动物营养学报 >

2025 , Vol. 37 >Issue 7: 4201 - 4208

DOI: https://doi.org/10.12418/CJAN2025.343

综述

支链氨基酸在肉鸡低蛋白质饲粮中应用的研究进展

  • 李彬 , 1, 2, 3 ,
  • 陈柳 1, 2, 3 ,
  • 虞洁 1, 2 ,
  • 梅绍锋 , 2, 3, *
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  • 1 四川农业大学,成都 611130
  • 2 四川特驱农牧科技集团有限公司,成都 610207
  • 3 新津邦得科技有限公司,成都 611434
* 梅绍锋,高级畜牧师,E-mail:

李 彬(1989—),女,四川成都人,初级畜牧师,硕士,从事动物饲料与营养科学研究。E-mail:

Copy editor: 菅景颖

收稿日期: 2024-12-08

  网络出版日期: 2025-07-12

基金资助

成都市重点研发支撑计划(2024-YF05-02195-SN)

成都市产业建圈强链人才计划

收起

Advances in Branched-Chain Amino Acids in Low-Protein Diets for Broilers

  • LI Bin , 1, 2, 3 ,
  • CHEN Liu 1, 2, 3 ,
  • YU Jie 1, 2 ,
  • MEI Shaofeng , 2, 3, *
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  • 1 Sichuan Agricultural University, Chengdu 611130, China
  • 2 Sichuan Tequ Agriculture and Animal Husbandry Technology Group Co., Ltd., Chengdu 610207, China
  • 3 Xinjin Bangde Technology Co., Ltd., Chengdu 611434, China
* senior livestock specialist, E-mail:

Received date: 2024-12-08

  Online published: 2025-07-12

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摘要

维持支链氨基酸(BCAA)平衡对于调节肉鸡的糖、脂代谢和蛋白质合成具有重要作用。低蛋白质饲粮容易引起BCAA比例失衡,对肉鸡生长性能产生负面影响。肉鸡的BCAA需要量受饲粮组成、生长发育阶段及生产效率等因素的影响。本文结合国内外的研究进展,系统阐述了BCAA在肉鸡体内的代谢途径及其推荐比例,并基于BCAA的拮抗效应以及不同形式氨基酸的消化动力学差异,对不同饲料原料的BCAA组成特征及其在肉鸡低蛋白质饲粮中的应用进行了综述,旨在为精准评估促进肉鸡生长以及肌肉蛋白质沉积的BCAA需要量提供参考。

关键词： 肉鸡; 支链氨基酸; 低蛋白质饲粮; 代谢机制; 平衡比例

本文引用格式

李彬 , 陈柳 , 虞洁 , 梅绍锋 . 支链氨基酸在肉鸡低蛋白质饲粮中应用的研究进展[J]. 动物营养学报, 2025 , 37(7) : 4201 -4208 . DOI: 10.12418/CJAN2025.343

Abstract

Maintaining the balance of branched-chain amino acids (BCAA) is crucial for regulating the sugar metabolism, lipid metabolism and protein synthesis in broilers. Low-protein diets are prone to causing an imbalance in the BCAA ratio, negatively affecting growth performance of broilers. The BCAA requirements of broilers are influenced by variables such as feed ingredients, growth and development stage, and productivity target. Consequently, this paper synthesized the research advancements from domestic and international sources to delineate the metabolic mechanism of BCAA and recommended BCAA ratios in broilers. Considering the antagonistic effects of BCAA and the differences in the digestive dynamics of various amino acid forms, this review summarized the characteristics of BCAA contents in feed ingredients and their application in low-protein diets for broilers, thereby offering theoretical support for future assessments of BCAA requirements in enhancing broiler growth and muscle protein deposition.

Key words： broilers; branched-chain amino acids; low-protein diet; metabolic mechanisms; balanced ratio

氨基酸是蛋白质的主要组成单位,也是动物机体多种含氮化合物的前体,对肉鸡糖、脂代谢以及抗氧化酶、促反应等生理过程具有重要作用,能够显著影响肉鸡的体增重和饲料转化率等生产指标[1-4]。利用理想氨基酸比例配制肉鸡饲粮,可有效提高饲粮中蛋白质(氨基酸)的利用率,降低蛋白质水平,减少未消化蛋白质的体外排出量[5-6]。亮氨酸(Leu)、异亮氨酸(Ile)和缬氨酸(Val)在结构上与支链脂肪酸类似,具有疏水侧链,被称为支链氨基酸(BCAA)[7]。其中,Leu的化学名称为2-氨基-4-甲基戊酸,Ile为2-氨基-3-甲基戊酸,Val为2-氨基-3-甲基丁酸。已有研究证实,BCAA能够通过哺乳动物雷帕霉素靶蛋白(mTOR)信号通路促进肌肉蛋白质合成[8-10]。但由于BCAA之间存在拮抗效应,某一BCAA过量可能导致其他水平较低的BCAA代谢分解加剧,诱发BCAA继发性缺乏,对肉鸡生产性能造成不利影响[10-11]。因此,维持BCAA动态平衡已成为肉鸡低蛋白质饲粮技术体系的重要研究方向。本文系统阐述了BCAA在肉鸡体内的代谢途径及其推荐比例,归纳了不同饲料原料中BCAA的组成特征,深入探讨了低蛋白质饲粮模式下BCAA对肉鸡生长性能、氨基酸消化率以及胴体和肌肉品质等方面的调控效应,旨在为后续评估调节肉鸡生长性能和肌肉蛋白质沉积等功能的BCAA需要量提供参考。

1 BCAA在肉鸡体内的代谢途径

BCAA在肉鸡体内的代谢主要发生在肌肉和肝脏中。研究发现,Leu、Val和Ile首先在肌肉中经支链氨基转移酶催化进行可逆的转氨基反应,分别生成α-酮异己酸、α-酮异戊酸和α-酮-β-甲基戊酸;随后转运至肝脏,在支链α-酮酸脱氢酶复合物作用下进行不可逆的氧化脱羧,其中Leu生成乙酰-辅酶A,Val生成琥珀酰-辅酶A,Ile则同时生成乙酰-辅酶A和琥珀酰-辅酶A,参与三羧酸循环,并通过生糖途径和生酮途径参与机体的糖、脂和蛋白质的合成与转换[11-13]。但在代谢过程中,支链α-酮酸脱氢酶复合物活性受丙酮酸脱氢酶的磷酸化-去磷酸化机制调控。支链α-酮酸脱氢酶激酶介导的磷酸化会导致支链α-酮酸脱氢酶复合物失活,而支链α-酮酸脱氢酶磷酸酶可以脱去支链α-酮酸脱氢酶的磷酸基团激活支链α-酮酸脱氢酶复合物[13]。因此,Leu过量会导致α-酮异己酸含量增加,触发负反馈机制抑制支链α-酮酸脱氢酶激酶,并在支链α-酮酸脱氢酶磷酸酶的去磷酸化作用下增强支链α-酮酸脱氢酶复合物活性,加剧Val和Ile的分解代谢,导致继发性BCAA缺乏[14-15]。此外,α-酮异戊酸和α-酮-β-甲基戊酸也能抑制支链α-酮酸脱氢酶激酶,但抑制效果弱于α-酮异己酸[12]。Val或Ile过量仅引起其他BCAA的血浆浓度略微下降,表明Val、Ile增加不会显著加剧BCAA分解代谢[16]。综上可知,Leu水平是BCAA拮抗效应和平衡模型的关键因素,在低蛋白质饲粮中,Val和Ile的最佳水平应基于Leu水平调整。

2 肉鸡饲粮中BCAA的推荐比例

目前,关于肉鸡饲粮中蛋氨酸(Met)、Lys和苏氨酸(Thr)这3种限制性氨基酸平衡的相关研究已经较为成熟,但由于BCAA之间存在拮抗作用,因此,有研究认为BCAA也可能成为限制因素。NRC(1994)推荐的肉鸡各生长阶段Leu:Ile:Val:Lys均为109:73:82:100,其中0~3周龄的Leu、Ile和Val需要量分别为1.20%、0.80%和0.90%,3~6周龄分别为1.09%、0.73%和0.82%,6~8周龄分别为0.93%、0.62%和0.70%[17]。在现有理想氨基酸模式的研究中,本文重点总结了BCAA在肉鸡饲粮中的推荐比例[18-26](表1),结果显示,Leu:Ile:Val:Lys在肉小鸡阶段的平均为110:74:82:100,与NRC(1994)的推荐比例接近,但在肉大鸡阶段的提高至116:74:83:100,表明NRC(1994)的推荐比例可能不能满足肉大鸡对Leu的需要量。此外,Sklan等[24]研究表明,在肉鸡1~42日龄的饲养期内,每周的BCAA推荐比例均存在一定差异,Leu变化范围为96~123,Ile为77~91,Val为83~100,其中肉鸡在1~7日龄对Leu和Val的需要量最低,在22~28日龄对Ile的需要量最低,在15~21日龄对 Leu、Ile和Val 的需要量达到最高。因此,为了节约蛋白质类原料、实现肉鸡的最佳生长性能,后期可根据肉鸡的不同生长阶段调整饲粮中BCAA的比例,并关注BCAA与其他氨基酸之间的相互作用,以实现肉鸡饲粮中氨基酸的总体平衡。
表1 支链氨基酸在肉鸡饲粮中的推荐比例

Table 1 Recommended proportion of BCAA in broiler diet

阶段
Phases		 赖氨酸
Lys		 支链氨基酸 BCAA 参考文献
References
亮氨酸 Leu 异亮氨酸 Ile 缬氨酸 Val
肉小鸡 Broiler chicks
1~21日龄 1 to 21 days of age 100 107 71 73 [18]
1~21日龄 1 to 21 days of age 100 126 63 86 [19]
1~21日龄 1 to 21 days of age 100 110 66 76 [20]
1~21日龄 1 to 21 days of age 100 109 67 77 [21]
1~21日龄 1 to 21 days of age 100 109 66 76 [22]
1~21日龄 1 to 21 days of age 100 109 67 77 [23]
1~7日龄 1 to 7 days of age 100 96 85 83 [24]
8~14日龄 8 to 14 days of age 100 103 90 86 [24]
15~21日龄 15 to 21 days of age 100 123 91 100 [24]
7~28日龄 7 to 28 days of age 100 108 63 81 [25]
平均值 Mean 100 110 74 82
肉大鸡 Broiler chickens
22~42日龄 22 to 42 days of age 100 109 69 80 [23]
22~42日龄 22 to 42 days of age 100 128 72 78 [26]
22~42日龄 22 to 42 days of age 100 123 71 79 [26]
22~42日龄 22 to 42 days of age 100 127 71 79 [26]
22~28日龄 22 to 28 days of age 100 106 77 86 [24]
29~35日龄 29 to 35 days of age 100 111 79 91 [24]
36~42日龄 36 to 42 days of age 100 113 83 91 [24]
平均值 Mean 100 116 74 83

3 不同饲料原料中BCAA的组成特征

在低蛋白质饲粮中,主要利用非结合氨基酸(合成氨基酸)配制氨基酸平衡饲粮,以提高蛋白质(氨基酸)利用率并降低饲料成本[27-28]。在肉鸡低蛋白质饲粮(粗蛋白质含量在18.75%)中,Greenhalgh等[29]发现高水平BCAA在小麦型饲粮中显著降低肉鸡的体增重,而在高粱型饲粮中显著增加肉鸡的体增重,该差异可能与饲料原料中非结合氨基酸的含量有关,小麦型饲粮中非结合BCAA的含量较高,占总BCAA含量的52.6%,而高粱型饲粮中仅占27.2%。过量的非结合BCAA可能引发肉鸡肠道内氨基酸氧化损失,造成非结合氨基酸和蛋白质结合氨基酸之间的不平衡,以及过剩氨基酸的脱氨[29];并且,非结合氨基酸和蛋白质结合氨基酸在消化动力学上可能存在差异,导致蛋白质合成部位的氨基酸不平衡,阻碍肉鸡的生长发育[30-31]。因此,配制低蛋白质氨基酸平衡饲粮应考虑饲粮中非结合氨基酸和蛋白质结合氨基酸含量及其肠道吸收率。Li等[32]分析了不同饲料原料在非结合BCAA、蛋白质结合BCAA和总BCAA含量上的差异(表2),结果发现,动物性原料的非结合BCAA含量明显高于植物性原料,其中以喷雾干燥蛋白胨的非结合BCAA含量最高;相反,喷雾干燥禽血浆和羽毛粉的非结合BCAA含量较低,但蛋白质结合Leu和Val含量较高;此外,螺旋藻粉和大豆浓缩蛋白中蛋白质结合Ile含量也较高。另外,玉米蛋白粉的Leu含量接近10%,远高于Ile和Val含量[14]。综上可知,在配制肉鸡低蛋白质BCAA平衡饲粮时,喷雾干燥禽血浆、羽毛粉、螺旋藻粉、大豆浓缩蛋白以及玉米蛋白粉均可作为理想原料,用于平衡蛋白质结合BCAA,防止肉鸡体内蛋白质合成部位的氨基酸不平衡。
表2 不同饲料原料中非结合支链氨基酸、蛋白质结合支链氨基酸及总支链氨基酸含量

Table 2 Contents of non-bound BCAA, protein-bound BCAA and total BCAA in different feedstuffs[32]g/kg

项目
Items		 非结合支链氨基酸
Non-bound BCAA		 蛋白质结合支链氨基酸
Protein-bound BCAA		 总支链氨基酸
Total BCAA
亮氨酸
Leu		 异亮氨酸
Ile		 缬氨酸
Val		 亮氨酸
Leu		 异亮氨酸
Ile		 缬氨酸
Val		 亮氨酸
Leu		 异亮氨酸
Ile		 缬氨酸
Val
黑水虻幼虫粉
Black soldier fly larvae meal		 0.80 0.25 0.45 31.73 19.95 28.21 32.54 20.21 28.65
鸡副产物粉
Chicken by-product meal		 0.35 0.25 0.32 52.40 26.94 32.89 52.75 27.19 33.21
鸡内脏消化物
Chicken visceral digest		 5.42 3.07 3.15 55.61 35.30 52.53 61.03 38.37 55.68
羽毛粉 Feather meal 0.27 0.12 0.20 68.90 37.68 57.94 69.17 37.80 58.13
鱼粉(美国)
Fish meal (USA)		 0.82 0.38 0.57 44.19 23.81 29.63 45.01 24.19 30.20
鱼粉(秘鲁)
Fish meal (Peruvian)		 1.81 0.83 1.08 50.92 26.69 35.45 52.73 27.52 36.53
鱼粉(东南亚)
Fish meal (Southeast Asian)		 4.56 2.91 3.85 41.36 24.99 28.69 45.92 27.90 32.53
喷雾干燥蛋白胨
Spray-dried peptone		 22.16 12.42 16.81 20.86 12.01 13.65 43.01 24.43 30.48
禽副产物粉
Poultry by-product meal		 0.74 0.42 0.73 42.58 23.36 28.37 43.32 23.78 29.10
喷雾干燥禽血浆
Spray-dried poultry plasma		 0.19 0.05 0.12 75.40 27.68 49.49 75.59 27.73 49.62
喷雾干燥蛋制品
Spray-dried egg product		 0.10 0.06 0.07 46.02 29.68 35.64 46.11 29.74 35.71
螺旋藻粉
Algae spirulina meal		 0.09 0.04 0.07 63.13 40.82 43.53 63.22 40.87 43.61
豆粕
Soybean meal		 0.09 0.05 0.08 34.43 20.25 20.71 34.52 20.30 20.78
大豆浓缩蛋白
Soy protein concentrate		 0.04 0.02 0.03 57.87 37.70 38.18 57.91 37.72 38.21

4 BCAA在肉鸡低蛋白质饲粮中的应用研究

4.1 低蛋白质饲粮中BCAA对肉鸡生长性能的影响

针对不同的生产目标,肉鸡低蛋白质饲粮的BCAA推荐比例也有差异。在20~35日龄肉鸡低蛋白质饲粮(粗蛋白质含量为18%)研究中,Kriseldi等[33]发现,饲粮可消化Lys含量为1.10%,且Leu:Ile:Val:Lys为110:66:78:100时,肉鸡的生长性能达到最佳,但仅增加Leu含量可能会对肉鸡增重和饲料转化率产生负面影响;Ospina-Rojas等[34]对21~42日龄科宝肉鸡饲喂低蛋白质饲粮(粗蛋白质含量为16%),发现可消化Leu和Val含量分别为1.19%和0.86%时,饲料转化率最佳,可消化Leu和Val含量分别为1.15%和0.86%时,体增重最大。另外,在玉米型饲粮中,玉米中Leu含量较高,Ile和Val含量相对较低,降低饲粮蛋白质水平并增加玉米含量会导致饲粮中Leu过量,引发BCAA之间的拮抗效应,补充适量或高水平的Val和Ile可以改善Leu过量对肉鸡生长性能的负面影响[9,35-36]。此外,不同类型饲粮中饲料原料的粗蛋白质和BCAA含量的差异可能影响饲粮中非结合BCAA和蛋白质结合BCAA的比例,进而影响肉鸡的生长性能[37]。例如,在以小麦或高粱为基础的饲粮中,小麦的粗蛋白质含量高于高粱,但Leu含量却低于高粱,饲喂高粱型饲粮的肉鸡体重增加9.26%,而饲喂小麦型饲粮的肉鸡体重减少9.49%;进一步分析发现,小麦型饲粮中非结合BCAA含量高于高粱型饲粮,表明小麦型饲粮比高粱型饲粮更容易出现非结合BCAA和蛋白质结合BCAA不平衡[29],该发现与Chrystal等[38]的研究结果一致。与玉米型低蛋白质饲粮(粗蛋白质含量为16.5%)相比,饲喂小麦型低蛋白质饲粮的肉鸡的生长性能下降[38]。这可能与小麦型饲粮中非结合BCAA含量较高有关,非结合氨基酸的吸收速度比蛋白质结合氨基酸更快,进而影响蛋白质合成效率和生长性能。由此可见,肉鸡的BCAA需要量因生产目标和饲粮类型不同而异,且非结合BCAA与蛋白质结合BCAA比例失衡的不利影响在饲喂低蛋白质饲粮的肉鸡中更容易发生,建议应针对原料类型平衡饲粮中的非结合BCAA与蛋白质结合BCAA。

4.2 低蛋白质饲粮中BCAA对肉鸡氨基酸消化率的影响

低蛋白质饲粮中补充非结合氨基酸(合成氨基酸)可满足肉鸡对可消化氨基酸的需要量,增强氨基酸消化率,并有效降低氮排泄量[39-40]。但由于BCAA之间的拮抗作用,以及非结合和蛋白质结合形式的氨基酸在消化动力学上的差异,过量补充可能无法实现饲喂低蛋白质饲粮减少氮排泄的预期效益。例如,在小麦型低蛋白质饲粮中增加Ile+Val含量(20.32 g/kg)可使氮沉积率降低3.06%[41]。类似地,Greenhalgh等[29]研究指出,肉鸡饲喂低蛋白质饲粮并补充较高水平的BCAA,即Leu、Ile和Val含量分别为16.50、8.25和9.35 g/kg(Leu:Ile:Val:Lys为150:75:85:100),氮沉积率降低8.78%。尽管以上研究均未评估粪便中的氮含量,但预计氮沉积率降低将导致排泄量增加[42]。另外,Greenhalgh等[41]在小麦型低蛋白质饲粮中补充高于推荐水平的Leu(15.02 g/kg),可显著提高Leu(4.74%)、Ile(4.56%)、Val(2.90%)、Lys(3.02%)、Met(1.60%)及Thr(4.96%)等氨基酸的回肠消化率,但由于分解代谢过程需要消耗能量,因此,会大大降低能量利用率,使表观代谢能(AME)和氮校正表观代谢能(AMEn)分别降低2.98%和3.25%,从而对生长性能产生负面影响。综上可知,在肉鸡低蛋白质饲粮中补充非结合BCAA可以提高其他氨基酸的消化率,但BCAA的拮抗作用和不同形式氨基酸的消化动力学差异可能导致氮沉积率降低,进而影响肉鸡的生长性能,后期需进一步研究添加比例以满足肉鸡的生长需要。

4.3 低蛋白质饲粮中BCAA对肉鸡胴体及肌肉品质的影响

BCAA中的Leu能够通过激活mTOR信号通路,增强肌肉蛋白质合成[43]。Chen等[44]报道,肉鸡的胸肌重量因感染因黄曲霉毒素B1(AFB1)而显著降低,将饲粮Leu含量从1.16%增加至2.73%,且Ile和Val含量分别为1.60%和1.88%(Leu:Ile:Val:Lys估计为186:109:128:100),能显著增加肉鸡的胸肌重量,但饲粮中AFB1与BCAA含量无显著交互作用。Erwan等[45]研究表明,在3~6周龄肉鸡饲粮中额外添加0.5% Leu,肉鸡的胴体重量增加9%。但在固定的Leu:Ile:Val比例下,饲粮中Leu含量超过肉鸡推荐量60%后对肌肉增长无显著促进作用[46-47]。此外,低蛋白质饲粮可引起肉鸡胴体中的脂肪含量增加[48-50],但Leu能够降低血液和脂肪组织中的甘油三酯含量,从而降低脂肪含量,Val可通过抑制脂肪酸合成途径,减少腹部脂肪沉积[51-52]。因此,在低蛋白质饲粮中补充Leu和Val,可以有效减少脂肪沉积,改善胴体品质[33,51-52]。但在BCAA比例不平衡的低蛋白质饲粮中添加过量的Leu,容易引起BCAA的拮抗作用。另外,Ile可以提高肌肉产量,缺乏Ile容易导致肉鸡的肌肉产量下降[53-54]。Kriseldi等[33]研究证实,当Leu:Ile:Val:Lys为190:70:82:100,肉鸡的胴体重量最大;当Leu:Ile:Val:Lys为190:68:78:100,肉鸡的胴体率最高;当Leu:Ile:Val:Lys为190:72:81:100,肉鸡的胸肌肉重量最大;当Leu:Ile:Val:Lys为190:74:75:100,肉鸡的胸肌肉率最高。以上研究表明,高水平的Leu、Val和Ile可以促进肉鸡的肌肉增长,但需注意BCAA之间的拮抗效应;但当饲粮Leu含量超出肉鸡推荐量的特定比例,对肌肉增长的促进作用不再显著。

5 小结

综上所述,NRC(1994)的BCAA推荐量及比例可能不能满足肉大鸡对Leu的需要量。配制肉鸡低蛋白质BCAA平衡饲粮应考虑BCAA的拮抗作用和消化动力学差异,关注饲料原料中非结合氨基酸和蛋白质结合氨基酸含量。尽管补充非结合BCAA能提高其他氨基酸的消化率,但对肉鸡的氮沉积率及生长性能可能存在不利影响。此外,促进肌肉增长的BCAA需要量高于体重增长的需要量,但饲粮中Leu含量超过特定阈值后,对肌肉的促增长作用不再显著。总之,应该根据饲粮组成、生长发育阶段以及生产效率合理地调整饲粮中BCAA比例。目前,关于肉鸡低蛋白质饲粮BCAA平衡的研究主要集中在白羽肉鸡上,而对于黄羽肉鸡的研究尚显不足,后期可开展黄羽肉鸡相关研究试验,以期为BCAA在肉鸡低蛋白质饲粮体系中的深入应用提供理论依据。

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