骼肌细胞摄取、储存和利用葡萄糖,通过中枢和周围靶点提高胰岛素敏感性[5],这是脂肪细胞因子特殊而又共同的信号途径。脂联素是1995年运用分子生物学技术发现的由成熟脂肪细胞分泌的胶原样蛋白产物,由244个氨基酸组成的多肽。在分子结构中都包含一个氨基端成胶原区域和一个羧基端球状三聚区域。脂联素是一种在脂肪容量增大时分泌反而减少的白色脂肪组织蛋白,作为一种肥胖负调控因子,脂联素在体内的生理作用主要是增加胰岛素敏感性,促进糖脂代谢。脂联素的生物学功能表现在胰岛素抵抗增敏、代谢调节、抗炎、血管保护、抗心肌缺血和心肌保护功能。脂联素通过抑制单核细胞黏附,减低其细胞周期活性,降低血管壁脂蛋白的沉积,而抑制动脉硬化过程。近年来研究发现,脂联素在血细胞形成和免疫反应中发挥负调控作用[6-9],另外,脂联素单核苷酸多态性也与脂联素的生成和分泌的减少有关,这可能也是导致胰岛素抵抗及2型糖尿病发生的原因。在动物和人类的实验中证实其具有增加脂肪酸氧化、提高葡萄糖摄取量的作用[10-12]。脂联素参与调节糖代谢的作用可能与其增强骨骼肌和肝脏的脂肪酸氧化,促进骨骼肌对葡萄糖的摄取,抑制肝糖输出和糖异生有关。 胰岛素抵抗是由免疫系统介导的慢性、非特异性炎症过程。脂联素具有抗炎、促炎的双重作用,其与炎症因子的相互作用可能是脂联素增强胰岛素敏感性的机制之一,也可能是胰岛素抵抗时低脂联素血症的原因。以往研究证实脂联素有胰岛素增敏作用,能拮抗胰岛素抵抗[13]。脂联素与空腹胰岛素浓度负相关,与胰岛素敏感指数正相关,吡格列酮治疗可使血浆脂联素水平明显增加,游离脂肪酸水平降低,改善胰岛素抵抗,使胰岛素敏感性增加[14]。马慧娟等[15]研究结果说明非糖尿病一级亲属组已经存在胰岛素抵抗,并且血清游离脂肪酸水平升高,脂联素水平降低;血清游离脂肪酸、脂联素与胰岛素抵抗密切相关,可作为评估胰岛素抵抗的较好指标。由此可见对脂联素的研究将为探讨胰岛素抵抗和2型糖尿病的发病机制及治疗方案提供新的线索。 脂联素是目前发现的惟一对人体有保护作用的脂源性激素,脂联素的胰岛素增敏作用是通过增加骨骼肌脂肪酸氧化以及抑制肝糖异生实现的。潘云龙等[16]通过临床研究发现在糖尿病患者中血浆脂联素与体质量指数、胰岛素抵抗指数呈负相关,与高密度脂蛋白胆固醇呈正相关,与空腹血糖、糖化血红蛋白、三酰甘油、总胆固醇及胰岛β细胞功能指数相关性不明显,其浓度下降与糖尿病及其大血管病变密切相关。 脂联素水平受到多种因素调节,其在健康人体血清中含量丰富,在肥胖及2型糖尿病患者中含量降低。运动训练能够改善糖尿病或代谢综合征患者的胰岛素抵抗,提高脂联素的水平;运动训练改善胰岛素敏感性是由于与运动训练改善了血浆中脂联素的浓度有关[17-19]。因此,提高体质水平的运动能够提高脂联素浓度。寻找一个有效干预脂联素水平的运动处方是一个值得关注的问题。李世成等[20]观察中等强度运动对2型糖尿病肥胖患者血浆脂联素水平和胰岛素敏感性及体质量的影响,发现中等强度运动可改善2型糖尿病肥胖患者胰岛素敏感性,有助于血糖稳定,推测该种作用可能与血浆脂联素水平增高有关。牛燕媚等[21]研究证实有氧运动通过增加骨骼肌中腺苷酸活化蛋白激酶蛋白表达,从而促进骨骼肌细胞的糖脂代谢,达到改善高脂饮食诱发的胰岛素抵抗症状的效果。 2.2 瘦素与骨骼肌细胞和胰岛β细胞 瘦素是近期发现的一种存在于啮齿动物及人体的主要调节食物摄取和能量消耗的激素。它在1994年肥胖基因分离后被发现,是肥胖基因编码的蛋白产物,并由脂肪组织分泌进入血液循环,瘦素以游离及结合的形式存在于血浆中。瘦素在成肌细胞中发挥重要作用,可以促进骨骼肌成肌细胞的增殖,并呈剂量依赖性,但其抑制成肌细胞分化,对骨骼肌细胞生长和发育有重要作用。 过去认为脂肪细胞在外界刺激下被动地储存和释放能量。瘦素的发现使人们认识到脂肪组织是主动参与能量代谢调节的组织。现在已经共同认识到高血压、肥胖、糖尿病、脂代谢紊乱、高半胱氨酸血症、心脏病、吸烟是脑梗死发病的高危因素,胰岛素抵抗也日益受到重视,瘦素参与了上述疾病的发生发展[22-24]。瘦素是由肥胖基因编码的一种多肽激素,具有调节能量代谢和抑制食欲的作用,其调节作用由瘦素受体所介导。瘦素受体属于Ⅰ类细胞因子受体家族,瘦素具有广泛的生物学特性,作用于下丘脑的体质量调节中枢,研究表明瘦素与胰岛素抵抗有关[25-26],高瘦素血症与胰岛素抵抗显著相关,高瘦素血症与葡萄糖耐量异常有关,而妊娠糖尿病患者存在高瘦素血症,因此妊娠糖尿病的胰岛素抵抗与高瘦素水平有关。 瘦素在胰岛素抵抗中的作用备受关注,多数研究表明,血清瘦素水平与肥胖呈显著正相关;瘦素能够控制体质量,纠正糖、脂代谢紊乱,增加胰岛素敏感性。高丽萍等[27]研究结果显示糖尿病肥胖患者的胰岛素抵抗指数明显增高,随着瘦素的降低,胰岛素抵抗指数呈上升趋势,说明胰岛素抵抗指数与瘦素间呈显著负相关,瘦素有望成为反映胰岛素抵抗的新指标。瘦素与胰岛素互为负反馈关系,二者在产生机制上可能互为因果,并且相互作用,共同参与了多种疾病的发生发展。阚卫军等[28]以对数胰岛素抵抗指数为因变量,以体质量指数、瘦素、三酰甘油、总胆固醇、高密度脂蛋白为自变量,进行多元线性逐步回归分析,显示三酰甘油和瘦素进入回归方程,当三酰甘油不变的情况下,对数胰岛素抵抗指数仍与瘦素显著正相关。以对数胰岛β细胞功能指数为因变量同上做多元线性逐步回归分析,仅瘦素进入回归方程,对数胰岛β细胞功能指数与瘦素呈负相关。结果表明2型糖尿病的一级亲属与正常对照组相比存在瘦素高表达,且瘦素水平与胰岛素抵抗指数呈正相关,而与胰岛β细胞功能指数呈负相关,提示瘦素与胰岛素抵抗密切相关。 血清瘦素调节代谢与多种因素有关,瘦素的最主要作用是对能量平衡的影响,运动通过改变系统能量流动和平衡影响瘦素水平,正常生理状态下运动降低血清瘦素水平是机体为维持自身体质量稳定和能量平衡的一种适应性反应[29-30]。赵美鲁等[31]研究表明长期有氧运动可有效降低体脂,缓解瘦素抵抗,但其产生机制并不太清楚。最新研究表明运动与瘦素信号传导通路有关,运动作为刺激手段通过不同途径影响血清瘦素水平。 2.3 内脂素与骨骼肌细胞和胰岛β细胞 内脂素是一种多效细胞因子,广泛存在于骨骼肌、肝脏、骨髓中,具有调节炎症反应和免疫功能的作用,对骨骼肌细胞应激反应和信号转导有重要作用,促进骨骼肌的葡萄糖吸收和葡萄糖代谢,还能促进炎性反应因子表达及诱导血管生成[6]。内脂素是一种主要由内脏脂肪细胞分泌的脂肪细胞因子,它同时也在骨骼肌、肝脏、骨髓和淋巴细胞等组织中被发现,并被定义为前B细胞集落增强因子。它具有类胰岛素样作用,能降低血糖和促进脂肪组织的分化与合成。内脂素还可以调节血管平滑肌的成熟和影响胰岛细胞的胰岛素分泌,亦具有调节炎症反应和免疫功能的作用[32-34]。随着研究的发展,人们对内脂素的结构特性、分布、表达调控及其生物学功能有了更加深入的认识。内脂素在葡萄糖和脂质代谢中发挥着重要作用,对内脂素的研究为探讨胰岛素抵抗和妊娠糖尿病的发病机制提供新的思路和方法[35-36]。 国内外众多研究均揭示了内脂素在2型糖尿病和肥胖中发挥重要作用,而内脂素与胰岛素抵抗密切相关。王玉环等[37]通过对2型糖尿病患者、正常健康人群中血浆内脂素、脂联素水平及其他临床指标的观察,分析血浆内脂素与一般临床资料及其他代谢和生化指标的关系;同时研究血浆内脂素与血糖、胰岛素抵抗及冠状动脉粥样硬化等之间的关系,研究显示血清内脂素可能参与了血糖、血脂代谢的调节,且与胰岛素抵抗密切相关,血清内脂素可能是2型糖尿病的危险因素之一。 内皮素与空腹胰岛素、稳态模型评估的胰岛素抵抗指数、稳态模型评估的胰岛β细胞功能、内脂素呈正相关,内脂素与内皮素、空腹胰岛素、胰岛素抵抗指数呈正相关,胰岛素抵抗指数是血浆内脂素的独立相关因素,内脂素与内皮素、胰岛索抵抗有关[38],内皮素与2型糖尿病及胰岛素抵抗的发病机制密切相关。运动可以降低血浆内脂素水平,提高脂肪组织中内脂素mRNA的表达。内脂素可能是运动改善机体代谢状况、防治代谢疾病的信号分子[39-41]。 2.4 其他细胞因子 肿瘤坏死因子α是一种多功能的细胞因子,在不同的组织中发挥其不同的生物学作用。致炎因子肿瘤坏死因子α抑制成肌细胞分化和增强细胞凋亡。肿瘤坏死因子α在肥胖和胰岛素抵抗过程中起重要作用。肥胖和胰岛素抵抗时肿瘤坏死因子α表达增强,可能通过抑制各种基因转录,干扰受体信号转导等机制发挥作用。肿瘤坏死因子α参与了胰岛素抵抗的发生,肥胖者机体过度表达了肿瘤坏死因子α,并与胰岛素抵抗程度呈正相关[42-44]。慢性疾病或遗传性或退化性肌肉病中肿瘤坏死因子α水平升高,并可能导致肌肉萎缩,与此相反,胰岛素样生长因子1对骨骼肌生长有促进作用。肿瘤坏死因子α和胰岛素样生长因子1调节骨骼肌细胞体外分化时miRNA表达,miRNA通过靶向调节轴突导向相关基因、MAPK路径、神经营养因子路径对影响骨骼肌成肌能力的肿瘤坏死因子α产生负作用,成肌细胞分化和细胞因子作用有重要关系。低浓度肿瘤坏死因子α降低肌原纤维蛋白的含量,相反高浓度(100 U/mL以上)增加C2C12肌管肌原纤维蛋白质含量。骨骼肌细胞蛋白降解速率减少14%,蛋白质合成速度增加40%,肿瘤坏死因子α对肌管蛋白质合成的影响与胰岛素对肌管蛋白质合成的影响相似,都通过影响骨骼肌细胞蛋白质合成中MAPKs细胞内信号转导路径发挥作用。 吴静等[45]研究发现血浆肿瘤坏死因子α水平与胰岛素敏感性指数呈负相关,肿瘤坏死因子α参与了胰岛素敏感性指数的变异,血浆总胆固醇、体质量指数、肿瘤坏死因子α对胰岛素敏感性指数总的影响程度为39%。同样也有研究证实肿瘤坏死因子α是影响糖尿病患者胰岛素敏感性指数最为显著的因素[46],与糖尿病患者胰岛素敏感性指数呈独立负相关,瘦素与肿瘤坏死因子α呈显著正相关,且两者均与胰岛素、胰岛素敏感性密切相关,肿瘤坏死因子α可能在2型糖尿病患者胰岛素抵抗中起着重要作用。骨骼肌卫星细胞增殖、分化并融合为新的肌管可以促进肌肉修复和生长,肿瘤坏死因子α水平升高对骨骼肌卫星细胞增殖、分化有抑制作用,在骨骼肌病理过程中有重要影响,肿瘤坏死因子α增加了NF-κB活性,减少PPARγ表达,导致受损肌管形成和活化,二十碳五烯酸(EPA)可以上调PPARγ表达,抑制肿瘤坏死因子α介导的白细胞介素6的转录和分泌,在骨骼肌损伤再生中起潜在的治疗作用。 抵抗素是由脂肪组织特异表达分泌的细胞因子,啮齿类动物实验和体外实验研究显示抵抗素可以诱导机体的胰岛素抵抗,引起糖脂代谢紊乱。多种因素可影响抵抗素的表达水平。抵抗素的表达受胰岛素增敏剂、营养状态及抵抗素拮抗剂等调节[47-50]。 郝奉民等[51]进行相关分析显示,糖尿病肾病患者血清抵抗素水平与空腹胰岛素、空腹血糖、胰岛素抵抗指数均呈显著正相关,并证实抵抗素与糖尿病肾病患者的胰岛素抵抗密切相关,通过检测2型糖尿病患者的血清抵抗素变化,在一定程度上可反映其肾脏病变。研究发现2型糖尿病患者血清抵抗素水平明显升高,血清抵抗素水平与胰岛素抵抗指数、体质量指数、三酰甘油密切相关[52-53]。胆固醇为影响抵抗素水平最明显的因素,其次为葡萄糖,说明2型糖尿病患者血清抵抗素水平升高,且与肥胖及胰岛素抵抗程度相关,因而抵抗素可能在2型糖尿病的发生发展中起重要作用。"
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