综述

胍基乙酸的生物学功能及改善畜禽肉品质研究进展

  • 罗喜平 ,
  • 张文彦 ,
  • 瞿明仁 ,
  • 许兰娇 , *
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  • 江西农业大学动物科学技术学院, 动物营养与饲料江西省重点实验室, 南昌 330045
*许兰娇,副研究员,硕士生导师,E-mail:

罗喜平(2002—),男,湖南邵阳人,硕士研究生,从事反刍动物营养研究。E-mail:

收稿日期: 2025-08-29

  网络出版日期: 2026-03-16

基金资助

国家肉牛牦牛产业技术体系(CARS-37)

江西省2023年度研究生创新专项资金项目(YC2023-S358)

Research Progress on Biological Functions of Guanidinoacetic Acid and Its Improvement on Meat Quality in Livestock and Poultry

  • LUO Xiping ,
  • ZHANG Wenyan ,
  • QU Mingren ,
  • XU Lanjiao , *
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  • Jiangxi Provincial Key Laboratory of Animal Nutrition and Feed, College of Animal Science and Technology, Jiangxi Agricultural University, Nanchang 330045, China
*associate professor, E-mail:

Received date: 2025-08-29

  Online published: 2026-03-16

摘要

胍基乙酸是肌酸合成的唯一前体,具有抗氧化、调节能量代谢、促进肌纤维发育和胰岛素激素释放以及提高动物生长性能等多种生物学功能。本文系统阐述了胍基乙酸的理化特性、体内合成与代谢途径及其生物学功能,综述了胍基乙酸在猪、禽及反刍动物生产中改善肉品质的效果并深入探讨了其作用机理,以期为胍基乙酸在畜禽生产中肉品质调控方面的应用提供理论参考。

本文引用格式

罗喜平 , 张文彦 , 瞿明仁 , 许兰娇 . 胍基乙酸的生物学功能及改善畜禽肉品质研究进展[J]. 动物营养学报, 2026 , 38(3) : 1689 -1695 . DOI: 10.12418/CJAN2026.136

Abstract

Guanidinoacetic acid is the sole precursor for creatine synthesis and possesses multiple biological functions, including antioxidation, regulation of energy metabolism, promotion of muscle fiber development and insulin hormone release, as well as improvement of animal growth performance. This article systematically expounds the physicochemical properties, in vivo synthesis and metabolic pathways, as well as the biological functions of guanidinoacetic acid. It reviews the effects of guanidinoacetic acid on improving meat quality in pig, poultry and ruminant production and deeply explores its mechanism of action, with the aim of providing theoretical references for the application of guanidinoacetic acid in meat quality regulation in livestock and poultry production.

胍基乙酸(guanidinoacetic acid,GAA)是肌酐及其磷酸化衍生物磷酸肌酐的重要前体。肌酸在动物生产中不仅具有促进生长发育的作用,还可以提高抗氧化酶活性、延缓宰后糖酵解以及改善畜禽肉品质[1]。但由于肌酸具有溶解性差、利用率不高的问题,从而研发出了新型的肌酸补充剂GAA[2]。研究表明,GAA能够显著提高组织中肌酸水平[3]。在饲粮中添加GAA能够促进胰岛素激素的释放[4]、提高机体的抗氧化水平[5]、调节动物体内能量代谢[6]、促进肌纤维发育[7]、调节体内糖酵解[8]以及改善肉品质[9]。本文综述了GAA的理化特性、体内合成与代谢途径及其生物学功能,探讨了GAA对畜禽肉品质的改善作用及其作用机理,旨在为畜牧生产中GAA的应用提供参考。

1 GAA的理化特性以及合成与代谢途径

GAA结构稳定,室温条件下可稳定保存2年[10]。在肾脏和胰腺中,脒基转移酶催化L-精氨酸与甘氨酸发生反应,进而合成GAA[11]。然后,GAA通过血液循环转运至肝脏,GAA在S-腺苷甲硫氨酸合酶作用下进行腺苷酸化,生成S-腺苷甲硫氨酸,进而通过甲基化过程产生肌酸和S-腺苷同型半胱氨酸[12]。随后,生成的肌酸由肌酸转运蛋白1(creatine transporter 1,CT1)介导进入血液循环,供肝脏外组织吸收和代谢[13]。肌酸会被进一步磷酸化为磷酸肌酸(phosphocreatine,PCr),参与能量代谢的调控;而肌酸与PCr的最终代谢产物——肌酸酐,则经肾脏随尿液排出[14]

2 GAA的生物学功能

2.1 促进激素的释放

GAA具有调节胰岛素的功能,它能促进胰岛素分泌从而降低机体血糖水平;血糖水平的下降进而刺激动物采食,导致采食量上升,促进动物生长[15-16]。此外,GAA还具有促进生长激素分泌的作用。Ostojic等[12]研究表明,GAA对生长激素的分泌具有促进作用,通过提升γ-氨基丁酸水平以激活下丘脑,进而刺激促生长激素释放激素的释放并作用于垂体,促进生长激素生成和分泌。

2.2 抗氧化作用

GAA可以提高血清中超氧化物歧化酶(SOD)和过氧化氢酶(CAT)活性,而SOD能够清除细胞中多余的自由基[17];过氧化氢被CAT催化分解为水,从而清除自由基氧化所引起的损害[18]。此外,邱明科[19]研究发现,在肉牛饲粮中添加GAA还可提高谷胱甘肽过氧化物酶(GSH-Px)活性,而GSH-Px可以识别并催化过氧化氢等有机过氧化物反应,从而达到清除活性氧(ROS)的作用[20-21]

2.3 能量代谢相关功能

GAA在调节脊椎动物细胞能量代谢中扮演重要角色,其主要通过作为肌酸的唯一前体来发挥作用[22]。肌酸及其磷酸化形式PCr是调节能量转移和维持三磷酸腺苷(ATP)恒定水平的关键物质[23]。GAA可以促进肌酸的合成,增加体内肌酸和PCr含量并通过磷酸激酶反应与二磷酸腺苷(ADP)作用,有效再生ATP,从而维持细胞内ATP的稳定水平,为机体提供持续的能量供应[4,24]

2.4 促进肌纤维发育

GAA促进肌纤维发育,主要通过以下2个方面。一方面,GAA可以增强相关基因的表达,研究发现GAA通过上调背最长肌中肌球蛋白重链4(MYH4)、成肌分化因子(MyoD)和生肌因子5(Myf5)等基因的表达,从而有效促进肌肉的生长和发育[25]。此外,GAA还能提高体内胰岛素样生长因子-1(IGF-1)水平,IGF-1作为一种强效的促生长因子,对肌肉内蛋白质的合成和沉积具有促进作用,进而促进肌肉的生长和肌纤维的肥大[4]。另一方面,GAA可以抑制一些阻碍肌纤维发育基因的表达,如肌肉生长抑制素对肌肉生长和力量提升起负向调控,其水平的高低会影响肌肉的发育和功能[26]。Gustafsson等[27]研究表明,GAA及其代谢物肌酸能够有效下调肌肉生长抑制素的表达,从而缓解其对肌肉生长的抑制作用,促进肌肉的生长和发育。

3 GAA对畜禽肉品质的影响

近年来,许多研究指出在畜禽饲粮中添加GAA可以提高畜禽生产性能和抗氧化能力,促进肌纤维发育,调节脂质代谢,减缓pH下降,降低肌肉的滴水损失和剪切力,改善肌肉嫩度。辛均平[28]在锦江黄牛饲粮中添加0.2%和0.4% GAA能显著提高平均日增重,降低料重比,提高锦江黄牛生长性能。王亚琼[29]研究发现,在樱桃谷肉鸭饲粮中添加0.025%、0.050%和0.100% GAA,血清总抗氧化能力(T-AOC)及CAT和SOD活性均有提高的趋势,血清丙二醛(MDA)含量有所降低,且宰后24 h胸肌滴水损失降低。上述研究表明,饲粮添加GAA通过提高一些与自由基代谢相关酶的活性来提高pH,从而减少脂质过氧化和肌肉的滴水损失,进而提升肉品质。Li等[30]研究表明,在羔羊饲粮中添加0.09% GAA,通过激活IGF-1/蛋白激酶B(Akt)/哺乳动物雷帕霉素靶蛋白(mTOR)信号通路促进肌原纤维蛋白合成,并通过调控肌肉生长抑制素和磷酸化叉头框蛋白O1(FoxO1)丰度来抑制蛋白质分解,使得屠宰后羔羊肉品质得到改善。晁雅琳等[31]研究表明,饲粮添加0.08% GAA不仅显著降低尾脂重、胴体重和胴体脂肪含量,还显著提高肌肉糖原以及肌内脂肪和蛋白质含量,表明GAA能够调节脂质代谢,促进肌内脂肪沉积,减少舍饲滩羊的胴体脂肪含量,从而提升滩羊肉品质。GAA还可以提高肌肉中肌酸和PCr含量,抑制能量代谢单磷酸腺苷活化蛋白激酶(AMPK)信号通路中肝脏激酶B1(LKB1)和AMPKα的表达,延缓宰后糖酵解从而减缓pH下降,对肉品质有一定的改善作用[8]。Zhu等[32]研究发现,在生长育肥猪饲粮中添加450 mg/kg GAA不仅能够使背最长肌重量增加,同时还降低了其剪切力、黄度(b*)值和滴水损失,从而改善其肉品质和产肉性能。
此外,也有研究发现在家禽饲粮中添加高剂量GAA以及在饲粮中甲硫氨酸不平衡时会对家禽生长性能和肉品质产生负面影响。Majdeddin等[33]研究发现,在25~42日龄雄性罗斯(Ross)308肉仔鸡饲粮中添加0.06% GAA能显著提高平均日增重和饲料转化率,然而在饲粮甲硫氨酸含量过低或过高时,添加高剂量GAA(0.12%)则会对肉鸡生长性能产生负面影响。Xiao等[34]研究发现,在黄羽肉鸡饲粮中添加1 200 mg/kg GAA时,肌肉滴水损失和血清谷丙转氨酶(ALT)活性会显著上升,表明高剂量GAA对肉鸡肉品质和肝脏功能存在负面影响。

4 GAA改善畜禽肉品质的作用机理

GAA作为功能性饲料添加剂可有效改善肉品质,其可能的作用机理总结为以下几个方面。

4.1 改善畜禽抗氧化能力

在畜禽养殖过程中,许多因素会直接或间接引起动物的氧化应激反应,而氧化应激会损害畜禽的健康导致生长变缓和生产性能的下降,从而增加养殖成本[35]。而氧化应激对肌肉的主要影响包括蛋白质变性、脂质过氧化以及风味物质和肉色的改变。脂质氧化产生的醛类也是畜禽肉产品产生异味的主要原因[36],而蛋白质氧化也会通过降低蛋白质溶解度来加剧肌肉的滴水损失[37]
目前,研究已证实GAA具有减轻畜禽养殖中氧化应激的作用。一方面,GAA能够提升组织中肌酸水平[3]。肌酸在抗氧化和抗凋亡反应中发挥着重要作用,能够清除自由基并增强体内抗氧化酶的活性[38]。因此,GAA通过提高肌酸水平,间接增强机体的抗氧化能力。同时,肌酸会在线粒体内转化为PCr,后者在细胞内的循环(从线粒体到细胞质)过程中,也有助于减少ROS的形成,从而减轻氧化应激[39-40]。另一方面,GAA能够提高动物血清中SOD和CAT等关键抗氧化酶的活性[41-43]。这些酶在抗氧化防御体系中起着核心作用,通过催化自由基的分解来增强机体的抗氧化能力;同时,GAA也参与清除由肌酸和精氨酸代谢过程中产生的氧自由基,会进一步降低体内氧化应激水平。此外,还有研究表明通过精氨酸保留效应,GAA促进一氧化氮和谷胱甘肽的合成途径,间接提升谷胱甘肽(GSH)水平[5]。GSH是重要的抗氧化剂,能有效清除自由基,增强抗氧化防御系统。GAA通过多种途径和机制,包括提高组织中肌酸和抗氧化剂水平、抗氧化酶活性以及清除自由基和减少ROS生成等,综合增强畜禽抗氧化能力,从而有利于畜禽肉品质的改善。

4.2 调节机体糖酵解过程

研究表明,宰后糖酵解对肌肉宰后变化过程有着重要的影响,如肌肉宰后的pH。pH是辨别肉品质优劣的重要指标之一,其数值越低,说明肉的酸味越强,肉品和风味就越差,肌肉pH在一定范围内偏高时肉品质较好,较高的pH有助于维持肌肉的系水力、肉色和剪切力,是提升肉品质的关键因素[44]。GAA调控糖酵解过程和肌肉pH来改善肉品质,主要通过以下几个方面来体现。一方面,GAA能够提高肌肉中的肌酸、PCr和ATP储备,这些物质是参与肌肉能量代谢的重要化合物[45]。通过增加这些能量储备,GAA有效延缓了肌肉在屠宰后的糖酵解过程,减少了乳酸的积累,避免了肌肉pH的急剧下降[8]。另一方面,GAA作为肌酸的前体能够促进PCr和ATP的合成,降低细胞内一磷酸腺苷(AMP)/ATP值,间接抑制AMPK的过度激活,从而减缓糖酵解过程,改善畜禽的肉品质[46]。研究发现,AMPK是细胞能量代谢的核心调控因子,对糖酵解具有重要影响;AMPK可以通过感知细胞内能量状态(AMP/ATP值)来调节糖酵解,主要作用包括促进糖酵解快速生成ATP,同时协调其他代谢途径以维持能量平衡[47]。当畜禽被屠宰后,细胞内AMP/ATP值开始升高(即能量不足),AMPK被激活并促进糖酵解过程,导致畜禽肉品质下降[48]。已有试验证明,在畜禽饲粮中添加GAA对AMPK有一定的调节作用[49]。此外,GAA可能通过AMPK信号通路增强线粒体的氧化磷酸化,减少对无氧糖酵解的依赖从而改善畜禽肉品质[50-51]。在缺氧条件下,AMPK可能通过抑制缺氧诱导因子-1α(HIF-1α)活性,减少糖酵解相关基因的表达,从而抑制糖酵解[52-53]。因此,GAA通过促进肌肉能量代谢重要化合物的合成以及通过对AMPK信号通路的影响,抑制和减缓糖酵解过程,并提高肌肉pH、系水力、剪切力以及肉色等关键指标,进而改善肉品质。

5 小结

GAA作为一种饲料添加剂,具有促进畜禽肌肉生长、改善能量代谢、增强机体抗氧化能力、调节氨基酸代谢以及改善肉品质等生物学功能。本文探讨了GAA及其代谢产物通过改善能量代谢、增强抗氧化能力等多途径影响肉品质的机理,为GAA在畜禽生产中的应用提供了参考依据。目前,GAA的生物学功能及其对肉品质的调控已引起广泛关注,但是仍有许多问题有待探索,如GAA对不同生长阶段畜禽肌肉生长的调控机理、GAA精细调控能量代谢的途径等。总得来说,GAA调控肉品质的机制十分复杂,未来研究仍需进一步揭示其分子层面的作用机理,以推动其在畜禽生产中的科学应用。
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