动物营养学报 >

2023 , Vol. 35 >Issue 7: 4108 - 4118

DOI: https://doi.org/10.12418/CJAN2023.380

综述

支链氨基酸的生物学作用机制及在猪和鸡生产中的应用

  • 王宗伟 , 1, 2 ,
  • 庄佳荣 3 ,
  • 李洪涛 3 ,
  • 隋美霞 , 1, *
展开
  • 1 潍坊学院生物与海洋学院,潍坊 261061
  • 2 寿光市政务服务中心,寿光 262700
  • 3 临沂布恩生物科技有限公司,临沂 276036
* 隋美霞,副教授,E-mail:

王宗伟(1982—),男,山东寿光人,高级畜牧师、执业兽医师,硕士,主要从事动物营养与畜禽疾病的研究。E-mail:

Office editor: 田艳明

收稿日期: 2023-04-24

  网络出版日期: 2023-07-11

基金资助

山东省重点研发计划(重大科技创新工程)项目(2019JZZY020609)

收起

Biological Mechanism of Branched-Chain Amino Acids and Their Application in Pig and Poultry Production

  • WANG Zongwei , 1, 2 ,
  • ZHUANG Jiarong 3 ,
  • LI Hongtao 3 ,
  • SUI Meixia , 1, *
Expand
  • 1 College of Biology and Oceanography, Weifang University, Weifang 261061, China
  • 2 Shouguang Government Service Center, Shouguang 262700, China
  • 3 Linyi Backbone Biotechnology Co., Ltd., Linyi 276036, China
* associate professor, E-mail:

Received date: 2023-04-24

  Online published: 2023-07-11

Fold

摘要

支链氨基酸(亮氨酸、缬氨酸、异亮氨酸)结构相似,在人和动物体内代谢途径相同、功能相近。支链氨基酸具有多种生物学作用,例如亮氨酸能够刺激蛋白质合成,增强免疫功能;缬氨酸、异亮氨酸能够提升抗氧化性能,促进乳蛋白和乳脂合成等。本文综述了支链氨基酸的物理特性、吸收和代谢,以及生物学作用机制及其在猪和鸡生产中的应用,以期为利用支链氨基酸配制畜禽功能性和低蛋白质饲粮提供理论和实践参考。

关键词： 支链氨基酸; 代谢; 低蛋白质饲粮; 蛋白质合成; 免疫功能; 抗应激

本文引用格式

王宗伟 , 庄佳荣 , 李洪涛 , 隋美霞 . 支链氨基酸的生物学作用机制及在猪和鸡生产中的应用[J]. 动物营养学报, 2023 , 35(7) : 4108 -4118 . DOI: 10.12418/CJAN2023.380

Abstract

Branched-chain amino acids (leucine, valine, isoleucine) have similar structure, same metabolic pathway and similar functions in human and animal bodies. Branched-chain amino acids have a variety of biological functions, such as leucine can stimulate protein synthesis and enhance immune function, valine and isoleucine can improve antioxidant performance, promote milk protein and milk fat synthesis. This paper summarized the physical properties, absorption and metabolism, and the biological mechanism of branched-chain amino acids and their application in pig and poultry production, in order to provide theoretical and practical reference for formulating functional and low protein diets for livestock and poultry using branched-chain amino acids.

Key words： branched-chain amino acids; metabolism; low protein diet; protein synthesis; immune function; anti-stress

支链氨基酸(branched-chain amino acids,BCAAs)具有多种生物学作用,例如亮氨酸(leucine,Leu)能刺激蛋白质合成[1],增强免疫功能[2];缬氨酸(valine,Val)和异亮氨酸(isoleucine,Ile)能提升抗氧化性能[3],促进乳蛋白和乳脂合成[4⇓-6]。但是,BCAAs比例失衡会严重影响畜禽的生长性能,例如过多的Leu会增加Val和Ile的分解代谢,减少蛋白质沉积率[7];降低血液中色氨酸和下丘脑5-羟色胺含量,降低采食量[8]。L-Leu[9]、L-Val[10]和L-Ile[11]的规模化生产为精准配制不同养殖条件下的BCAAs平衡饲粮提供了必要条件,在饲粮中添加适当比例的BCAAs,能够在保持生产性能的前提下,显著降低饲粮粗蛋白质水平[12-13],有效节约饲用蛋白质;在饲粮中添加适量Leu能够缓解育肥猪热应激[14],添加适量Val能够改善蛋鸡肠道健康[15],提升畜禽健康水平。本文综述了BCAAs的物理特性、吸收和代谢,以及生物学作用机制及其在猪和鸡生产中的应用研究进展,以期为利用BCAAs配制畜禽功能性和低蛋白质饲粮提供理论和实践参考。

1 BCAAs的物理特性、吸收和代谢

L-Leu(化学名称为L-2-氨基-4-甲基戊酸,化学式为C6H13NO2)、L-Val(化学名称为L-2-氨基-3-甲基丁酸,化学式为C5H11NO2)、L-Ile(化学名称为L-2-氨基-3-甲基戊酸,化学式为C6H13NO2)均为白色晶体或结晶性粉末,无臭,略有苦味。由于Leu、Val和Ile的分子结构中都有1个甲基侧链,所以被称为BCAAs。D-BCAAs的营养价值低于L-BCAAs[16],因此添加到饲粮中的BCAAs均为L型,含量在98.5%以上。
Leu、Val和Ile的主要吸收部位在小肠[17⇓-19],通过肠道氨基酸转运体主动转运到肠道细胞[20]。Leu、Val、Ile在吸收部位的拮抗作用可能不显著,因为BCAAs能上调肠道丙氨酸-丝氨酸-半胱氨酸转运蛋白(alanine-serine-cysteine transporter 2,ASCT2)等mRNA的表达,增强对氨基酸的吸收[20-21]。
BCAAs吸收进入机体后,部分参与肌肉等部位的蛋白质合成,也有部分BCAAs被机体代谢分解。BCAAs分解代谢的第1步是Leu、Val、Ile通过支链氨基酸转氨酶(branched-chain amino acid transaminase,BCAT)在肌肉等部位进行可逆的转氨基作用,形成支链α-酮酸(branched-chain α-keto-acids,BCKAs),即α-酮异己酸(α-keto isocaproic acid,KIC)、α-酮异戊酸(α-keto isovaleric acid,KIV)和α-酮-β-甲基戊酸(α-keto-β-methyl-n-valeric acid,KMV)[22];第2步是BCKAs在肝脏支链α-酮酸脱氢酶(branched-chain α-keto acid dehydrogenase,BCKDH)的催化下进行不可逆的氧化脱羧进入三羧酸(tricarboxylic acid,TCA)循环[3,23],释放能量,供机体使用,或者重新进入糖类、脂肪和蛋白质代谢。其中,Leu分解代谢为乙酰辅酶A(acetyl-CoA)[24],属于生酮氨基酸;Val代谢为琥珀酰辅酶A(succinyl-CoA)[25],属于生糖氨基酸;Ile代谢为乙酰辅酶A、琥珀酰辅酶A[26],属于生酮生糖氨基酸;不同的生酮生糖特性,使得Leu、Val、Ile能够差异化调控机体三大物质代谢,以及细胞线粒体的功能。
过量Leu通过转录后调控,增强BCKDH活性,促进Ile和Val分解[23];过量Val或Ile也能通过影响肝脏BCAAs代谢基因,促进BCAAs分解[3]。在疾病等条件下,BCAAs代谢也会发生改变,例如鸡传染性支气管炎病毒、脂多糖(lipopolysaccharide,LPS)刺激分别增加鸡肾脏、肝脏BCAAs的分解[27-28]。

2 BCAAs的生物学作用机制

2.1 调节消化、吸收和内分泌功能

BCAAs能够增加胰岛素和胰岛素样生长因子(insulin-like growth factor,IGF)的分泌,促进肠道发育和消化酶分泌,优化肠道菌群。
Leu通过变构激活胰腺β细胞内的谷氨酸脱氢酶,诱导胰岛素释放[29],KIC也能够促进小鼠胰岛中胰岛素的分泌[30];Val和Ile通过上调猪肝脏中IGF1 mRNA表达水平,下调胰岛素样生长因子结合蛋白1(insulin-like growth factor-binding protein 1,IGFBP1)mRNA表达水平,发挥IGF1的促生长作用[12]。Val和Ile通过上调肠道表皮生长因子受体(epidermal growth factor receptor,EGFR)和半胱天冬酶-9(Caspase-9)mRNA表达水平,改善仔猪小肠的发育[12];适宜比例的BCAAs能够显著促进爱拔益加(AA)肉鸡肠道生长发育[31];Leu通过增强乳牛胰腺腺泡细胞线粒体TCA循环中的柠檬酸合酶和ATP酶活性,促进一般分泌(general secretory,Sec)信号通路,增加α-淀粉酶的分泌[32]。Val和Ile能够提高断奶仔猪结肠中放线菌门(Actinobacteria)、肠球菌属(Enterococcus)和短芽孢杆菌属(Brevibacillus)丰度[33]。
过量的BCAAs及其代谢物能够降低采食量,抑制葡萄糖的吸收和利用,不利于动物的生长。过量的Leu、Ile或BCAAs混合物通过上调味觉二聚体受体1型成员1/3(taste dimeric receptor type 1 members 1/3,T1R1/T1R3)mRNA和蛋白表达,显著增加体外猪空肠模型胆囊收缩素(cholecystokinin,CCK)的分泌[34],而CCK有降低采食量的作用;Val能够降低大鼠大脑色氨酸含量[35],而色氨酸对采食量有促进作用。过量的KIC和KMV在骨骼肌细胞中通过上调雷帕霉素复合物1的机制靶点(mechanistic target of rapamycin complex 1,mTORC1)和下游蛋白翻译信号,抑制胰岛素诱导的蛋白激酶B(protein kinase B,Akt)磷酸化,降低葡萄糖摄取和线粒体耗氧[36]。

2.2 调节免疫功能

2.2.1 增强免疫功能

BCAAs及其代谢物能够增强肠道上皮细胞的免疫功能,促进肠道分泌型免疫球蛋白A(secretory immunoglobulin A,sIgA)分泌,提高粒细胞活性和血清免疫球蛋白M(immunoglobulin M,IgM)含量,增强免疫功能。Leu通过激活哺乳动物雷帕霉素靶蛋白(mammalian target of rapamycin,mTOR)信号通路促进猪肠上皮细胞IPEC-J2信号转导与转录激活因子1(signal transducer and activator of transcription 1,STAT1)和干扰素刺激基因(interferon-stimulated genes,ISGs)的蛋白表达,预防猪传染性胃肠炎病毒感染[2]。Leu通过T细胞依赖途径、T细胞不依赖途径和肠道菌群的非排他性机制促进小鼠肠道sIgA的分泌[37]。Leu代谢物——β-羟基-β-甲基丁酸(β-hydroxy-β-methyl butyrate,HMB)显著提高初生犊牛血液中粒细胞趋化和吞噬活性[38]。高温、潮湿环境下母猪补充Val,能够提高仔猪血清IgM含量[39]。

2.2.2 降低炎症水平

BCAAs通过促进Foxp3+调节性T细胞(regulatory T cells,Tregs)增殖,改善Toll样受体4(Toll-like receptor 4,TLR4)信号通路,调节促炎因子和抗炎因子表达,降低炎症水平。BCAAs通过Foxp3+ Tregs细胞膜表面氨基酸转运体——溶质载体家族3成员2(solute carrier family 3 member 2,SLC3A2)/mTORC1信号通路,促进Foxp3+ Tregs增殖,减少小鼠胃肠道、肝脏等器官的炎症[40]。Ile通过改善结肠细胞TLR4/髓样分化因子88(myeloid differentiation factor 88,MyD88)/核转录因子-κB(nuclear factor-kappa B,NF-κB)通路,缓解葡聚糖硫酸钠(dextran sodium sulfate,DSS)诱导的大鼠生长发育迟缓和结肠损伤[41]。Ile显著降低肥胖小鼠肝脏中的促炎因子——啮齿动物单核细胞趋化蛋白1(monocyte chemoattractant protein 1,MCP1)、C-C基序趋化因子配体2(C-C motif chemokine ligand 2,CCL2)等表达;Val或Ile通过显著提高抗炎因子——白细胞介素-4(interleukin-4,IL-4)含量,减少肝脏的慢性炎症[3]。

2.3 提升抗氧化性能

BCAAs能够促进抗氧化酶表达,增加细胞能量供应,增强过氧化物酶体和线粒体功能等,提升抗氧化性能。Leu通过激活核因子红系2相关因子2(nuclear factor erythroid 2-related factor 2,Nrf2)信号通路,显著促进牛肠上皮细胞中超氧化物歧化酶2(superoxide dismutase 2,SOD2)等抗氧化酶和Ⅱ相解毒酶——血红素加氧酶1(heme oxygenase 1,HO1)mRNA和蛋白表达[42]。Leu通过调节丙酸盐代谢、BCAAs降解和产热途径,产生更多的ATP,保护牛乳腺上皮细胞免受过氧化氢(H2O2)诱导的氧化应激;Ile通过调节过氧化物酶体途径改善过氧化物酶体运输的缺陷,以及调节丙酸代谢途径促进乙酰辅酶A的产生,提高牛乳腺上皮细胞的抗氧化性能[43]。Val或Ile通过激活控制线粒体生物发生关键步骤的过氧化物酶体增殖物激活受体γ辅激活因子-1α(peroxisome proliferator-activated receptor γ coactivator-1α,PGC-1α)和簇状线粒体同源蛋白(clustered mitochondriahomolog protein,CLUH)表达,提高线粒体的功能和数量,减少肥胖小鼠肝脏的氧化应激和脂质过氧化[3]。

2.4 促进肌肉生成

BCAAs及其代谢物能够促进肌细胞的增殖和分化,改善肌肉纤维类型,增强蛋白质合成,抑制蛋白质降解,促进肌肉生成。Ile可以促进小鼠成肌细胞的增殖和分化[44]。Leu通过脂联素/腺苷酸活化蛋白激酶(AMP-activated protein kinase,AMPK)/PGC-1α[45]、Akt/叉头框蛋白O1(forkhead box 1,FoxO1)信号通路和微RNA-27a(microRNA-27a)[46]促进猪骨骼肌由快肌纤维向慢肌纤维转变;通过沉默交配型信息调节因子2同源蛋白1(silent mating type information regulation 2 homolog 1,Sirt1)/AMPK信号通路诱导猪骨骼肌卫星细胞慢肌纤维表达并改善线粒体功能[47]。Leu通过改变mTOR磷酸化位点、mTOR与调控相关蛋白——Raptor之间的相互作用,促进L6成肌细胞内蛋白质合成[1];通过上调新生雏鸡骨骼肌卫星细胞雷帕霉素靶蛋白(target of rapamycin,TOR)及其下游信号分子——核糖体蛋白S6激酶1(ribosomal protein S6 kinase 1,S6K1)和真核翻译起始因子4E结合蛋白1(eukaryotic translation initiation factor 4E binding protein 1,4E-BP1)mRNA表达,激活mTOR转录信号通路,刺激骨骼肌蛋白合成[48];通过促进应激诱导蛋白——Sestrin2/营养感应枢纽——GATOR2复合物解离,增加Rag A/C与mTOR结合,激活mTORC1,刺激新生仔猪骨骼肌蛋白质合成;HMB也能通过mTORC1信号通路刺激新生仔猪骨骼肌蛋白质合成[49]。Leu通过下调组蛋白去乙酰化酶4(histone deacetylase 4,HDAC4)mRNA表达,抑制大鼠骨骼肌萎缩[50];HMB通过改善线粒体氧化能力,抑制肌肉蛋白质降解[51]。

2.5 提高泌乳性能

BCAAs能够促进乳腺发育,增加乳蛋白、乳脂和乳糖含量,提高泌乳性能。Val通过调节mTOR和大鼠肉瘤蛋白(sarcoma,Ras)/细胞外信号调节激酶(extracellular signal-regulated kinase,ERK)信号通路,促进猪乳腺上皮细胞(porcine mammary epithelial cells,PMECs)生长和乳蛋白合成[52]。BCAAs通过刺激mTOR和抑制泛素-蛋白酶体信号通路,调节PMECs中的蛋白质合成和降解,促进净蛋白质合成[4]。Ile通过磷脂酰肌醇3激酶(phosphoinositide 3-kinase,PI3K)/Brahma相关基因1(Brahma related gene 1,BRG1)/mTOR/固醇调控元件结合蛋白1c(sterol regulatory element binding protein 1c,SREBP1c)通路刺激乳腺上皮细胞中乳蛋白和脂肪的合成[6]。妊娠后期饲粮添加Val通过Akt/mTOR/固醇调控元件结合蛋白1(sterol regulatory element binding protein 1,SREBP1)信号通路,促进母猪初乳中乳脂的合成[5];还能提高高温、潮湿环境下母猪初乳乳糖含量[39]。

2.6 改善脂肪代谢

BCAAs及其代谢产物能够调控脂肪代谢。Leu通过独立于mTORC1信号通路上调与脂肪酸代谢相关的基因表达,并经mTORC1依赖和mTORC1独立的途径上调线粒体生物合成,降低小鼠成肌细胞C2C12内的脂质[53];通过上调白色脂肪组织(white adipose tissue,WAT)褐变相关选择性标记物的基因和蛋白表达,下调脂肪酸合成相关基因的表达,改善小鼠脂质代谢[54];此外,KIC、HMB分别通过SREBP1c等转录因子、AMPK/mTOR信号通路,改变骨骼肌脂肪代谢,改善育肥猪肌肉脂肪酸组成[55]。Val或Ile通过失活SREBP1等与脂肪变性合成代谢相关的主调控因子,激活PGC-1α等与线粒体生物合成和酰基辅酶A氧化酶-1(acyl-coenzyme A oxidase 1,ACOX1)等与脂质分解代谢相关的主调控因子,减少肥胖小鼠肝脏脂肪积累[3]。但是,Liu等[44]研究发现,Ile通过上调脂肪酸合酶(fatty acid synthase,FAS)和过氧化物酶体增殖物激活受体γ(peroxisome proliferator-activated receptor γ,PPARγ)蛋白水平,显著下调脂肪甘油三酯脂酶(adipose triglyceride lipase,ATGL)和脂蛋白脂肪酶(lipoprotein lipase,LPL)等重要脂解基因的蛋白水平,造成线粒体功能障碍,促进高脂肪喂养小鼠骨骼肌细胞内脂肪沉积;Bishop等[56]研究发现,Val通过过氧化物酶体增殖物激活受体α(peroxisome proliferator-activated receptor α,PPARα)诱导的α氧化和增加丙酰辅酶A供应从头生成脂肪2种途径提高雄性小鼠肝脏和循环中奇链脂肪酸水平。Gart等[3]与Liu等[44]、Bishop等[56]的研究结果不一致,造成差异的原因可能与研究部位、BCAAs添加量等有关。

2.7 其他

Val能够促进小鼠造血干细胞的增殖[57];增强鸡精子的抗冻性和受精能力[58];促进神经突生长[59];还通过增强中国地鼠卵巢细胞(Chinese hamster ovary cell,CHO)中TCA循环,加速CHO中氨以丙氨酸的形式排泄,促进CHO的生长[60]。此外,Leu可以改善中年小鼠衰老诱导的血管重塑和功能障碍[61]。

3 BCAAs在猪生产中的应用

3.1 仔猪

对弱仔猪而言,Leu添加量越高,体重增长越快。Zhang等[62]在基础饲粮中额外添加0.35%Leu,结果发现,Leu能够通过增强肝脏mTOR磷酸化,缓解14~35日龄宫内发育迟缓(intrauterine growth retardation,IUGR)引起的“长大”仔猪(公母各占1/2)蛋白质代谢紊乱,改善血液参数,促进蛋白质合成。Bertocchi等[63]研究发现,初始体重(initial body weight,IBW)为7.9 kg的轻型仔猪,在38~59日龄饲粮中标准回肠可消化(standard ileal digestible,SID)Leu∶SID赖氨酸(lysine,Lys)=1.105~1.154时,平均日增重最高;IBW为10.5 kg的重型仔猪,在38~59日龄饲粮中SID Leu∶SID Lys=0.949~1.102时,平均日增重最高。
在正常和低蛋白质饲粮中,适宜比例的BCAAs都能够促进仔猪的生长。Habibi等[12]研究发现,在29~42日龄和43~63日龄“杜长大”阉公猪2阶段低蛋白质饲粮(粗蛋白质14% vs. 20%,SID Val、SID Ile分别为0.50%、0.43%;粗蛋白质13% vs. 19%,SID Val、SID Ile分别为0.46%、0.37%)中添加Val和Ile,结果发现,SID Val、SID Ile分别为0.81%、0.73%以及0.77%、0.68%时,低蛋白质组能够通过改善肠道发育、肝脏IGF1表达和生物利用度以及血浆代谢组学特征,逆转低蛋白质饲粮引起的生长性能下降。Stas等[64]研究发现,发酵玉米蛋白(fermented corn protein,FCP)中支链氨基酸不平衡,Leu比例过大,饲粮中添加FCP,如果不平衡BCAAs,会降低仔猪的生长性能;当FCP添加量为10%时,高SID Val+SID Ile∶SID Lys饲粮(1.09 vs. 1.00)显著提高45~66日龄仔猪饲粮效率;当FCP添加量为20%时,高SID Val+SID Ile∶SID Lys饲粮(1.04 vs. 0.88)也具有相似的效果。

3.2 育肥猪

饲粮中额外添加Leu能够缓解育肥猪应激。Yin等[14]在“杜长大”育肥公猪(IBW为68.33 kg)饲粮(SID Leu为1.12%)中添加0.5%Leu,试验期42 d,结果发现,Leu通过Kelch样ECH关联蛋白1(Kelch-like ECH-associated protein 1,Keap1)/Nrf2和PGC-1α/线粒体转录因子A(mitochondrial transcription factor A,TFAM)信号通路改善热应激育肥公猪的生长性能和肉品质。Rudar等[65]研究发现,当免疫系统刺激的大白阉公猪(IBW为14.5 kg)饲粮中SID Leu(2.72%)为NRC(2012)推荐水平的200%时,有降低肝脏蛋白质合成率和增加骨骼肌——腓肠肌蛋白合成率的趋势;在免疫系统刺激期间,补充高于估计需求的Leu(1.36%)可能支持氨基酸从内脏到外周蛋白质沉积的再分配。
在含有谷物干酒糟及其可溶物(distillers dried grains with solubles,DDGS)的饲粮中添加Val和Ile能够缓解高Leu带来的负面影响,提高育肥猪的生长性能。Kerkaert等[66]研究发现,分别在33.6~50.8 kg、50.8~74.9 kg、74.9~90.0 kg 3阶段均添加30%DDGS的饲粮(SID Val分别为0.69%、0.61%、0.53%)中添加Val,结果发现,3阶段饲粮中SID Val分别为0.74%、0.67%、0.59%时,育肥猪(PIC TR4×Fast LW×PIC L02,阉公猪、母猪各占1/2)平均日增重、平均日采食量和饲料效率均有提高的趋势,生长性能与高蛋白质饲粮组接近;分别在90.0~98.5 kg(DDGS为30%,SID Ile为0.44%)、98.5~136.0 kg(DDGS为20%,SID Ile为0.39%)2阶段饲粮中添加Ile,结果发现,当2阶段饲粮中SID Ile分别为0.51%、0.46%时,90.0~136.0 kg育肥猪具有相似的试验结果。Clizer等[67]研究发现,在39~68 kg育肥猪(DNA600×Topigs Norsvin 70,阉公猪、母猪各占1/2)饲粮(DDGS为30%,SID Val为0.59%)中添加Val,结果发现,当SID Val=0.67%(SID Val∶SID Lys=0.68)时,获得最大日增重和饲料效率的置信概率分别达到99%和97%以上。Clizer等[68]研究发现,82.3~130.0 kg育肥猪(DNA600×Topigs Norsvin 70,阉公猪、母猪各占1/2)饲粮(DDGS为20%)中SID Ile∶SID Lys=0.70时,可获得与高蛋白质玉米大豆饲粮(DDGS为20%)相似的生长性能。
适当比例的BCAAs能够降低脂肪含量,提高猪肉品质。Zhang等[69]研究发现,59.11~95.00 kg“杜长大”阉公猪低蛋白质饲粮(粗蛋白质12% vs. 16%)中,SID Leu∶SID Val∶SID Ile=2∶1∶1~2∶2∶1(推荐2∶2∶1)时,能够通过脂肪因子/AMPK/Sirt1/PPARγ通路,抑制脂肪酸、甘油三酯合成,加速脂肪酸氧化。Xu等[70]研究发现,与高、低比例组(SID Val∶SID Ile分别为2.60、0.58)相比,75~100 kg“杜长大”阉公猪低蛋白质饲粮(粗蛋白质为10.39%,SID Val为0.48%)中SID Val∶SID Ile=1.23时能够显著降低背膘厚度,提高无脂瘦肉指数,增加风味氨基酸含量,但显著降低肌内脂肪含量,增加熟肉的剪切力;同样,与高、低比例组(SID Val∶SID Ile分别为2.39、0.70)相比,100~135 kg“杜长大”阉公猪饲粮(粗蛋白质为9.97%,SID Val为0.41%)中SID Val∶SID Ile=1.24时,也得到相似的试验结果;且与较重的猪(130 kg)相比,较轻的猪(105 kg)的胴体性状和肉色更易受SID Val∶SID Ile的影响。

3.3 母猪

高温状态下,BCAAs能够促进母猪乳腺发育,提高乳汁品质,增强母猪胎盘代谢。Zhao等[39]在产前29 d~产后21 d“长大”母猪饲粮(Lys为0.96%,Val为0.72%)中添加Val,结果发现,当Val∶Lys=1.01时,可通过提高母猪妊娠血糖、初乳乳糖和仔猪血清IgM含量,改善高温、潮湿条件下母猪和仔猪的生产性能;Che等[71]研究发现,妊娠75 d~分娩“长大”母猪分别饲喂低Val(Val∶Lys=0.63∶1.00,Lys为0.85%)、中Val(Val∶Lys=0.73∶1.00)、高Val(Val∶Lys=0.93∶1.00)3种饲粮,结果发现,Val能够通过上调乳腺组织中乙酰辅酶A羧化酶α等脂肪酸合成相关蛋白的表达来改善乳脂合成;Che等[72]在妊娠75 d~分娩“长大”母猪饲粮(Lys为0.85%,Val为0.54%)中添加Val,结果发现,当Lys∶Val=0.93时,能够通过促进母猪乳腺发育,显著提高断奶仔猪体重和平均日增重,同时增加母猪采食量。但是,Greiner等[73]在妊娠112 d~哺乳23 d期间饲粮(SID Lys为0.95%,SID Val为0.48%)中添加Val,结果发现,Val(SID Val∶SID Lys=0.50~1.00)没有改变母猪繁殖性能或仔猪生长速度。出现不同试验结果的原因可能与饲养温度有关,Zhao等[39](22~31 ℃)、Che等[71](30.4~32.3 ℃)高于Greiner等[73](18~24 ℃)的饲养温度,高温状态下母猪的代谢功能可能与常温下不同。Cui等[74]在妊娠90 d~分娩“长大”母猪饲粮(Leu为1.30%)中额外添加0.4%~0.8%Leu,饲养温度为30~32 ℃,结果发现,Leu会全面改变胎盘代谢,并通过PI3K/AKT/mTOR信号通路促进母胎营养物质(氨基酸、葡萄糖和脂肪酸)运输。

4 BCAAs在鸡生产中的应用

4.1 肉鸡

现代白羽肉鸡对低蛋白质饲粮中BCAAs的需要量有所增加。Sedghi等[75]采用正交L16(43)田口法(Taguchi method,TM)优化1~7日龄雄性罗斯308肉鸡饲粮SID Leu、SID Val、SID Ile分别为1.345%、1.045%、0.850%时,平均日增重和料重比最佳,并在19.93%粗蛋白质水平上验证确认。Kriseldi等[76]采用23全因子中心复合设计(central composite design,CCD)优化20~34日龄雄性罗斯344×708肉鸡低蛋白质饲粮(粗蛋白质为18%)可消化(digestible,Dig)Leu、Dig Val、Dig Ile分别为1.21%、0.86%~0.87%、0.72%~0.73%时,体增重最高,料重比最低;并认为低BCAAs∶Lys可改善体增重和采食量,而高BCAAs∶Lys可使屠宰率最大化。Ospina-Rojas等[77]研究发现,雄性科宝肉鸡1~14日龄低蛋白质饲粮(粗蛋白质为19.2%)中SID Leu、SID Val、SID Ile分别为1.37%、0.94%、0.87%,15~28日龄(粗蛋白质为17.8%)分别为1.23%、0.82%、0.75%,29~42日龄(粗蛋白质为16%)分别为1.15%、0.77%、0.70%时,体增重最高;并认为在预估低蛋白质饲粮中理想Val和Ile水平时,应考虑Leu水平。Sedghi等[75]、Kriseldi等[76]、Ospina-Rojas等[77]研究的肉鸡对Leu、Val和Ile的需要高于《鸡饲养标准》(NY/T 33—2004)[78]、NRC(1994)[79],与罗斯肉鸡(2022)[80]推荐的营养需要接近,原因可能是一方面育种技术提高了肉鸡的生长速度,肉鸡对饲粮中氨基酸的需求增加;另一方面,低蛋白质饲粮增加了肉鸡对BCAAs的需要量。
BCAAs比例对胸肌率、腹部脂肪、胫骨以及不同性别肉鸡的生长速度都有一定的影响。Kriseldi等[76]研究发现,在低蛋白质饲粮(粗蛋白质为18%)中Dig Leu∶Dig Lys=1.70~1.90时,提高Dig Ile∶Dig Lys(0.56~0.68)可提高20~34日龄雄性罗斯344×708肉鸡胸肌率,但提高Dig Val∶Dig Lys(0.64~0.76)对胸肌率没有影响;Agostini等[81]也发现,低蛋白质饲粮(1~12日龄粗蛋白质为19%,13~28日龄粗蛋白质为17.5%,29~42日龄粗蛋白质为16%)中Dig Val∶Dig Lys=0.63~0.93时,对1~42日龄雄性科宝500肉鸡的胴体率和胸肌率没有影响。Chrystal等[13]研究发现,低蛋白质饲粮(粗蛋白质为17.5%)中提高Dig Leu(1.25%~1.83%)会抑制7~28日龄雄性罗斯308阉割肉鸡体增重,但当Dig Leu为1.499%时,料重比最低;同时,增加Val和Ile会降低腹部脂肪的相对重量,当Dig Ile为1.09%、Dig Val为1.25%时,腹部脂肪的相对重量最轻。Ospina-Rojas等[82]研究发现,1~21日龄雄性科宝500肉仔鸡低蛋白质饲粮(粗蛋白质为18.5%)中SID Leu、SID Val分别为1.42%、0.90%时,肉仔鸡胫骨断裂强度在21日龄达到最大;此外,Leu能够减少1~21日龄肉仔鸡软骨肥大。Kidd等[83]研究发现,22~35日龄雌性洛曼印第安河肉鸡对BCAAs比例的敏感性高于雄性,当雌性肉鸡饲粮中Leu提高时,需要提高Ile来抵消负面影响。

4.2 蛋鸡

适宜比例的BCAAs能够增强蛋鸡肠道屏障,降低炎症反应,提高产蛋性能。Jian等[15]在34~42周龄丰达1号蛋鸡饲粮(Val为0.59%)添加Val,结果发现,Val为0.64%~0.69%时,可以改善肠道绒毛形态,增强肠道屏障,降低盲肠致病菌丰度,抑制一般控制非抑制2(general control nonderepressible 2,GCN2)激酶介导的炎症反应,改善肠道健康;但是,长期摄入0.74%~0.79%的Val可促进GCN2激酶介导的肝脏脂肪生成,抑制脂肪酸氧化,加速蛋鸡非酒精性脂肪肝的发展。Wen等[84]研究发现,随着低蛋白质饲粮(粗蛋白质为13.38%)中Val添加量的提高(0.515%~0.865%),41~60周龄海兰蛋鸡(品系W-36)产蛋率极显著上升;采用线性折线模型,以蛋重、产蛋量、料蛋比为指标,Val的需要量分别为597.3、591.9、500.5 mg/d。Liaqat等[85]在33~40周龄罗曼蛋鸡(品系LSL-LITE)饲粮(Dig Ile为0.72%,Dig Val为0.72%)中添加Val,结果发现,Dig Val为0.87%时,能够显著提高产蛋率,降低料蛋比。Ullah等[86]在23~30周龄罗曼蛋鸡(品系LSL-LITE)饲粮(Ile为0.66%)中添加Ile,结果发现,Ile为0.72%时,料蛋比最低。

5 小结与展望

综上所述,BCAAs参与和调控糖类、脂肪和蛋白质代谢,具有刺激蛋白质合成、增强免疫功能等多种生物学作用;在猪和鸡生产中能够降低饲粮粗蛋白质水平、抗应激和提高生产性能等。然而,Val调控脂肪代谢的机理还不完善,高温下BCAAs提高母猪生产性能的原因和机制等还缺乏了解,需要在将来进一步探究为BCAAs合理应用提供基础理论;此外,考虑到BCAAs与Lys的比例、与色氨酸的拮抗作用以及BCAAs之间的协同和拮抗作用,将来还需通过科学、合理的饲粮设计,确定不同畜禽品种、养殖目标下BCAAs的适宜添加量和添加比例。

参考文献

原文顺序 | 文献年度倒序 | 文中引用次数倒序
[1]
ROBERSON P A, JEFFERSON L S, KIMBALL S R. Convergence of signaling pathways in mediating actions of leucine and IGF-1 on mTORC1 in L6 myoblasts[J]. American Journal of Physiology:Cell Physiology, 2022, 323(3):C804-C812.

[2]
DU J, CHEN D W, YU B, et al. L-leucine promotes STAT1 and ISGs expression in TGEV-infected IPEC-J2 cells via mTOR activation[J]. Frontiers in Immunology, 2021, 12:656573.

DOI

[3]
GART E, VAN DUYVENVOORDE W, CASPERS M P M, et al. Intervention with isoleucine or valine corrects hyperinsulinemia and reduces intrahepatic diacylglycerols,liver steatosis,and inflammation in Ldlr-/-.Leiden mice with manifest obesity-associated NASH[J]. The FASEB Journal, 2022, 36(8):e22435.

[4]
REZAEI R, WU G Y. Branched-chain amino acids regulate intracellular protein turnover in porcine mammary epithelial cells[J]. Amino Acids, 2022, 54(11):1491-1504.

DOI PMID

[5]
CHE L, XU M M, GAO K G, et al. Valine increases milk fat synthesis in mammary gland of gilts through stimulating AKT/MTOR/SREBP1 pathway[J]. Biology of Reproduction, 2019, 101(1):126-137.

DOI

[6]
HAO Q, WANG Z, WANG L L, et al. Isoleucine stimulates mTOR and SREBP-1c gene expression for milk synthesis in mammary epithelial cells through BRG1-mediated chromatin remodelling[J]. British Journal of Nutrition, 2023, 129(4):553-563.

DOI

[7]
KWON W B, TOUCHETTE K J, SIMONGIOVANNI A, et al. Excess dietary leucine in diets for growing pigs reduces growth performance,biological value of protein,protein retention,and serotonin synthesis1[J]. Journal of Animal Science, 2019, 97(10):4282-4292.

DOI

[8]
KWON W B, SOTO J A, STEIN H H. Effects of dietary leucine and tryptophan on serotonin metabolism and growth performance of growing pigs[J]. Journal of Animal Science, 2022, 100(1):skab356.

[9]
ZHANG Y F, XIONG H B, CHEN Z C, et al. Effect of fed-batch and chemostat cultivation processes of C. glutamicum CP for L-leucine production[J]. Bioengineered, 2021, 12(1):426-439.

DOI

[10]
赵阔, 程金宇, 郭亮, 等. 谷氨酸棒杆菌代谢工程高效生产L-缬氨酸[J/OL]. 生物工程学报:1-23[2023-03-10].https://doi.org/10.13345/j.cjb.220998.

ZHAO K, CHENG J Y, GUO L, et al. Highly efficient production of L-valine by multiplex metabolic engineering of Corynebacterium glutamicum[J/OL]. Chinese Journal of Biotechnology:1-23[2023-03-10].https://doi.org/10.13345/j.cjb.220998. (in Chinese)

[11]
孔帅, 陈敏, 郑美娟, 等. 常温常压等离子体诱变选育高产L-异亮氨酸谷氨酸棒杆菌[J]. 中国酿造, 2019, 38(7):76-79.

KONG S, CHEN M, ZHENG M J, et al. Mutation breeding of high-yield L-isoleucine Corynebacterium glutamicum by atmospheric and room temperature plasmas mutagenesis[J]. China Brewing, 2019, 38(7):76-79. (in Chinese)

[12]
HABIBI M, GOODARZI P, SHILI C N, et al. A mixture of valine and isoleucine restores the growth of protein-restricted pigs likely through improved gut development,hepatic IGF-1 pathway,and plasma metabolomic profile[J]. International Journal of Molecular Sciences, 2022, 23(6):3300.

DOI

[13]
CHRYSTAL P V, GREENHALGH S, MACELLINE S P, et al. A multivariate Box-Behnken assessment of elevated branched-chain amino acid concentrations in reduced crude protein diets offered to male broiler chickens[J]. PLoS One, 2022, 17(3):e0266080.

[14]
YIN Y J, LIU Y T, DUAN G Y, et al. The effect of dietary leucine supplementation on antioxidant capacity and meat quality of finishing pigs under heat stress[J]. Antioxidants, 2022, 11(7):1373.

DOI

[15]
JIAN H F, MIAO S S, LIU Y T, et al. Dietary valine ameliorated gut health and accelerated the development of nonalcoholic fatty liver disease of laying hens[J]. Oxidative Medicine and Cellular Longevity, 2021, 2021:4704771.

[16]
SUGAHARA M, MORIMOTO T, KOBAYASHI T, et al. The nutritional value of D-amino acid in the chick nutrition[J]. Agricultural and Biological Chemistry, 1967, 31(1):77-84.

DOI

[17]
冯子坛, 陈元方, 温淑豪, 等. 研究大鼠小肠氨基酸吸收的实验模型[J]. 中国医学科学院学报, 1989(3):174.

FENG Z T, CHEN Y F, WEN S H, et al. Research of the experimental model of amino acid absorption in rat small intestine[J]. Acta Academiae Medicinae Sinicae, 1989(3):174. (in Chinese)

[18]
LEBZIEN P, ROHR K. Bestimmung der wahren verdaulichkeit von aminosäuren aus dem dünndarm von milchkühen[J]. Archiv für Tierernaehrung, 1994, 45(4):355-362.

DOI

[19]
YOUNOSZAI M K. Intestinal maturation:effect of growth retardation on L-valine absorption[J]. American Journal of Clinical Nutrition, 1984, 40(6):1243-1249.

DOI

[20]
ZHANG S H, QIAO S Y, REN M, et al. Supplementation with branched-chain amino acids to a low-protein diet regulates intestinal expression of amino acid and peptide transporters in weanling pigs[J]. Amino Acids, 2013, 45(5):1191-1205.

DOI PMID

[21]
张宇峰, 田敏, 恒景会, 等. 支链氨基酸对断奶仔猪肠道和肌肉氨基酸转运载体表达的影响[J]. 中国畜牧杂志, 2020, 56(4):92-99.

ZHANG Y F, TIAN M, HENG J H, et al. Effects of branched-chain amino acids on intestinal and muscular amino acid transporter expression of weaning piglets[J]. Chinese Journal of Animal Science, 2020, 56(4):92-99. (in Chinese)

[22]
TEN HAVE G A M, JANSEN L, SCHOONEMAN M G, et al. Metabolic flux analysis of branched-chain amino and keto acids (BCAA,BCKA) and β-hydroxy β-methylbutyric acid across multiple organs in the pig[J]. American Journal of Physiology:Endocrinology and Metabolism, 2021, 320(3):E629-E640.

[23]
WILTAFSKY M K, PFAFFL M W, ROTH F X. The effects of branched-chain amino acid interactions on growth performance,blood metabolites,enzyme kinetics and transcriptomics in weaned pigs[J]. British Journal of Nutrition, 2010, 103(7):964-976.

DOI

[24]
SON S M, PARK S J, LEE H, et al. Leucine signals to mTORC1 via its metabolite acetyl-coenzyme A[J]. Cell Metabolism, 2019, 29(1):192-201.e7.

DOI PMID

[25]
ADEVA-ANDANY M M, LÓPEZ-MASIDE L, DONAPETRY-GARCÍA C, et al. Enzymes involved in branched-chain amino acid metabolism in humans[J]. Amino Acids, 2017, 49(6):1005-1028.

DOI

[26]
JOHANSEN M L, BAK L K, SCHOUSBOE A, et al. The metabolic role of isoleucine in detoxification of ammonia in cultured mouse neurons and astrocytes[J]. Neurochemistry International, 2007, 50(7/8):1042-1051.

DOI

[27]
DAI J, WANG H, LIAN Y, et al. Non-targeted metabolomic analysis of chicken kidneys in response to coronavirus IBV infection under stress induced by dexamethasone[J]. Frontiers in Cellular and Infection Microbiology, 2022, 12:945865.

DOI

[28]
BI S C, SHAO J J, QU Y W, et al. Hepatic transcriptomics and metabolomics indicated pathways associated with immune stress of broilers induced by lipopolysaccharide[J]. Poultry Science, 2022, 101(12):102199.

DOI

[29]
SENER A, MALAISSE W J. L-leucine and a nonmetabolized analogue activate pancreatic islet glutamate dehydrogenase[J]. Nature, 1980, 288(5787):187-189.

DOI

[30]
PANTEN U. Effects of alpha-ketomonocarboxylic acids upon insulin secretion and metabolism of isolated pancreatic islets[J]. Naunyn-Schmiedeberg’s Archives of Pharmacology, 1975, 291(4):405-420.

DOI

[31]
常银莲. 支链氨基酸对肉鸡小肠发育及TOR通路相关基因表达的影响[D]. 硕士学位论文. 北京: 中国农业科学院, 2016.

CHANG Y L. Effects of BCAAs on small intestinal development and relative genes expression of TOR pathway in broilers[D]. Master’s Thesis. Beijing: Chinese Academy of Agricultural Sciences, 2016. (in Chinese)

[32]
GUO L, TIAN H B, YAO J H, et al. Leucine improves α-amylase secretion through the general secretory signaling pathway in pancreatic acinar cells of dairy calves[J]. American Journal of Physiology:Cell Physiology, 2020, 318(6):C1284-C1293.

[33]
GOODARZI P, WILEMAN C M, HABIBI M, et al. Effect of isoleucine and added valine on performance,nutrients digestibility and gut microbiota composition of pigs fed with very low protein diets[J]. International Journal of Molecular Sciences, 2022, 23(23):14886.

DOI

[34]
TIAN M, HENG J H, SONG H Q, et al. Branched chain amino acids stimulate gut satiety hormone cholecystokinin secretion through activation of the umami taste receptor T1R1/T1R3 using an in vitro porcine jejunum model[J]. Food&Function, 2019, 10(6):3356-3367.

[35]
YOUNGBLOOD B D, SMAGIN G N, ELKINS P D, et al. The effects of paradoxical sleep deprivation and valine on spatial learning and brain 5-HT metabolism[J]. Physiology&Behavior, 1999, 67(5):643-649.

[36]
BISWAS D, DAO K T, MERCER A, et al. Branched-chain ketoacid overload inhibits insulin action in the muscle[J]. Journal of Biological Chemistry, 2020, 295(46):15597-15621.

DOI

[37]
SONG B, ZHENG C, ZHA C, et al. Dietary leucine supplementation improves intestinal health of mice through intestinal SIgA secretion[J]. Journal of Applied Microbiology, 2020, 128(2):574-583.

DOI PMID

[38]
WÓJCIK R, MAŁACZEWSKA J, ZWIERZCHOWSKI G, et al. The influence of dietary supplementation with the leucine metabolite β-hydroxy-β-methylbutyrate (HMB) on the chemotaxis,phagocytosis and respiratory burst of peripheral blood granulocytes and monocytes in calves[J]. BMC Veterinary Research, 2020, 16(1):171.

DOI

[39]
ZHAO L Y, LI Y S, LI Z J, et al. Effect of the valine-to-lysine ratio on the performance of sows and piglets in a hot,humid environment[J]. Journal of Thermal Biology, 2019, 81:89-97.

DOI

[40]
IKEDA K, KINOSHITA M, KAYAMA H, et al. Slc3a2 mediates branched-chain amino-acid-dependent maintenance of regulatory T cells[J]. Cell Reports, 2017, 21(7):1824-1838.

DOI PMID

[41]
MAO X B, SUN R, WANG Q X, et al. L-isoleucine administration alleviates DSS-induced colitis by regulating TLR4/MyD88/NF-κB pathway in rats[J]. Frontiers in Immunology, 2022, 12:817583.

DOI

[42]
MAO H L, ZHANG Y F, JI W W, et al. Leucine protects bovine intestinal epithelial cells from hydrogen peroxide-induced apoptosis by alleviating oxidative damage[J]. Journal of the Science of Food and Agriculture, 2022, 102(13):5903-5912.

DOI

[43]
WU S N, LIU X Z, CHENG L, et al. Protective mechanism of leucine and isoleucine against H2O2-induced oxidative damage in bovine mammary epithelial cells[J]. Oxidative Medicine and Cellular Longevity, 2022, 2022:4013575.

[44]
LIU S G, SUN Y M, ZHAO R, et al. Isoleucine increases muscle mass through promoting myogenesis and intramyocellular fat deposition[J]. Food&Function, 2021, 12(1):144-153.

[45]
CHEN X L, XIANG L, HUANG Z Q, et al. Effect of dietary leucine supplementation on skeletal muscle fiber type transformation in weaning piglets[J]. Animal Biotechnology, 2022, 33(3):546-554.

DOI

[46]
ZHANG S R, CHEN X L, HUANG Z Q, et al. Leucine promotes porcine myofibre type transformation from fast-twitch to slow-twitch through the protein kinase B (Akt)/forkhead box 1 signalling pathway and microRNA-27a[J]. British Journal of Nutrition, 2019, 121(1):1-8.

DOI

[47]
CHEN X L, XIANG L, JIA G, et al. Leucine regulates slow-twitch muscle fibers expression and mitochondrial function by Sirt1/AMPK signaling in porcine skeletal muscle satellite cells[J]. Animal Science Journal, 2019, 90(2):255-263.

DOI PMID

[48]
SHE Y, DENG H L, CAI H Y, et al. Regulation of the expression of key signalling molecules in mTOR pathway of skeletal muscle satellite cells in neonatal chicks:effects of leucine and glycine-leucine peptide[J]. Journal of Animal Physiology and Animal Nutrition, 2019, 103(3):786-790.

DOI

[49]
SURYAWAN A, RUDAR M, FIOROTTO M L, et al. Differential regulation of mTORC1 activation by leucine and β-hydroxy-β-methylbutyrate in skeletal muscle of neonatal pigs[J]. Journal of Applied Physiology, 2020, 128(2):286-295.

DOI PMID

[50]
ALVES P K N, CRUZ A, SILVA W J, et al. Leucine supplementation decreases HDAC4 expression and nuclear localization in skeletal muscle fiber of rats submitted to hindlimb immobilization[J]. Cells, 2020, 9(12):2582.

DOI

[51]
DUAN Y H, ZHONG Y Z, SONG B, et al. Suppression of protein degradation by leucine requires its conversion to β-hydroxy-β-methyl butyrate in C2C12 myotubes[J]. Aging, 2019, 11(24):11922-11936.

DOI

[52]
ZHANG J M, HE W L, YI D, et al. Regulation of protein synthesis in porcine mammary epithelial cells by L-valine[J]. Amino Acids, 2019, 51(4):717-726.

DOI

[53]
WU H X U, DRIDI S, HUANG Y, et al. Leucine decreases intramyocellular lipid deposition in an mTORC1-independent manner in palmitate-treated C2C12 myotubes[J]. American Journal of Physiology:Endocrinology and Metabolism, 2020, 318(2):E152-E163.

[54]
ZHOU X B, CHEN J Y, SUN B, et al. Leucine,but not isoleucine or valine,affects serum lipid profiles and browning of WAT in mice[J]. Food&Function, 2021, 12(15):6712-6724.

[55]
ZHONG Y Z, SONG B, ZHENG C B, et al. α-ketoisocaproate and β-hydroxy-β-methyl butyrate regulate fatty acid composition and lipid metabolism in skeletal muscle of growing pigs[J]. Journal of Animal Physiology and Animal Nutrition, 2019, 103(3):846-857.

DOI PMID

[56]
BISHOP C A, SCHULZE M B, KLAUS S, et al. The branched-chain amino acids valine and leucine have differential effects on hepatic lipid metabolism[J]. The FASEB Journal, 2020, 34(7):9727-9739.

DOI

[57]
TAYA Y, OTA Y, WILKINSON A C, et al. Depleting dietary valine permits nonmyeloablative mouse hematopoietic stem cell transplantation[J]. Science, 2016, 354(6316):1152-1155.

PMID

[58]
BERNAL B, IGLESIAS-CABEZA N, SÁNCHEZ-RIVERA U, et al. Effect of supplementation of valine to chicken extender on sperm cryoresistance and post-thaw fertilization capacity[J]. Poultry Science, 2020, 99(12):7133-7141.

DOI PMID

[59]
KOGA T, TAI A. Neurite outgrowth-promoting compounds from cockscomb hydrolysate[J]. Nutrients, 2022, 14(7):1422.

DOI

[60]
SAVIZI I S P, MAGHSOUDI N, MOTAMEDIAN E, et al. Valine feeding reduces ammonia production through rearrangement of metabolic fluxes in central carbon metabolism of CHO cells[J]. Applied Microbiology and Biotechnology, 2022, 106(3):1113-1126.

DOI

[61]
HAO Z J, XU G W, YUAN M Y, et al. Leucine supplementation in middle-aged male mice improved aging-induced vascular remodeling and dysfunction via activating the Sirt1-Foxo1 axis[J]. Nutrients, 2022, 14(18):3856.

DOI

[62]
ZHANG J F, XU W, YANG Y X, et al. Leucine alters blood parameters and regulates hepatic protein synthesis via mammalian/mechanistic target of rapamycin activation in intrauterine growth-restricted piglets[J]. Journal of Animal Science, 2022, 100(4):skac109.

[63]
BERTOCCHI M, BOSI P, LUISE D, et al. Dose-response of different dietary leucine levels on growth performance and amino acid metabolism in piglets differing for aminoadipate-semialdehyde synthase genotypes[J]. Scientific Reports, 2019, 9(1):18496.

DOI PMID

[64]
STAS E B, CHANCE J A, GOODBAND R D, et al. Evaluation of fermented corn protein and its effects in either high or low branch chain amino acid to leucine ratio diets on nursery pig performance and feed intake preference[J]. Translational Animal Science, 2022, 6(4):txac134.

[65]
RUDAR M, HUBER L A, ZHU C L, et al. Effects of dietary leucine supplementation and immune system stimulation on plasma AA concentrations and tissue protein synthesis in starter pigs[J]. Journal of Animal Science, 2019, 97(2):829-838.

DOI PMID

[66]
KERKAERT H R, CEMIN H S, WOODWORTH J C, et al. Improving performance of finishing pigs with added valine,isoleucine,and tryptophan:validating a meta-analysis model[J]. Journal of Animal Science, 2021, 99(1):skab006.

[67]
CLIZER D A, TOSTENSON B J, TAUER S K, et al. Impact of increasing standardized ileal digestible valine:lysine in diets containing 30% dried distiller grains with solubles on growing pig performance[J]. Journal of Animal Science, 2022, 100(9):skac228.

[68]
CLIZER D A, TOSTENSON B J, TAUER S K, et al. The effect of standardized ileal digestible isoleucine:lysine in diets containing 20% dried distillers grains with solubles on finishing pig performance and carcass characteristics[J]. Journal of Animal Science, 2022, 100(9):skac234.

[69]
ZHANG L Y, LI F N, GUO Q P, et al. Different proportions of branched-chain amino acids modulate lipid metabolism in a finishing pig model[J]. Journal of Agricultural and Food Chemistry, 2021, 69(25):7037-7048.

DOI PMID

[70]
XU D D, WANG Y B, ZHANG X, et al. Dietary valine/isoleucine ratio impact carcass characteristics,meat edible quality and nutritional values in finishing crossbred Duroc×Landrace×Yorkshire pigs with different slaughter weights[J]. Frontiers in Nutrition, 2022, 9:899871.

DOI

[71]
CHE L, XU M M, GAO K G, et al. Mammary tissue proteomics in a pig model indicates that dietary valine supplementation increases milk fat content via increased de novo synthesis of fatty acid[J]. Food Science&Nutrition, 2021, 9(11):6213-6223.

[72]
CHE L, XU M M, GAO K G, et al. Effects of dietary valine supplementation during late gestation on the reproductive performance and mammary gland development of gilts[J]. Journal of Animal Science and Biotechnology, 2020, 11(1):15.

DOI

[73]
GREINER L, GRAHAM A, GONCALVES M, et al. Evaluation of the optimal standardized ileal digestible valine:lysine ratio in lactating sow diets1[J]. Journal of Animal Science, 2019, 97(7):2965-2971.

DOI PMID

[74]
CUI C, WU C C, WANG J, et al. Leucine supplementation during late gestation globally alters placental metabolism and nutrient transport via modulation of the PI3K/AKT/mTOR signaling pathway in sows[J]. Food&Function, 2022, 13(4):2083-2097.

[75]
SEDGHI M, GOLIAN A, KOLAHAN F, et al. Optimisation of broiler chicken responses from 0 to 7 d of age to dietary leucine,isoleucine and valine using Taguchi and mathematical methods[J]. British Poultry Science, 2015, 56(6):696-707.

DOI

[76]
KRISELDI R, SILVA M, LEE J, et al. Understanding the interactive effects of dietary leucine with isoleucine and valine in the modern commercial broiler[J]. Poultry Science, 2022, 101(12):102140.

DOI

[77]
OSPINA-ROJAS I C, POZZA P C, RODRIGUEIRO R J B, et al. High leucine levels affecting valine and isoleucine recommendations in low-protein diets for broiler chickens[J]. Poultry Science, 2020, 99(11):5946-5959.

DOI

[78]
中华人民共和国农业部. 中华人民共和国农业行业标准-鸡饲养标准(NY/T 33—2004)[J]. 湖南饲料, 2006(4):19-27.

Ministry of Agriculture of the PRC. Agricultural industry standard of the People’s Republic of China-Feeding standard of chicken (NY/T 33—2004)[J]. Hunan Feed, 2006(4):19-27. (in Chinese)

[79]
张宏福. 美国NRC禽类营养需要量(1994年版)[J]. 中国饲料, 1995(2):36-38.

ZHANG H F. American national research council (NRC) nutrient requirements of poultry (1994)[J]. China Feed, 1995(2):36-38. (in Chinese)

[80]
Aviagen. ROSS broiler:nutrition specifications-all plant protein-based feeds[EB/OL].(2022-10-03)[2023-03-15].http://en.aviagen.com/assets/Tech_Center/Ross_Broiler/Ross-PlantProteinBasedBroilerNutritionSpecifications2022-EN.pdf.

[81]
AGOSTINI P S, SANTOS R R, KHAN D R, et al. The optimum valine:lysine ratios on performance and carcass traits of male broilers based on different regression approaches[J]. Poultry Science, 2019, 98(3):1310-1320.

DOI

[82]
OSPINA-ROJAS I C, MURAKAMI A E, DUARTE C R A, et al. Tibiotarsus bone characteristics and tibial dyschondroplasia incidence of broilers fed diets supplemented with leucine and valine[J]. Journal of Animal Physiology and Animal Nutrition, 2018, 102(2):e768-e776.

[83]
KIDD M T, POERNAMA F, WIBOWO T, et al. Dietary branched-chain amino acid assessment in broilers from 22 to 35 days of age[J]. Journal of Animal Science and Biotechnology, 2021, 12(1):6.

DOI

[84]
WEN J, HELMBRECHT A, ELLIOT M A, et al. Evaluation of the valine requirement of small-framed first cycle laying hens[J]. Poultry Science, 2019, 98(3):1272-1279.

DOI PMID

[85]
LIAQAT U, DITTA Y, NAVEED S, et al. Effects of L-valine in layer diets containing 0.72% isoleucine[J]. PLoS One, 2022, 17(4):e0258250.

[86]
ULLAH S, DITTA Y A, KING A J, et al. Varying isoleucine level to determine effects on performance,egg quality,serum biochemistry,and ileal protein digestibility in diets of young laying hens[J]. PLoS One, 2022, 17(1):e0261159.

Options
文章导航

/

〈 〉

    联系我们

  • 地址：北京市海淀区圆明园西路2号
    　　　中国农业大学动科动医大楼153室
  • 邮编：100193
  • 电话：010－62817823
  • 网址：http://www.chinajan.com
  • E-mail：cjan@chinajan.com
京公网安备 11010802041926号    京ICP备17050989号-2