2. 湖南畜禽安全生产协同创新中心, 长沙 410128
2. Hunan Co-Innovation Center of Animal Production Safety, Changsha 410128, China
硒(selenium)元素主要以硒蛋白的形式在动物体内发挥重要的生理功能[1],已发现的硒蛋白有谷胱甘肽过氧化物酶(GSH-Px)、硫氧蛋白还原酶(TrxR)、甲状腺素脱碘酶(DIOs)和硒蛋白P(SelP)等25种,在动物体内参与抗氧化、免疫及激素的运输和调节等[2]。在动物生产中,常需要在饲粮中添加硒以满足机体对硒的需要[3],硒可提高动物生产性能,改善机体抗氧化、免疫等性能[4-5]。茶多酚(tea polyphenols)是茶叶中提取多酚物质的总称,包括黄烷醇类、黄酮类、酚酸类等,具有多种生物性能,如抗氧化、免疫、抑菌等,广泛用于食品药品等行业[6-7]。我国茶资源丰富,具有一系列的茶叶产业链,每年产生大量的茶叶副产物,具有提取茶多酚的大量资源,随着我国“全面禁抗”的到来,“替抗”饲料添加剂的开发逐渐成为研究热点,茶多酚作为绿色“替抗”饲料添加剂可提高动物的生产性能、产品品质、抗氧化性能等[8-9]。
茶多酚的性能可被其他物质影响,如蛋白质、多糖、矿物元素等[10],而硒是动物体必需矿物元素,对机体生长发育、抗氧化等方面具有十分重要的意义。硒与茶多酚具有多种相似的生物学供能,有研究显示,二者联用效应强于其单一物质[11]。对于硒与茶多酚的研究多见于富硒茶,其多酚类化合物多于常规茶,具有更强的抗氧化性能[12-13]。本文就硒与茶多酚的生物学功能、细胞信号转导调控机制及其在畜禽上的应用进行综述,为微量元素的精准营养、绿色饲料添加剂的研发及资源节约提供思路。
1 硒与茶多酚的生物学功能 1.1 抗氧化功能自由基是体内物质发生氧化还原产生的高活性分子,可破坏蛋白质、DNA,造成脂质过氧化。体内自由基过多可影响动物生产,甚至诱发疾病,造成死亡[14]。
硒主要通过GSH-Px、TrxR、DIOs等含硒酶发挥抗氧化作用。GSH-Px可催化过氧化氢(H2O2)及有机过氧化物与还原型谷胱甘肽(GSH)的反应,进而清除自由基[15],是主要的抗氧化酶;DIOs通过参与调节甲状腺激素的活性,调节机体抗氧化等功能;TrxR通过对底物硫氧还蛋白(thioredoxin,Trx)的催化还原参与过氧化氢清除、DNA合成以及氧化还原信号传导等过程,进一步发挥在氧化还原中的重要性能[16]。茶多酚可通过以下几种方式发挥抗氧化作用:1)络合金属离子以阻止金属离子催化自由基的形成,减少产生自由基[9];2)直接还原氢氧自由基(·OH)、超氧阴离子(O2-)、一氧化氮(NO)等自由基[17];3)调节抗氧化酶的活性,通过相应信号通路激活和保护含硒酶GSH-Px等体内抗氧化酶活性。研究发现,茶多酚中的没食子酸酯(epigallocatechin-3-gallate,EGCG)在硒充足的小鼠体内可诱导TrxR1基因的表达[18]。体外研究表明,茶多酚与硒的联用可发挥更佳的抗氧化功能[11]。
1.2 免疫功能硒与茶多酚均能介导免疫反应[19-20]。硒可刺激血液中白细胞的活化,增加抗体的合成,提高免疫细胞的活性,增强机体非特异性免疫、细胞免疫及体液免疫。当体内缺硒或硒蛋白表达被抑制时,多种组织炎症细胞因子的含量会受到影响,如缺硒会降低机体血清白细胞介素-1(IL-1)以及升高肿瘤坏死因子-α(TNF-α)含量[21],其中增加的TNF-α可以诱导核转录因子-κB(nuclear factor-κB,NF-κB)的活化并增加肝细胞中C反应蛋白(C-reactive protein,CRP)的分泌[19, 22]。茶多酚可结合免疫细胞上相应的受体,调节免疫细胞的黏附和炎症过程,调节免疫因子的分泌。研究发现,EGCG的免疫细胞受体有3类,67 ku层黏连蛋白受体(67 ku laminin receptor,67LR)、zeta链结合70 ku蛋白(zeta chain associated protein kinase 70 ku,ZAP-70)以及维甲酸诱导基因Ⅰ(retinoic acid induction gene Ⅰ,RIG-Ⅰ)[23]。Kuo等[24]研究表明,儿茶素可增强T细胞亚群的活性,提高白细胞介素-2(IL-2)等的分泌;增强免疫细胞活性;于娟等[25]研究表明,红茶多酚可以增强小鼠的T细胞、B细胞、自然杀伤细胞以及巨噬细胞的活性,增加抗体水平及免疫球蛋白含量[26]。硒与茶多酚促进细胞因子分泌,提高抗体水平,提高动物体免疫能力,但二者的互作及机理需要深入探究。
1.3 抗肿瘤功能硒和茶多酚可扰乱肿瘤细胞代谢、诱导细胞凋亡和抑制血管生成等,表现出抗肿瘤作用。硒可以通过抗氧化功能发挥抗肿瘤作用,此外,TrxR可调节细胞增殖和生存以及刺激肿瘤细胞,硒化合物可抑制肿瘤细胞端粒酶活性以及调控转录因子NF-κB的激活和失活,抑制肿瘤细胞[27]。茶多酚通过调节相关酶,如谷胱甘肽-S-转移酶(glutathione S-transferase,GST)抑制多种癌细胞生长,同时,硒也可以调节GST活性。硒与茶多酚对肿瘤细胞周期的抑制作用机理基本相同。茶多酚可以通过抑制B淋巴细胞瘤-2(B-cell lymphoma-2,bcl-2)基因的表达,诱导bcl-2相关蛋白(bcl-2 associated x protein,bax)基因的表达,从而诱导癌细胞发生编程性死亡,通过调控细胞周期蛋白依赖性激酶(cyclin-dependent kinases,CDK)、细胞周期蛋白E、CDK相互作用蛋白(CDK interacting proteins,CIPs)、激酶抑制蛋白(kinase inhibitory proteins,KIPs,如p27KIP1、P57KIP2)等细胞周期调控蛋白,诱导肿瘤细胞周期停滞在G1期进而抑制其生长[28-29]。硒与茶多酚均可以通过调节某些信号通路进而发挥抗肿瘤作用,通过活化丝裂原活化蛋白激酶(mitogen-activated protein kinases,MAPK)信号通路,抑制蛋白质丝氨酸-苏氨酸激酶(protein-serine-threonine kinase,AKT)细胞信号通路,激活线粒体凋亡通路,降低连环蛋白(catenin)水平,激活半胱氨酸-天冬氨酸蛋白酶(cystein-asparate protease,Caspase)-9、Caspase-3,上调视黄酸受体(retinoic acid receptors,RXR)表达,通过影响细胞间隙连接蛋白以及通讯功能,抑制肿瘤细胞生长,加快其凋亡,也可通过抑制肿瘤组织血管的形成产生抗肿瘤作用[30-31]。
2 硒与茶多酚对细胞内信号通路的调节 2.1 核因子E2相关因子2(nuclear factor erythroid 2-related factor 2,Nrf2)/抗氧化反应元件(antioxidant response element,ARE)信号通路硒与茶多酚可通过调节多种信号通路发挥作用。Nrf2/ARE通路是调节抗氧化机制的重要通路,在未被激活时,Nrf2与Kelch样环氧氯丙烷相关蛋白-1(Kelch-like ECH-associated protein 1,Keap1)结合,激活的Nrf2与Keap1解离,结合ARE,进而启动抗氧化等基因的转录,如GSH-Px、超氧化物歧化酶(SOD)、还原型烟酰胺腺嘌呤二核苷酸磷酸(NADPH)脱氢酶1等抗氧化酶等[32],研究发现,EGCG能诱导增加Nrf2的转录活性,提高与ARE的结合能力,继而上调含硒酶GSH-Px和TrxR的表达[33],当体内硒含量充足时,EGCG可直接诱导TrxR1基因的表达,而使Nrf2不被激活,当硒缺乏时,Nrf2被EGCG激活[18],同时,硒也可激活Nrf2信号通路[26]。
2.2 MAPK信号通路MAPK信号通路参与细胞的炎症、应激等过程,可被应激、炎性细胞等激活,调节细胞因子的产出。MAPK通路通过MAPK、MAPK激酶和MAPK激酶激酶三级级联调控。MAPK信号通路主要有3条途径:细胞外信号调节激酶(extracellular signal-regulated kinases,ERK)、p38丝裂原激活蛋白激酶(p38 mitogen-activated protein kinases,p38)、Jun氨基末端激酶(jun N-terminal kinases,JNK)[34]。硒可通过调控MAPK信号通路,下调TNF-α、白细胞介素-1β(IL-1β)和白细胞介素-6(IL-6)基因的表达,发挥抗炎作用[35]。Ma等[36]研究表明,茶多酚通过激活MAPK磷酸化缓解氧化应激对牛乳腺上皮细胞的损伤;Siddiqui等[37]发现,EGCG可通过激活ERK1/2,抑制人前列腺癌细胞活性。
2.3 NF-κB信号通路NF-κB通路可调控免疫和炎症反应,NF-κB蛋白家族通过与顺式调控区的DNA位点结合,形成异型和同型二聚体,未被激活的NF-κB二聚体通过与NF-κB抑制蛋白(inhibitor of NF-κB,IκB)结合在细胞中,当有刺激源时,IκB激酶复合物通过激活IκB进而激活NF-κB,进行靶基因的转录[38]。研究显示,硒与茶多酚能激活NF-κB通路,进而上调前列腺素E2(prostaglandin E2,PGE2)及血栓素2(thromboxane 2,TXA2)的表达,PGE2和TXA2是活化巨噬细胞的物质,以此调节免疫功能[22, 39]。
2.4 磷脂酰肌醇-3激酶(phosphatidylinositide 3-kinase,PI3K)-AKT信号通路胞内PI3K-AKT参与细胞营养物质的摄取等功能,PI3K是特殊的胞内脂质激酶,其活化可磷酸化质膜上的磷脂酰肌醇环,产生的3,4,5-三磷酸磷脂酰肌醇与AKT结合使其激活,继而磷酸化多种酶和转录因子对相应功能进行调节,如激活细胞色素C(cytochrome C,Cyt C),激活Caspase-3,致使细胞凋亡,也可以调节血管内皮细胞的生长[40-41]。硒和茶多酚可诱导PI3K-AKT,降低氧化应激对细胞的损伤以及抑制癌细胞生长[38, 42]。
茶多酚与硒通过调控多条信号通路参与机体的生长代谢发育。研究证实Nrf2是AKT的靶蛋白之一,MAPK也可激活Nrf2,硒与茶多酚可调控PI3K-AKT及MAPK信号通路,也可通过这2条信号通路进一步激活Nrf2[43-44],二者通过多条信号通路及通路间的相互作用,协同发挥抗氧化、免疫、抗肿瘤等功能,但硒与茶多酚发挥协同作用的机制研究尚需深入探究。
3 硒与茶多酚在动物生产中的应用 3.1 硒在动物生产中的应用硒作为饲料添加剂添加到饲粮中,可提高动物的生产性能、产品品质、抗氧化以及免疫功能,有机硒的效果通常大于无机硒。表 1为硒在动物生产中的部分研究成果[45-51]。
茶多酚可促进动物的生产性能,提高产品品质,如改善肉品质和蛋品质,提高抗氧化能力,调节动物体脂质代谢。茶多酚在动物生产中的部分研究成果见表 2[52-56]。
目前,硒与茶多酚的联合应用已有的研究显示,二者具有协同作用,配伍效果优于单一物质,但目前关于两者的协同作用机制探讨相对较少。李卫春[57]指出,硒与茶多酚组合可降低肉鸡胸肌红度(a*)、亮度(L*)值,提高黄度(b*)值,提高肉品质以及血浆中抗氧化酶(GSH-Px、SOD等)活性,降低氧化物黄嘌呤氧化酶(XOD)活性,且二者具有交互作用,饲粮中硒和茶多酚添加水平分别为0.15、200 mg/kg可提高鸡肉的抗氧化能力,延长鸡肉的货架期。Long等[58]研究酵母硒与茶多酚对于武昌鱼的影响发现,饲料中酵母硒与茶多酚添加水平分别为0.5、50 mg/kg可促进武昌鱼的生长、抗氧化性能及组织的恢复能力,且效果优于添加单一的酵母硒或茶多酚。龙萌等[59]研究显示,在饲料中添加酵母硒和茶多酚的影响下,团头鲂鱼的生长性能以及鱼体抗病原菌的感染力均显著提高,饵料系数显著下降,且二者联用具有互作效应,同时,两者均具有抗氧化作用,可诱导相关抗氧化基因表达,当两者联用时,对抗氧化功能具有协同作用,可提高铜锌超氧化物歧化酶(Cu/Zn- SOD)和过氧化氢酶(CAT)的基因表达,综合分析,对于团头鲂幼鱼,饲料中酵母硒和茶多酚适宜添加水平分别为0.5和50 mg/kg。
4 小结硒与茶多酚均具有抗氧化、免疫、抗肿瘤等作用,共同调节多条信号通路,其联合应用可改善动物体的生长性能、抗氧化功能、免疫功能等,在大部分的研究中,二者的联合应用效果优于单一物质,但关于硒与茶多酚联合发挥功能的许多机制尚未清晰,如二者发挥抗氧化功能与免疫功能的机制尚未完全清楚,其联合应用对动物肠道健康的影响尚未明了,对不同动物的配伍剂量尚需进一步研究。鉴于目前二者联用的良好效果,并随着研究的不断深入,硒与茶多酚的联合应用在动物生产中拥有广阔的前景。
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