动物营养学报 >

2025 , Vol. 37 >Issue 11: 7285 - 7297

DOI: https://doi.org/10.12418/CJAN2025.593

综述

牛磺酸的生物学功能及其在动物生产中的应用研究进展

  • 刘新宇 , 1, 2 ,
  • 栾嘉明 1, 2 ,
  • 冯鑫 1, 2 ,
  • 耿春银 , 1, 2, *
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  • 1 延边大学农学院,延吉 133000
  • 2 延边大学,东北寒区肉牛科技创新教育部工程研究中心,延吉 133002
*耿春银,教授,博士生导师,E-mail:

刘新宇(2000—),男,吉林长岭人,博士研究生,研究方向为反刍动物营养。E-mail:

Copy editor: 菅景颖

收稿日期: 2025-04-20

  网络出版日期: 2025-11-14

基金资助

国家自然科学基金项目(32060763)

国家自然科学基金项目(31660669)

吉林省科技发展项目(YDZJ202203CGZH042)

吉林省科技发展项目(20220202048NC)

吉林省教育厅科学研究项目(JJKH20220542KJ)

收起

Research Progress on Biological Functions of Taurine and Its Application in Animal Production

  • LIU Xinyu , 1, 2 ,
  • LUAN Jiaming 1, 2 ,
  • FENG Xin 1, 2 ,
  • GENG Chunyin , 1, 2, *
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  • 1 College of Agriculture, Yanbian University, Yanji 133000, China
  • 2 Engineering Research Center of North-East Cold Region Beef Cattle Science and Technology Innovation, Ministry of Education, Yanbian University, Yanji 133002, China
*professor, E-mail:

Received date: 2025-04-20

  Online published: 2025-11-14

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摘要

牛磺酸是一种在哺乳动物组织中含量很高的游离氨基酸,具有消炎、抗氧化等广泛的生物学功能。牛磺酸作为饲料添加剂已逐步被验证具有改善畜禽生产性能、肠道健康以及产品品质等作用。尽管如此,目前牛磺酸在动物生产中的应用潜力未得到充分开发,尤其在调控反刍动物瘤胃微生物以及与其他益生素协同使用的效果与影响机制还尚不明确。本文综述了牛磺酸的生物学功能、作用机制以及其在动物生产中的应用进展,并对影响牛磺酸应用效果的因素进行了总结,旨在为牛磺酸的高效利用提供参考。

关键词： 牛磺酸; 生物学功能; 作用机制; 动物生产

本文引用格式

刘新宇 , 栾嘉明 , 冯鑫 , 耿春银 . 牛磺酸的生物学功能及其在动物生产中的应用研究进展[J]. 动物营养学报, 2025 , 37(11) : 7285 -7297 . DOI: 10.12418/CJAN2025.593

Abstract

Taurine is a free amino acid with a high content in mammalian tissues and has a wide range of biological functions such as anti-inflammatory and antioxidant properties. As a feed additive, taurine has been progressively demonstrated to improve livestock and poultry production performance, enhance intestinal health, and upgrade product quality. However, its full potential in animal production remains underexplored, particularly regarding its modulation of rumen microbiota in ruminants and synergistic interactions with probiotics; the underlying mechanisms of these effects require further investigation. This paper reviewed the biological functions and mechanisms of action of taurine, summarized its advancements in animal production applications, and synthesized factors influencing its efficacy, aiming to provide references for the efficient utilization of taurine.

Key words： taurine; biological functions; mechanism of action; animal production

牛磺酸是一种不参与蛋白质合成的含硫氨基酸,最早由Tiedemann和Gmelin于1827年从牛胆汁中成功分离,广泛分布于动物组织和多种食物中。研究表明,在心脏、视网膜、中枢神经系统及骨骼肌等代谢活性高的组织中,牛磺酸呈现显著富集现象[1]。牛磺酸在肝脏中由蛋氨酸或半胱氨酸合成,其在维持细胞稳态和代谢平衡中发挥关键作用,新生儿和幼儿无法像成年人那样合成牛磺酸或部分组织在特定生理状态下需依赖外源补充以满足其生长发育和生理功能的需求,所以牛磺酸也被称为条件性必需氨基酸[2-3]。在畜牧生产领域中,牛磺酸作为一种功能性饲料添加剂已获得广泛关注,已有研究证实其具有抗氧化[4]、抗炎症[5]、调节脂质代谢稳态[6]以及改善肠道微生态平衡[7]等作用,可使动物生长性能提升、饲料效率改善、肠道屏障功能完善及产品品质优化,同时能有效缓解各类应激反应。基于此,本文综述了牛磺酸的生理学功能、作用机制以及影响其应用效果的因素,为其在动物生产中的应用提供参考。

1 牛磺酸的理化性质

牛磺酸化学名称为2-氨基乙磺酸,是一种含硫的有机化合物,分子式为C2H7NO3S,常温下为白色结晶或结晶性粉末,熔点为300 ℃,易溶于水,不溶于有机溶剂[8],化学性质稳定,其化学结构式如图1所示。
图1 牛磺酸的化学结构式

Fig.1 Chemical structural formula of taurine

2 牛磺酸的生物学功能及其作用机制

2.1 抗氧化能力

活性氧(reactive oxygen species,ROS)的过量产生可导致氧化还原稳态失衡,进而引发蛋白质、脂质及DNA生物大分子的氧化损伤,最终造成细胞功能障碍[9]。牛磺酸具有抗氧化作用,可有效缓解体内氧化应激过程。研究表明,在赭曲霉毒素A诱导的PK-15细胞模型中,牛磺酸可显著降低细胞内ROS含量,下调环氧合酶-2(cyclooxygenase-2,COX-2)和诱导型一氧化氮合酶(iNOS)的mRNA表达水平,同时上调多种抗氧化基因的表达,从而减轻细胞焦亡和细胞毒性[10]。核因子E2相关因子2(nuclear factor erythroid 2 related factor 2,Nrf2)是调控抗氧化应激相关通路的关键转录因子。在生理条件下,Nrf2由细胞质中的Kelch样ECH相关蛋白1(Kelch like ECH associated protein 1,Keap1)锚定[11],氧化应激时,两者解偶联,然后转运到细胞核中,与核内抗氧化应答元件(antioxidant responsive element,ARE)结合,通过诱导抗氧化基因的转录,从而保护细胞免受氧化应激损伤[12-13]。研究报道,牛磺酸可抑制2型糖尿病大鼠氧化应激,反映在改善较低的超氧化物歧化酶(superoxide dismutase,SOD)活性和较高的丙二醛(mmalondialdehyde,MDA)含量上,同时下调Keap1的mRNA表达水平以及上调Nrf2和血红素加氧酶-1(heme oxygenase-1,HO-1)的mRNA表达水平,验证了牛磺酸可通过对Keap1-Nrf2信号传导的调节实现缓解糖尿病大鼠氧化应激[4]。此外,牛磺酸还可通过激活Nrf2信号通路及其下游抗氧化酶来逆转黄曲霉毒素B1、过氧化氢(H2O2)诱导线粒体受氧化应激的影响,从而缓解细胞损伤[14-15]。在胰岛素抵抗模型中,牛磺酸可通过激活磷脂酰肌醇3激酶(phosphatidylinositol 3 kinase,PI3K)/蛋白激酶B(protein kinase B,Akt)信号通路,显著降低ROS荧光强度和MDA含量,同时提高SOD活性和谷胱甘肽(glutathione,GSH)含量,从而改善糖尿病大鼠的氧化应激状态[16]。此外,在采用牛磺酸干预H2O2诱导的成骨细胞氧化应激模型的试验研究中进一步发现,牛磺酸能够通过激活Wnt/β-连环蛋白(β-catenin)信号通路,调控细胞外信号调节激酶(extracellular signal-regulated kinase,ERK)的磷酸化水平,进而显著增强细胞的抗氧化防御能力[17]。
综上可知,牛磺酸可通过多途径、多靶点来发挥抗氧化作用,其抗氧化作用机制复杂,可调控Keap1-Nrf2、PI3K/Akt、Wnt/β-catenin等信号通路来调节抗氧化酶系统,改善机体氧化应激反应。

2.2 抗炎能力

炎症反应作为机体应对感染和组织损伤的重要防御机制,其核心功能在于修复受损组织并恢复内环境稳态。适度的急性炎症反应可有效清除病原体,而持续的慢性炎症则导致组织病理损伤。核因子-κB(nuclear factor kappa-B,NF-κB)是炎症反应的关键转录调控因子,在静息状态下通过与其抑制蛋白IκB结合而被锚定于细胞质中[18]。当受到Toll样受体(Toll-like receptor,TLR)激动剂等细胞因子刺激时,IκB发生快速磷酸化降解,促使NF-κB易位进入细胞核,激活下游炎症因子基因的转录,进而引起炎症反应[19]。研究发现,在葡聚糖硫酸钠诱导的结肠炎模型中,半胱氨酸亚磺酸脱羧酶(cysteine sulfinic acid decarboxylase,CSAD)和牛磺酸转运蛋白(taurine transporter,TauT)的表达水平显著下调,且基因敲除试验证实CSAD和TauT缺失会加剧炎症反应;对作用机制的研究表明,牛磺酸可通过与TLR4结合,抑制NF-κB信号通路,从而降低肿瘤坏死因子-α(tumor necrosis factor-α,TNF-α)和白细胞介素-6(interleukin-6,IL-6)等促炎因子的表达[20]。此外,牛磺酸和乳酸菌联用可降低急性胰腺炎小鼠模型髓过氧化物酶(myeloperoxidase,MPO)、白细胞介素-33(interleukin-33,IL-33)、NF-κB p65和辣椒素受体(transient receptor potential vanilloid-1,TRPV-1)抗衡炎症,并明确作用机制是通过TRPV-1/IL-33/NF-κB信号转导缓解急性胰腺炎[5]。在伯氏疟原虫ANKA株感染小鼠模型中,补充牛磺酸可显著降低血清促炎细胞因子的含量,并抑制脾脏炎性小体相关基因的表达,强调了牛磺酸在青蒿素治疗患疟疾小鼠过程中的抗炎益处[21]。此外,信号转导和转录激活因子3(signal transducer and activator of transcription,STAT3)是一种调节炎症反应相关基因的转录因子,广泛参与促炎和抗炎机制[22]。在结肠炎模型中报道,牛磺酸可通过STAT3介导的促炎信号传导对结肠炎发挥保护作用[23]。在脂多糖(lipopolysaccharide,LPS)诱导的脓毒症小鼠模型中发现,牛磺酸可降低中性粒细胞和单核细胞中TNF-α和白细胞介素-1β(interleukin-1β,IL-1β)的蛋白表达水平,并通过减少中性粒细胞浸润来改善肺脏、肝脏和肾脏的损伤[24]。
由此可见,牛磺酸通过抑制促炎信号传导,同时增强抗炎和可能的促溶解机制,对宿主炎症具有保护作用。

2.3 调节脂代谢能力

脂质代谢紊乱如高甘油三酯血症和高胆固醇血症是诱导心血管疾病和动脉粥样硬化的重要危险因素。牛磺酸通过多种分子机制参与脂质代谢的调控。沉寂信息调节因子1(sirtuin1,SIRT1)作为一种烟酰胺腺嘌呤二核苷(nicotinamide adenine dinucleotide,NAD+)依赖的蛋白脱酰酶,能够催化多种非组蛋白的去乙酰化修饰,在脂质代谢调控中发挥关键作用[25]。研究证实,牛磺酸干预可显著抑制高脂饮食诱导的小鼠体重增加和肝脏肥大,改善血清胆固醇和甘油三酯含量;其分子机制涉及上调肝脏组织中SIRT1活性及NAD+水平,促进SIRT1 mRNA和蛋白表达,同时抑制脂肪生成关键基因[固醇调节元件结合蛋白1c(sterol regulatory element-binding protein 1c,SREBP1c)、脂肪酸合酶(fatty acid synthase,FAS)、过氧化物酶体增殖物激活受体γ(peroxisome proliferator-activated receptor γ,PPARγ)]的表达,并激活β氧化相关基因[过氧化物酶体增殖物激活受体α(peroxisome proliferator-activated receptor α,PPARα)、肝脏X受体β(liver X receptor β,LXRβ)、过氧化物酶体增殖物激活受体γ共激活因子1α(peroxisome proliferator-activated receptor gamma coactivator 1α,PGC1α)、AMP活化蛋白激酶(AMP-activated protein kinase,AMPK)]和脂溶性调控基因[叉头框蛋白O1转录因子(forkhead box protein O1,FOXO1)],从而通过激活SIRT1/AMPK/FOXO1信号通路发挥对肥胖小鼠的降脂作用[6]。此外,小鼠长期补充牛磺酸可降低血浆中胆固醇和胆汁酸含量,上调肝脏中胆固醇7α-羟化酶(cholesterol 7-alpha hydroxylase,CYP7A1)表达水平,同时抑制ERK和成纤维细胞生长因子21(fibroblast growth factor 21,FGF21)表达水平,体外细胞试验也得出相同结果,验证了牛磺酸通过抑制FGF21/ERK通路上调Cyp7A1表达促进胆固醇代谢的机制[26]。另一研究通过细胞热位移试验、药物亲和响应靶标稳定性和双荧光素酶报告基因测定技术验证出,腺苷A2A受体(adenosine A2A receptor,A2AR)是牛磺酸的直接作用靶标,牛磺酸通过抑制A2AR-固醇调节元件结合蛋白2(sterol-regulatory element binding protein 2,SREBP-2)/环磷腺苷效应元件结合蛋白(cAMP-response element binding protein,CREB)/3-羟基-3-甲基戊二酰辅酶A还原酶(3-hydroxy-3-methylglutaryl-CoA reductase,HMGCR)轴改善胆固醇代谢失调[27]。
综上可知,牛磺酸可通过SIRT1/AMPK/FOXO1、FGF21/ERK以及A2AR-SREBP-2/CREB/HMGCR等信号通路多途径、多靶点调节脂代谢紊乱,保护机体健康。

2.4 调节肠道菌群能力

肠道微生物群落作为人和动物体重要的微生物器官,其组成和功能与宿主健康密切相关。新一代测序已将其与宿主表型相关联,一系列脑/肝脏/肺脏/皮肤-肠道-微生物群轴相互作用通路正在逐渐阐明[28-30]。研究表明,牛磺酸能够显著影响肠道微生物的组成和代谢功能。在健康小鼠模型中,牛磺酸干预可特异性富集放线菌门、变形菌门、双歧杆菌属和嗜双歧杆菌属等有益菌群[31],这些菌群可能参与了牛磺酸的肠道代谢过程。值得注意的是,牛磺酸通过调节胆汁酸代谢影响肠道菌群构成,其中胆盐水解酶(bile salt hydrolase,BSH)催化的初级胆汁酸向次级胆汁酸转化是关键环节。由于乳酸菌富含BSH活性,因此在牛磺酸干预的非酒精性脂肪肝病和抗生素处理模型中表现出显著增殖优势,并促进微生物群落稳态的恢复[31-32]。研究发现,无论是天然来源还是化学合成的牛磺酸,均能提高小鼠肠道中乳杆菌属和另支菌属等有益菌的相对丰度,同时抑制幽门螺杆菌等致病菌的增殖[7]。在模拟人结肠的试验中发现,牛磺酸对门和属水平的微生物群落组成没有影响[33],这一结果与以上小鼠试验的结果形成了鲜明的对比。这种差异可能源于人类与小鼠肠道微生物组成的固有差异或试验剂量差异(小鼠研究常用165 mg/kg,而人体推荐剂量是约40 mg/kg)。值得注意的是,临床研究发现,当人类肠道微生态改善时,牛磺酸代谢通路呈现显著上调特征[34],提示牛磺酸可能在人类肠道微生态调节中发挥潜在作用。欧阳轶强等[35]研究发现,牛磺酸对乳腺癌大鼠的肠道菌群结构有调节作用,这可能与其抗肿瘤作用存在关联。
综上可知,牛磺酸可调节肠道菌群稳态,通过提高益生菌相对丰度和降低有害菌相对丰度,在维持宿主肠道健康方面发挥积极作用(表1)。
表1 牛磺酸对肠道菌群的影响

Table 1 Effects of taurine on gut microbiota

模型
Models		 添加量
Additive quantity		 添加途径
Additive path		 影响
Effect		 参考文献
References
健康小鼠
Healthy mice		 200 mmol/L 饮水 放线菌门、变形菌门、双歧杆菌属、
嗜双歧杆菌属↑		 [31]
健康BALB/C小鼠
Healthy BALB/C mice		 165 mg/(kg·d) 灌胃 另枝菌属、紫单胞菌科、梭状
芽孢杆菌目、乳酸菌目↑,毛螺菌科、
理研菌科、螺旋杆菌科↓		 [7]
抗生素致肠道菌群失调小鼠
Mice with intestinal flora imbalance caused by antibiotics		 200 mmol/L 饮水 变形菌门、疣微菌门、乳杆菌属、
肠杆菌属和变形杆菌属↑		 [31]
鼠疫杆菌感染小鼠
Infection of mice by Yersinia pestis		 200 mmol/L 饮水 变形菌门、疣微菌门、变形杆菌属、
摩根氏菌属、乳杆菌属↑		 [31]
非酒精性脂肪肝病大鼠
Rats with non-alcoholic fatty
liver disease		 40 mg/(kg·d) 灌胃 双歧杆菌属、乳杆菌属↑,
大肠杆菌属↓		 [32]
乳腺癌大鼠
Rats with breast cancer		 6% 饮水 消化链球菌、Blautia producta、无乳链球菌、
粪肠球菌、脱硫弧菌属、Adlercreutzia↑,
Rosebura faecis、柔膜菌门、颤螺菌属↓		 [35]

↑:提高 increase;↓:降低 decrease。下表同 the same as below。

3 牛磺酸在动物生产中的应用

3.1 牛磺酸对生长性能的影响

牛磺酸能有效提高动物生长以及饲料利用率等生长性能指标。在断奶仔猪的饲养试验中,饲粮中添加0.1%、0.3%和0.5%牛磺酸可剂量依赖性地提高断奶体重,其中0.3%和0.5%的添加量显著改善了仔猪的饲料转化率(FCR)[36]。在母猪上的研究表明,妊娠后期至哺乳期补充1%牛磺酸能显著提高泌乳期平均日采食量(ADFI)和仔猪断奶体重[37]。在育肥猪上的试验表明,0.2%牛磺酸添加组终末体重和平均日增重(ADG)显著提升,同时FCR呈现改善趋势[38]。在家禽生产方面,适宜剂量的牛磺酸可显著增加肉鸡[39]、肉鸭[40]以及笼养蛋雏鸭[41]的终末体重、ADG以及FCR,并能提高蛋鸡[42]的产蛋率以及平均蛋重。在反刍动物的研究中,饲粮中添加4 g/(d·头)的牛磺酸可显著提高犊牛的终末体重[43];以包被形式每天补饲25、50和75 g牛磺酸均可显著提高育肥牛的ADG[44],直接添加20和40 g牛磺酸可改善瘤胃微生物蛋白合成并提高纤维消化率,但对生长性能无显著影响[45]。
综合以上研究结果可知,牛磺酸可提高畜禽的生长性能,表现为提高体重、ADG和FCR(表2)。
表2 牛磺酸对动物生长性能的影响

Table 2 Effects of taurine on growth performance of animals

动物
Animals		 添加量
Additive quantity		 影响
Effects		 参考文献
References
猪Pigs
断奶仔猪Weaned piglets 0.5% BW↑,FCR↑ [36]
断奶仔猪Weaned piglets 1.5% ADG↓,ADFI↓ [46]
断奶仔猪Weaned piglets 0.3% BW↑,FCR↑ [47]
妊娠与哺乳期母猪Pregnant and lactating sows 1.0% BW↑,ADFI↑ [37]
育肥猪Fattening pigs 0.2% BW↑,ADG↑ [38]
家禽Poultry
肉鸡Broilers 0.5% BW↑,FCR↑ [39]
肉鸡Broilers 0.6% BW↑,FCR↑ [48]
肉鸡Broilers 0.1% ADG↑,FCR↑ [49]
蛋鸡Laying hens 0.1% 产蛋率↑,平均蛋重↑,FCR↑ [42]
肉鸭Meat ducks 0.1% BW↑,FCR↑ [40]
蛋雏鸭Laying ducklings 0.1% ADG↑,FCR↑ [41]
鹌鹑Quails 0.05% ADFI↓,FCR↑ [50]
牛Cattle
犊牛Calves 4 g/(d·头) BW↑ [43]
肉用阉牛Beef steers 75 g/(d·头) ADG↑ [44]
肉用阉牛Beef steers 40 g /(d·头)(包被形式) 对生长性能无显著影响 [45]

BW:体重 body weight;FCR:饲料转化率;feed conversion ratio;ADG:平均日增重 average daily gain;ADFI:平均日采食量:average daily feed intake。

3.2 牛磺酸对肉品质的影响

肉品质是由pH、肉色[红度(a*)、黄度(b*)、亮度(L*)值]、滴水损失、肌内脂肪(IMF)含量以及肌肉纤维类型等组成的综合性指标,是养殖业中一项重要的经济性状,直接或间接影响消费者的消费倾向。其中,肌肉纤维类型尤为重要,包括氧化型(Ⅰ型和Ⅱa型)肌纤维和糖酵解型(Ⅱb型)肌纤维,Ⅰ型肌纤维具有较低的糖原储存量和较高的pH,与滴失率呈负相关,并富含肌红蛋白,决定动物屠宰后肉的颜色。研究表明,饲粮中添加0.6%牛磺酸可通过促进Ⅱb型肌纤维向Ⅰ型肌纤维转化,显著提高pH45 min和a*值,降低滴水损失、b*和L*值,并趋于提高背最长肌中单不饱和脂肪酸和油酸(C18∶1n9c)的含量,从而改善猪肉品质[51]。此外,通过体内与体外联合研究发现,牛磺酸可通过抑制低氧诱导因子1α(hypoxia-inducible factor 1α,HIF-1α)信号通路降低猪骨骼肌糖酵解活性[52]。在肉鸡的试验中发现,牛磺酸通过介导钙调磷酸酶(calcineurin,CaN)-活化T细胞核因子(nuclear factor of activated T-cells,NFAT)信号通路诱导Ⅱb型肌纤维向Ⅰ型肌纤维转化,改善腿肌pH45 min、pH24 h、蒸煮损失以及剪切力,同时显著降低乳酸含量以及糖酵解潜力[53]。已有多项研究报道了牛磺酸在改善肉鸡[54-56]与育肥猪[38,57-58]肉品质方面的积极作用。此外,适宜剂量的牛磺酸还能提高肉鸡[59]与肉鸭[40]胸肌率和瘦肉率,降低皮下脂肪厚度,从而优化屠宰性能。
综合以上研究发现,牛磺酸可通过介导CaN-NFAT和HIF-1α信号通路促进肌纤维类型的有益转化,降低糖酵解潜力,在提高肉品质方面表现出显著效果(表3)。
表3 牛磺酸对动物肉品质的影响

Table 3 Effects of taurine on meat quality of animals

动物
Animals		 添加量
Additive quantity		 影响
Effects		 参考文献
References



育肥猪
Fattening pigs		 0.3% pH45 min↑,a*值↑,L*、b*值↓
滴水损失↓,乳酸含量↓,单不饱和脂肪酸含量↑,油酸含量↑		 [53]
2 000 mg/kg pH45 min↑,a*值↑,肌红蛋白含量↑ [38]
800 mg/kg pH45 min↑,a*值↑,滴水损失↓ [57]




肉鸡
Broilers		 0.5% pH45 min↑,蒸煮损失↓,剪切力↓,乳酸含量↓,
肌纤维直径↓,肌纤维密度↑		 [54]
0.5% pH45 min↑,b*值↓,蒸煮损失↓,滴水损失↓ [55]
0.15% pH45 min↑,a*值↑,滴水损失↓,蛋白质含量↑,
胸肌率↑,瘦肉率↑,皮脂厚↓		 [56]
肉鸭
Meat ducks		 0.15% 屠宰率↑,全净膛率↑,腿肌率↑,瘦肉率↑,
腹脂率↓,系水力↑,剪切力↓		 [40]

a*:红度 redness;b*:黄度 yellowness;L*:亮度 lightness。

3.3 牛磺酸对肠道健康的影响

肠道作为动物机体重要的消化吸收和免疫器官,其健康状态直接影响畜禽的生产性能。肠道屏障功能是抵御病原体和有害抗原入侵的关键防线,而肠道微生物稳态的维持对于预防组织损伤和疾病发生具有重要作用。因此,良好的肠道结构、平衡的菌群以及稳定的屏障系统是肠道生理功能的基础。在LPS以及敌草快诱导的仔猪模型试验中,牛磺酸可改善仔猪肠道的绒毛形态结构[60-63],逆转肠道紧密连接蛋白封闭蛋白-1(Claudin-1)、闭锁小带蛋白-1(ZO-1)和闭合蛋白(Occludin)mRNA表达的下调,并降低血清中二胺氧化酶(decentralized autonomous organization,DAO)活性[60-62],从而有效维护肠道形态结构与屏障功能。同时,牛磺酸能通过调节微生物群落组成(如下调拟杆菌门、螺旋体科和普雷沃氏菌科的丰度,上调厚壁菌门和乳杆菌科的丰度)来缓解LPS诱导的结肠微生态失衡[60]。在母猪[37]与育肥猪[64]试验中,牛磺酸在哺乳母猪后代与育肥猪肠道菌群稳态和肠道屏障功能方面也表现出显著的调控作用。在肉鸡试验中,饲粮中添加牛磺酸可使空肠绒毛长度以及绒毛长度与隐窝深度比值有最佳表现[65-66],并显著上调空肠ZO-1的mRNA表达水平[67]。瘤胃球菌属[68]、NK4A214_group[69]和毛螺菌属[70]可促进反刍动物体内纤维素的降解。在反刍动物上的研究显示,牛磺酸能特异性富集瘤胃中具有纤维素降解功能的微生物,如瘤胃球菌属、NK4A214_group和毛螺菌属,从而提高中性洗涤纤维的消化率[45]。
综上所述,牛磺酸可改善肠道形态结构、维持肠道屏障功能以及调节肠道微生物稳态,进而保持畜禽肠道健康状态,以发挥良好的生产状态。

3.4 牛磺酸对应激的缓解作用

应激反应作为机体应对内外环境变化的非特异性生理反应,在超过适应阈值时可导致内环境紊乱并诱发多种疾病。尤其是在现代规模化、集约化畜禽养殖中,对生产性能的高要求普遍伴随多种应激,如断奶、运输、热、冷应激,严重制约畜禽生产性能的发挥。肉鸡由于羽毛丰富、缺乏汗腺和高代谢活动,特别容易受到热应激的影响[71],进而损害生长性能和肉品质[72]。有研究报道,牛磺酸可显著提高热应激肉鸡的体重[73-75]以及FCR[74,76],促进脂肪分解和蛋白质合成,并抑制胸肌蛋白质降解[76],进而减轻慢性热应激诱导的胸肌蛋白质损失[74,77-79]。在一项蛋鸡热应激的试验中,牛磺酸可提高平均产蛋率和蛋重,降低平均料蛋比,同时降低死淘率[80],还能减轻蛋鸡急性热应激诱发的输卵管损伤并增强机体免疫与抗氧化机能[81]。对热应激鹌鹑的研究表明,牛磺酸可上调回肠中营养转运蛋白的基因表达,如肽转运蛋白1(peptide transporter 1,PEPT1)、兴奋性氨基酸转运体3(excitatory amino acid transporter 3,EAAT3)、阳离子氨基酸转运蛋白2(cationic amino acid transporter 2,CAT2)、钠-葡萄糖协同转运蛋白1(sodium/glucose cotransporter 1,SGLT1)、钠-葡萄糖协同转运蛋白5(sodium/glucose cotransporter 5,SGLT5)、葡萄糖转运蛋白2(glucose transporter 2,GLUT2)和葡萄糖转运蛋白5(glucose transporter 5,GLUT5)[82]。此外,包被牛磺酸还可改善热应激种公羊精液质量,提高其繁殖性能[83-84]。慢性冷应激可诱发肉鸡氧化应激,导致心肌活性氧积累和损伤,而牛磺酸可通过上调Nrf2信号通路增强心肌抗氧化能力,对低温环境下心肌起到保护作用[85]。在断奶仔猪的研究中发现,牛磺酸可从肠道形态结构[86]、黏膜屏障和微生物菌群3个方面缓解肠道氧化损伤,进而导致腹泻程度下降[87]以及FCR升高[88-89],改善生长性能[47,90-91],缓解断奶应激的负面影响。运输应激是影响动物健康和生产性能的关键死前因素,牛磺酸可减少牦牛运输导致的体重损失,提高抗氧化能力和免疫机能,并上调瘤胃优势菌群的丰度[92]。
综上所述,牛磺酸作为一种高效的畜禽饲料添加剂,在缓解应激方面展现出显著的应用潜力,其通过增强抗氧化能力、改善免疫机能、调节营养代谢和维持肠道健康等多重机制,有效提升畜禽的生长性能、繁殖能力和产品品质。

4 影响牛磺酸应用效果的因素

4.1 牛磺酸的添加剂量

饲粮中添加超过1.5%的牛磺酸会导致仔猪生长和肠道健康受损[46]。蛋氨酸和半胱氨酸的过量摄入会导致动物生长受到抑制,而牛磺酸作为蛋氨酸和半胱氨酸的代谢产物,产生类似功效并不奇怪[93]。因此,在生产应用中牛磺酸的剂量选择尤为重要。不同牛磺酸添加剂量应用效果的研究主要集中在猪和禽生产中,以生长性能相关指标为例,表2总结了不同剂量牛磺酸应用效果的相关研究。几乎所有研究表明,低剂量添加牛磺酸对动物生长性能具有显著改善效果,而过量添加的效果与不同种类动物的耐受量需深入探究。

4.2 动物生理状况

研究表明,在动物肠道菌群稳态紊乱以及机体遭受氧化应激时牛磺酸的应用效果尤为显著,主要包含动物断奶期或是应激期(温度、运输)。例如,在牛磺酸添加剂量相似时,最适温度下肉鸡FCR提高7.6%和11.1%,在热应激温度条件下提高12.3%[76]。也有研究表明,牛磺酸可显著改善断奶期仔猪生长与腹泻状况[36-37]。这主要归因于牛磺酸可提高肠道有益菌(双歧杆菌属与乳杆菌属等)丰度以及通过多途径发挥抗氧化的生物学功能。牛磺酸在动物应激状态下的调控效应更为显著,这具有重要的生产应用价值。

4.3 与其他添加剂的协同作用

研究发现,牛磺酸和二甲双胍结合使用比单独使用更能抵抗糖尿病诱导的大鼠红细胞代谢紊乱和氧化应激[94]。饲粮中添加4 mg/kg包被叶酸和40 mg/kg牛磺酸可显著提高热应激羔羊的ADG,并改善抗氧化能力和免疫水平[95]。此外,牛磺酸和乳酸菌的联用在急性胰腺炎小鼠模型中表现出显著的抗氧化与抗炎作用[5]。以上研究表明,牛磺酸与其他添加剂联合使用的互作效应在动物生产中可能具有更佳效果,但在畜牧领域相关研究报道鲜有,需进一步探究。

4.4 其他因素

其他因素包括试验动物本身的差异和牛磺酸的添加形式等。研究表明,以包被方式补充牛磺酸可提高育肥牛的ADG[44],但以直接方式补充牛磺酸对育肥牛的生长无显著影响[45]。结果产生差异可能归因于直接方式补充牛磺酸会导致大部分牛磺酸在瘤胃中被水解,剩余部分不足以发挥促生长作用。此外,动物的年龄、品种以及生长阶段不同导致的肠道菌群和吸收能力的差异都会影响牛磺酸的应用效果。

5 小结与展望

牛磺酸作为一种具有多功能的生物活性物质,在医学和其他领域的研究中已展现出其多方位的作用,包括抗氧化、抗炎、调节脂代谢和肠道菌群等。在畜牧领域,牛磺酸已被证实能够显著改善动物的生长性能、产品品质、肠道健康,并可效缓解多种应激反应,对提高动物的生产性能和健康水平具有重要应用价值。
然而,尽管牛磺酸在畜牧领域的研究已取得显著进展,但仍存在以下问题:1)牛磺酸具有多重生物学功能,其在畜禽体内的作用机制尚未完全阐明;2)牛磺酸与其他饲料添加剂的互作效应尚需研究;3)牛磺酸在反刍动物中的应用研究较少,其在反刍动物生产中的应用潜力需进一步探索。
因此,未来研究应重点关注牛磺酸在反刍动物中的应用,重点解析其对瘤胃微生物生态系统及营养物质代谢的调控规律及对肉用反刍动物脂质和蛋白质合成与分解的调节作用;应探究牛磺酸与其他益生素和抗氧化剂的联合使用效果;此外,还应整合基因组学、代谢组学和蛋白质组学等系统生物学方法,结合生物信息学分析技术,精准鉴定其分子作用靶点,深入揭示其多途径调控网络,为牛磺酸在畜牧业中的精准应用提供理论依据。

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