动物营养学报 >

2024 , Vol. 36 >Issue 8: 4761 - 4772

DOI: https://doi.org/10.12418/CJAN2024.408

综述

MicroRNA在畜禽生产中的研究进展

  • 王芳 ,
  • 吴康乐 ,
  • 姚康 , * ,
  • 印遇龙
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  • 中国科学院亚热带农业生态研究所,亚热带农业生态过程重点实验室,湖南省畜禽健康养殖工程技术中心,长沙 410125
*姚 康,研究员,博士生导师,E-mail:

王 芳(1998—),女,湖南郴州人,博士研究生,从事动物生理与代谢调控研究。E-mail:

收稿日期: 2024-02-26

  网络出版日期: 2024-08-12

基金资助

国家重点研发计划项目(2023YFD1301005)

泰山产业领军人才工程项目(tscy20190121)

吉林省与中国科学院科技合作高技术产业化专项资金项目(2022SYHZ0017)

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Research Progress of MicroRNA in Livestock and Poultry Production

  • WANG Fang ,
  • WU Kangle ,
  • YAO Kang , * ,
  • YIN Yulong
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  • Hunan Engineering Technology Center for Healthy Animal Husbandry, Key Laboratory of Agro-Ecological Processes in Subtropical Region, Subtropical Agro-Ecology Institute, Chinese Academy of Sciences, Changsha 410125, China
*professor, E-mail:

Received date: 2024-02-26

  Online published: 2024-08-12

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摘要

MicroRNA(miRNA)是一类短小的非编码RNA分子,在细胞调控和生物学过程中的多个方面引起广泛关注。近年来,研究发现miRNA在动物生理过程中扮演着重要的调控角色,尤其是与肌肉发育、脂肪代谢、肉品质、激素分泌和肠道健康等领域的关系。本文将深入探讨miRNA在这些生理过程中的作用及其潜在机制,以期为畜牧业和食品工业提供有益的科学依据。

关键词： MicroRNA; 肌肉发育; 脂肪代谢; 肉品质; 肠道健康

本文引用格式

王芳 , 吴康乐 , 姚康 , 印遇龙 . MicroRNA在畜禽生产中的研究进展[J]. 动物营养学报, 2024 , 36(8) : 4761 -4772 . DOI: 10.12418/CJAN2024.408

Abstract

MicroRNA (miRNA) is a class of short non-coding RNA molecules, which has attracted wide attention in many aspects of cell regulation and biological processes. In recent years, researchers have found that miRNA play important regulatory roles in animal physiological processes, especially in the fields of muscle development, fat metabolism, meat quality, hormone secretion and intestinal health. In this review, the roles of miRNA in these physiological processes and potential mechanisms will be thoroughly reviewed in order to provide useful scientific evidence for animal husbandry and food industry.

Key words： microRNA; muscle development; lipid metabolism; meat quality; intestinal health

MicroRNA(miRNA)是一类短链非编码RNA,通常含有19~25 nt的核酸碱基,广泛存在于动植物细胞内[1]。这类miRNA通常与蛋白质编码的mRNA序列中的3'非翻译区(3'UTR)互补,从而对转录后翻译水平进行调节。在细胞核内,miRNA可以与目标基因的启动子区域结合,影响靶基因表达[2]。首次发现miRNA可以追溯到1993年,在秀丽隐杆线虫中,LIN-14与转录因子LIN-4的mRNA的3'UTR序列结合,并抑制LIN-4的mRNA翻译[3]。miRNA通过靶基因,参与细胞分裂、分化、增殖、代谢等多种生物学过程。
哺乳动物体内约有60%的蛋白质编码基因能够被miRNA调节,miRNA最常研究的功能是在细胞质内沉默目标基因mRNA的表达[4]。近年来,miRNA在畜牧学研究领域中的研究不断增加,涉及肌肉发育、脂肪代谢、肉品质、激素分泌和肠道健康等方面[5-6]。因此,本文旨在综述miRNA在这些生理过程中的作用及其潜在机制,为解析miRNA在动物中的分子作用及调控机制提供理论依据。

1 miRNA与肌肉发育

1.1 miRNA对肌细胞增殖的调节

肌肉发育过程中,miRNA主要通过调控肌细胞的增殖和分化来发挥作用[7]。这一过程始于骨骼肌卫星细胞的激活,激活后的细胞进入增殖分化阶段,进而分化为肌细胞的一类成肌细胞[8]。骨骼肌中的成肌细胞作为成肌前体细胞,具有自我更新和生成新肌纤维的能力。它们沿着特定方向增殖生长,并通过融合形成肌管,最终分化成肌纤维[9]。研究表明,miRNA在肌细胞分化的不同阶段存在差异表达,这表明miRNA在肌细胞分化的各个阶段都可能有特定的调控作用[10⇓⇓⇓-14]。
在骨骼肌卫星细胞的调控中,miR-145-5p能靶向胰岛素样生长因子1受体(insulin-like growth factor 1 receptor, IGF1R)的3'UTR,调控IGF1R mRNA和蛋白的相对表达水平,并通过影响蛋白激酶B(protein kinase B, AKT)通路,进而抑制骨骼肌卫星细胞的增殖和分化[15]。另外,miR-143通过靶向胰岛素样生长因子结合蛋白5(insulin-like growth factor binding protein 5, IGFBP5)的表达促进骨骼肌卫星细胞分化[16]。miR-128在牛骨骼肌卫星细胞分化过程中上调表达,其过表达可靶向特异性蛋白 1(specificity protein 1, Sp1)基因抑制牛骨骼肌卫星细胞增殖和分化[17]。Song等[18]报道,miRNA Y-65通过靶向IGF1R激活AKT和细胞外信号调节激酶(extracellular signal-regulated kinases,ERK)通路,从而抑制猪肌肉卫星细胞的增殖和细胞周期进程。Zhu等[19]发现,miR-128-3p通过靶向法尼基二磷酸合酶(farnesyl diphosphate synthase, FDPS)促进鸡肌内脂肪细胞增殖并抑制其分化。此外,在卫星细胞分化过程中,miR-1和miR-206的表达量较高,是肌肉特异性miRNA[20]。研究指出,一方面,miR-206及miR-1调控的靶基因为配对盒基因7(paired box 7, PAX7)、miR-206及miR-1可结合PAX7 mRNA的3'UTR,通过抑制PAX7的表达来达到限制卫星细胞的增殖潜能,进而促进卫星细胞的分化作用[21]。另一方面,针对不同品种猪的肌纤维类型和肌肉发育程度的差异特征,Chen等[22]对荣昌猪断奶和屠宰时期的背最长肌组织进行了miRNA测序,发现miR-127在屠宰期显著下调,而在断奶期显著上调,表明miR-127可以抑制猪背最长肌卫星细胞的增殖和成肌。同时,研究证实miR-127通过靶向抑制铁调节蛋白2(aconitase 2, ACO2)基因表达来促进肌细胞增殖[23]。
如图1-B所示,在成肌细胞的增殖分化成为肌管过程中,miR-452靶向血管生成素1(angiopoietins 1, ANGPT1)促进成肌细胞增殖,抑制成肌分化[24]。miR-378可以调控生长因子受体结合蛋白2(growth factor receptor bound protein 2, GRB2)的3'UTR,其促进成肌细胞分化的机制可能是通过下调该基因间接抑制丝裂原活化蛋白激酶(mitogen-activated protein kinase, MAPK)通路,进而影响成肌细胞的增殖,促进其分化[25]。Sirt1可通过肌肉生长抑制素(myostatin, MSTN)信号通路促进成肌细胞的增殖,而miR-196b-5p通过靶向沉默调节蛋白1(sirtuin 1, SIRT1)促进成肌细胞分化[26]。在牛骨骼肌卫星细胞体外成肌诱导分化模型中,miR-128可以直接调控Sp1基因,miR-128过表达可降低Sp1蛋白水平,导致肌卫星细胞增殖过程受到抑制,导致生肌决定因子(myogenic determination gene, MyoD)表达量降低,从而使成肌分化过程受到抑制[27]。gga-miR-223在家禽骨骼肌生长发育中的功能是靶向胰岛素样生长因子2(insulin-like growth factor 2, IGF2)抑制成肌细胞的增殖,靶向锌指增强子结合蛋白1(zinc-finger-enhancer binding protein 1, ZEB1)促进成肌细胞分化[28]。综上所述,这些研究表明,miRNA在肌肉发育的不同阶段发挥着双重功能,意味着在肌细胞分化的不同阶段可能存在不同的调控靶点(图1-A和图1-B)。因此,miRNA在调节肌细胞增殖和分化方面具有不可忽视的作用,但是其具体的作用机制有待进一步研究。
图1 MicroRNA影响肌肉发育过程

A: 卫星细胞增殖和分化proliferation and differentiation of satellite cells;B: 成肌细胞增殖和分化myoblasts proliferated and differentiated;C: 肌纤维类型转换 muscle fiber type conversion。
PAX7: 配对盒基因7 paired box 7; ACO2: 铁调节蛋白2 aconitase 2; FDPS: 法尼基二磷酸合酶 farnesyl diphosphate synthase; IGFBP5: 胰岛素样生长因子结合蛋白5 insulin-like growth factor binding protein 5; IGF1R: 胰岛素样生长因子1受体 insulin-like growth factor 1 receptor; Sp1: 特异性蛋白 1 specificity protein 1; IGF2: 胰岛素样生长因子2 insulin-like growth factor 2; ZEB1: 锌指增强子结合蛋白1 zinc-finger-enhancer binding protein 1; SIRT1: 沉默调节蛋白1 sirtuin 1; ANGPT1: 血管生成素1 angiopoietin 1; SOX6: SRY-box转录因子6 SRY-box transcription factor 6; HDAC3: 组蛋白脱乙酰酶3 histone deacetylase 3; MYOZ1: 肌增生素 1 myozenin 1; UCP3: 解偶联蛋白3 uncoupling protein 3; ATP2a2: 钙离子转运ATP酶A2 ATPase sarcoplasmic/endoplasmic reticulum Ca2+ transporting 2。

Fig.1 MicroRNA affects process of muscle development

1.2 miRNA对肌纤维类型的调节

成年哺乳动物骨骼肌肌纤维根据肌球蛋白重链(myosin heavy chain, MyHC)的多态性表达差异可分为MyHCⅠ、Ⅱa、Ⅱx和Ⅱb 4种类型,代谢上分别与慢速氧化型、快速氧化型、快速酵解型和中间类型相对应[29]。Wang等[30]研究发现,过表达miR-449-5p能够靶向作用SRY-box转录因子6(SRY-box transcription factor 6, SOX6)促进MyHC Ⅰ和MyHC Ⅱa mRNA表达,表明miR-449-5p在促进氧化型肌纤维形成方面有显著的调节作用。张勇[31]研究发现,miR-378b-3p通过靶向抑制组蛋白脱乙酰酶3(histone deacetylase 3, HDAC3)导致肌细胞特异性增强因子2D(myocyte enhancer factor 2D, MEF2D)去乙酰化,从而抑制过氧化物酶体增殖受体γ辅激活因子-1α(peroxisome proliferator-activated receptor gamma coactivator-1 alpha, PGC-1α)对MEF2D的转录激活依赖,进而促进慢肌或氧化型肌纤维的形成并增加线粒体的生物合成。刘慧[32]研究表明,miR-432对肌细胞增殖及肌纤维类型转换可产生影响,敲降miR-432的表达可通过促进其靶基因肌增生素1(myozenin 1, MYOZ1)表达导致快肌生成量增多。Zhang等[33]研究发现,miR-152可通过靶向调节解偶联蛋白3(uncoupling protein 3, UCP3)基因,调节线粒体的质子梯度和ATP产生,从而影响能量代谢的效率,促进猪慢肌纤维和骨骼肌形成。另外,miR-151-3p通过靶向调节钙离子转运ATP酶A2(ATPase sarcoplasmic/endoplasmic reticulum Ca2+ transporting 2, ATP2a2)基因,抑制慢肌纤维标志基因肌球蛋白重链β(myosin heavy chain β, MHC β)表达,进而调节慢肌纤维的形成[34]。
综上所述,miRNA通过靶向靶标基因,进而调节肌球蛋白重链组成、线粒体的生物合成、能量代谢的效率等途径使肌纤维类型的形成和转换发生改变,这些研究揭示了miRNA在肌肉纤维调节中的多样作用,有望为肌纤维可变性提供新的视角和理论依据(图1-C)。

2 miRNA与肉品质

2.1 miRNA对肌内脂肪的调节

多种miRNA在调控动物脂肪沉积中具有潜在作用,其中包括通过调控Kruppel样转录因子(Kruppel-like factors, KLF)家族、过氧化物酶体增殖物激活受体γ(peroxisome proliferator-activated receptor gamma,PPARγ)等靶向调控路径[35-36]。
转录因子KLF家族成员(包括KLF3、KLF4、KLF9及KLF13)被认为是影响猪肌内脂肪沉积的关键因素[36]。Peng等[37]研究发现,在猪肌内脂肪细胞中,miR-429可以降低成脂标志基因表达及减少甘油三酯的积累,在此过程中,KLF9和p27是miR-429在这些过程中的有效靶点。miR-425-5p可靶向抑制KLF13,进而对猪肌内脂肪细胞的形成产生抑制作用[38]。Du等[39]研究发现,在猪肌内前脂肪细胞分化过程中,KLF13的3'UTR和超长链脂肪酸延伸酶6(elongase of verylong chain fatty acids 6, ELOVL6)的3'UTR均存在miR-125a-5p的靶基因位点,表明miR-125a-5p可能是猪肌内脂肪生成和猪肌内脂肪脂肪酸组成的一个新的调节因子。谭镇东[40]研究表明,miR-125a-5p通过抑制肌内前脂肪细胞脂肪合成基因表达,同时,增强猪肌内脂肪前体细胞的线粒体生成和能量代谢能力,加速脂肪细胞的能量支出,抑制脂质沉积。miR-206通过靶向类固醇急性调节蛋白相关脂质转运体7(StAR related lipid transfer 7, STARD7)来调控肌内脂肪细胞增殖,同时可抑制KLF4的表达,抑制肌内脂肪形成[41]。此外,PPARγ作为影响脂肪细胞分化和脂质代谢的关键基因,已被证实是多种miRNAs的靶向结合因子[42⇓-44]。马雪瑶[45]研究表明,Bta-miR-130a/b直接靶定PPARγ和细胞色素P450家族成员2 U1(cytochrome P450 family 2 subfamily U member 1, CYP2U1)调控秦川牛前体脂肪细胞的成脂分化,过表达Bta-miR-130a/b可以通过抑制PPARG mRNA及蛋白表达或促进CYP2U1 mRNA和蛋白的表达来降低甘油三酯的含量。
与此同时,miRNA可通过调节其他脂质代谢通路及基因,对动物肌内脂肪合成产生影响。miR-146a-5p通过靶向Smad同源物4(Smad homolog 4, SMAD4)(细胞周期基因)介导转化生长因子-β(transforming growth factor-β, TGF-β)信号通路,从而抑制猪肌内前脂肪细胞增殖;同时,靶向TRAF6介导AKT/雷帕霉素靶蛋白(mechanistic target of rapamycin, mTOR)信号通路来抑制肌内前体脂肪细胞分化[46]。miR-148a-3p通过抑制AMP活化蛋白激酶(AMP-activated protein kinase, AMPK)/乙酰辅酶A羧化酶(acetyl-CoA carboxylase, ACC)/肉毒碱棕榈酰基转移酶1C(carnitine palmitoyltransferase 1C, CPT1C)信号通路促进肌内前脂肪细胞分化,表明miR-148a-3p可作为贵州丛江象猪肌内脂肪形成的调控因子[47]。此外,Zhu等[19]研究报道,miR-128-3p靶向FDPS的3'UTR,介导脂代谢相关通路,如MAPK和TGF-β信号通路,这对于抑制肌内脂肪细胞的分化起到了关键作用。
综上所述,这些研究表明,miRNA调控畜禽肌内脂肪沉积的机制主要通过对肌内脂肪细胞增殖分化、脂质代谢通路及脂质代谢中关键基因等的调节,揭示了miRNA对于肌内脂肪沉积调节的差异性,为改善畜禽肉类风味品质提供了理论依据。

2.2 miRNA对其他肉品质指标的调节

miRNA对肌肉乳酸、滴水损失、肌肉脂肪酸组成、肉色及肌内脂肪含量和硫胺素含量等指标具有潜在调控作用。研究表明,miR-152通过靶向调控猪体内编码糖酵解关键限速酶丙酮酸激酶(propionone acid kinase,PKM)的基因来调控乳酸生成,从而影响猪肉品质[48]。Wei等[49]通过表达模式分析,初步证实了miRNA-499和miRNA-22是影响滴水损失性状的潜在候选基因。Wang等[50]研究发现,miR-22通过靶向ELOVL6调节骨骼肌细胞内钙离子(Ca2+)水平,而Ca2+水平可导致肉色变化。张彩荣等[51]通过对miRNA在牛背最长肌中表达及其与肉质形状相关性研究发现,miR-320a、miR-152、miR-143、miR-125a表达量增加促进肌肉组织脂肪沉积,其中miR-152、miR-143、miR-125a通过改变脂肪酸组成从而增加脂肪含量;miR-320a、miR-152、miR-143表达量增加会显著降低背最长肌失水率,有助于提高肉出品率;miR-320a及miR-143表达量增加可使肉色变深,是影响肉类色泽的重要遗传因素。杨雪梅等[52]研究表明,miR-1、miR-27a、miR-369和miR-378在猪背最长肌中的表达水平与肌内脂肪含量和硫胺素含量呈正相关,这表明miRNA对于猪肉品质具有潜在调控作用。综上所述,miRNA在肉质调控方面具有调控作用,通过miRNA的功能研究,可促进肉类品质提升,但miRNA对于肉品质的研究目前仅局限于miRNA对肉质及其风味品质的关联研究,其潜在的作用途径及发生机制并不明晰。

3 miRNA与脂代谢

3.1 miRNA对脂滴形成的调节

脂肪细胞中脂滴的正常形态和大小与脂肪细胞分化和脂肪组织的发育的调节具有相关性[53]。在脂肪细胞分化过程中,miR-215-5p直接靶作用于核受体辅激活蛋白3(nuclear receptor coactivator 3, NCOA3)抑制固始鸡腹部前脂肪细胞的增殖和分化,减少脂肪细胞内脂滴生成和甘油三酯含量[54]。miR-181a和miR-181d-5p可以分别靶向结合甘油-3-磷酸脱氢酶2(glycerol-3-phosphate dehydrogenase 2, GPD2)和环腺苷酸反应元件结合蛋白1(cAMP responsive element binding protein 1, CREB1)基因3'UTR序列,以促进未分化脂肪细胞脂滴数目增加,表明两者对于猪前体脂肪细胞成脂分化具有促进作用[55]。Shan等[56]报道,miR-218-5p通过抑制长链脂酰辅酶A合成酶1(long-chain acyl-CoA synthetase 1, ACSL1)的表达对猪前脂肪细胞分化产生抑制作用,其机制可能与其抑制甘油三酯和成脂相关标志物合成作用有关。此外,miR-130b对脂肪生成的抵抗作用已有报道,miR-130b双链可以直接靶向抑制KLF3的表达,进而减弱脂肪生成和甘油三酯合成相关基因的表达,从而导致其抗脂作用[13]。对分化4 d的鸭脂肪细胞miRNA表达谱分析发现,miR-214可调控鸭脂肪沉积,靶向肉毒碱棕榈酰基转移酶2(carnitine palmitoyl transferase 2, CPT2)的3'UTR,降低CPT2的蛋白丰度,促进脂滴形成[57]。

3.2 miRNA对肝脏糖、脂代谢的调节

Li等[58]研究发现,miR-1可通过增加靶向调节肝X受体α(liver X receptor α, LXRα)的表达促进肝脏脂质合成与积累;而miR-34a可通过靶向抑制SIRT1的表达促进肝脏糖异生过程。王星果等[59]研究表明,miR-1682和miR-1684a-3p通过调节肝脏中脂肪酸延长、非不饱和脂肪酸生物合成与脂代谢相关通路,最终调控蛋鸡肝脏脂代谢并影响生产性能。贾逸敏[60]研究表明,miRNA-1285在肝脏中靶向乙酰辅酶A羧化酶(acetyl-CoA carboxylase, ACC)和miRNA-let-7e靶向硬脂酰辅酶A去饱和酶(stearoyl-CoA desaturase, SCD),调节肝脏糖异生相关基因如磷酸烯醇丙酮酸羧激酶2(phosphoenolpyruvate carboxykinase 2, PCK2)和果糖1,6-二磷酸酶(fructose-1,6-bisphosphatase, FBP)水平,进而增强新生仔猪肝脏中的糖异生,调节新生仔猪肝脏能量代谢。综上所述,miRNA在肝脏中可通过靶向肝脏糖、脂代谢相关过程,进而调控机体的脂质沉积,但其具体的作用机制仍不明晰,需要进一步的研究加以验证其具体的作用机制。

4 miRNA与激素分泌

miRNA可通过靶向调节激素分泌水平及状态,主要体现在:1)miRNA可通过靶基因调节激素分泌相关的信号通路,进而调控激素的合成及分泌;2)通过靶向结合对应靶基因的3'UTR或靶基因的不同转录本及靶向剪接因子,调节激素水平;3)在细胞水平上,通过调控细胞不同时期的运转及代谢,对激素合成和分泌产生影响。

4.1 miRNA对生长激素的调节

生长激素是一种蛋白激素,它由垂体前叶的生长激素细胞合成和分泌,通过血液运输到靶器官和组织。研究发现,猪垂体存在的miRNA(let-7e和miR-328-5p)通过靶向生长激素释放激素受体(growth hormone releasing hormone receptor, GHRHR)不同转录本及靶向剪接因子实现对其的直接及间接调控作用,进而介导生长激素合成的作用[61-62]。不仅如此,ssc-let-7c也参与了猪垂体前叶细胞生长激素的分泌调节,它同时靶向生长激素1(growth hormone 1, GH1)和GHRHR mRNA的3'UTR,以此调节生长激素分泌;而过表达ssc-let-7c会导致生长激素分泌减少[63]。张睿等[64]发现,在延边黄牛的垂体细胞中,miR-23b-3p与生长激素分泌之间存在负调控关系,miR-23b-3p靶向结合垂体特异性转录因子1(pituitary specific transcription factor 1, POU1F1)的3'UTR,调控POU1F1 mRNA的转录,影响其蛋白的表达,以此调控延边黄牛垂体细胞生长激素分泌,进而影响延边黄牛的生长发育。

4.2 miRNA对性激素的调节

促卵泡素和促黄体生成素的主要作用为刺激卵泡发育和成熟。前期研究表明,miR-7通过靶向高尔基糖蛋白1(golgi apparatus protein 1, GLG1)促进卵泡刺激素和促黄体生成素的合成,这表明miR-7介导了猪垂体中卵泡刺激素和促黄体生成素合成的作用,从而调控卵泡发育[65]。体外试验证实,miR-429在ZEB1的3'UTR有结合位点,而转录因子ZEB1和黄体激素β(luteinizing hormone β, LHβ)之间存在靶向关系;因此,miR-429在仔猪的垂体中可能通过下调ZEB1基因的表达间接上调LHβ基因的表达,从而影响促黄体生成素生成来影响母猪的排卵过程[66]。miR-7可参与调控卵泡刺激素的表达、合成和分泌[67]。miR-361-3p直接靶向卵泡刺激素β (follicle stimulating hormone β, FSHβ)的3'UTR调控卵泡刺激素的合成和分泌,为miRNA参与卵泡刺激素的直接调控提供了依据[68]。
促性腺激素主要作用于垂体前叶,使得垂体前叶的促卵泡素、促黄体生成素分泌,有助于激活卵巢功能。不仅如此,禄欢等[69]研究了miRNA调控香猪卵巢功能及繁殖性状的分子机制,结果表明,miR-23b、let-7i-5p、miR-103、miR-30e-5p和miR-1271-5p的靶基因参与了卵母细胞的发育成熟相关的通路,包括促性腺激素释放激素受体通路。雌二醇是一种甾体雌激素,能促进母畜雌性器官和副性征的正常生长和发育。Pan等[70]研究发现,miR-378可以通过抑制猪卵丘细胞中的芳香化酶减少雌二醇的产生来调控卵母细胞成熟。徐盛玉[71]研究指出,在猪卵巢颗粒细胞中miR-378通过靶定结合芳香化酶的3'UTR直接影响雌二醇的合成,表明miR-378是实现调控繁殖内分泌的途径之一。miR-126通过靶向FSHR增加了雄激素受体诱导的猪卵泡颗粒细胞凋亡,表明其在调控FSHR表达和猪卵泡颗粒细胞凋亡中发挥积极作用[72]。但这些不同miRNA之间是否存在协同调控机制,还有待进一步探究。此外,深入解析miRNA调控性激素合成和分泌规律的机制,也将对动物的繁殖生理具有重要帮助。

4.3 miRNA对其他激素的调节

miRNA对于褪黑素、糖皮质激素及类固醇激素也具有潜在的调控作用。研究表明,miR-7作为负性调控分子,通过靶向RAF1抑制原癌基因丝氨酸/苏氨酸-蛋白激酶1(RAF-1 proto-oncogene, serine/threonine kinase 1, RAF1)/丝裂原活化蛋白激酶激酶(mitogen-activated protein kinase kinase, MEK)/ERK信号通路,进而调控褪黑素的表达、合成和分泌[73]。miR-128通过抑制SIRT1表达,上调糖皮质激素分泌相关基因表达和酶活性,表明其在糖皮质激素相关调控中发挥的重要作用[74]。李敬[75]研究指出,gga-miR-449a能够靶向胰岛素样生长因子2-mRNA结合蛋白3(insulin-like growth factor 2 mRNA-binding protein 3, IG2BP3)的3'UTR降低荧光素酶活性, 进而调控蛋鸡卵巢颗粒细胞类固醇激素E2/P4合成和分泌。

5 miRNA与肠道健康

本实验室前期研究发现,粪便中的miRNA可以缓解结肠炎症反应,表明miRNA对炎症性疾病具有预防或治疗作用[76]。本实验室最新研究指出,断奶应激导致miRNAs组成改变,ssc-miRNA-425-5p和ssc-miRNA-423-3p一方面通过抑制琥珀酸在肠道内的累积所引起的仔猪腹泻;另一方面通过调节猪肠道上皮细胞的氯离子分泌抑制巨噬细胞中髓样分化因子88(myeloid differentiation factor 88, MyD88)依赖的TLR4信号通路,从而降低仔猪腹泻率[77]。宫内发育迟缓仔猪最典型的特征是肠道屏障功能受损。Zhu等[78]研究发现,miR-29a与宫内发育迟缓所引起的肠道屏障受损有关,miR-29a抑制了细胞外基质相关基因(整合素α1、胰岛素样生长因子1、Ⅳ型胶原蛋白及纤连蛋白)和内皮紧密连接蛋白1的表达,表明miR-29a为预防及治疗仔猪宫内发育迟缓提供了潜在的治疗方案。在猪乳外泌体中,miR-4334和miR-219可通过核因子-κB(nuclear factor kappa-B,NF-κB)通路减少由脂多糖诱导的肠道炎症[79]。Wang等[80]研究发现,miR-450a通过下调脂多糖诱导的TNF因子(lipopolysaccharide-induced TNF factor, LITAF)的基因表达改善仔猪肠道抗炎能力,证明了miR-450a对于动物健康的有益作用。研究发现,miR-185可直接靶向细胞分裂周期42(cell division cycle protein 42, CDC42),促进猪小肠上皮细胞细胞增殖,表明miR-185有益于肠道健康,然而miR-185/CDC42轴在仔猪抵抗腹泻中的具体调控机制仍有待进一步研究[81]。
综上所述,miRNA可通过多途径影响机体肠道健康,主要体现在:1)miRNA可通过减少肠道内代谢产物水平,如减少肠道内琥珀酸累积水平,调控腹泻发生;2)调节肠道上皮细胞的离子分泌状态,如调节氯离子分泌进而抑制炎性通路的激活,降低腹泻率;3)在细胞水平上,通过调节细胞炎症和凋亡水平,增强上皮屏障功能。

6 小结及展望

综上所述,miRNA作为生理过程发展中不可或缺的调节器,在调节畜禽肌肉发育、脂肪代谢、肉品质、繁殖生理、激素分泌和肠道健康等方面扮演着重要角色。目前,miRNA对动物生理机能的调控网络初步形成,揭示了miRNA与靶基因的潜在效应,这对于研究与畜禽生产有关的产业和食品工业的健康稳态具有重要科学意义。然而,大部分研究仅局限于解析miRNA与靶基因作用关联上,部分主效应靶基因与生理机能作用发挥之间的靶向调控分子机制仍需深度挖掘。因此,在鉴于miRNA和靶基因的相互作用基础上,对两者所参与的信号调控通路与途径的作用机制进行功能研究及表型调控的研究仍需进一步探索。

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