动物营养学报 >

2025 , Vol. 37 >Issue 11: 7241 - 7250

DOI: https://doi.org/10.12418/CJAN2025.589

综述

半胱胺的生理功能及其在反刍动物生产中的应用

  • 吴铁成 , 1, 2 ,
  • 王涛 1 ,
  • 刘俊阳 1 ,
  • 刘斌 , 1, *
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  • 1 内蒙古自治区农牧业科学院,呼和浩特 010031
  • 2 肉羊遗传资源评价与繁育技术内蒙古自治区工程实验室,呼和浩特 010031
*刘斌,研究员,硕士生导师,E-mail:

吴铁成(1988—),男,甘肃天水人,副研究员,博士,主要从事营养学与饲料资源开发利用研究。E-mail:

Copy editor: 陈鑫

收稿日期: 2025-04-18

  网络出版日期: 2025-11-14

基金资助

内蒙古自治区自然科学基金项目(2024QN03007)

内蒙古农牧业青年创新基金项目(2021QNJJM02)

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Physiological Function of Cysteamine and Its Application in Ruminant Production

  • WU Tiecheng , 1, 2 ,
  • WANG Tao 1 ,
  • LIU Junyang 1 ,
  • LIU Bin , 1, *
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  • 1 Institute of Animal Husbandry, Inner Mongolia Autonomous Region Academy of Agricultural and Husbandry Science, Hohhot 010031, China
  • 2 Engineering Laboratory for Genetic Resources evaluation and Breeding Technology of Mutton Sheep in Inner Mongolia Autonomous Region, Hohhot 010031, China
*professor, E-mail:

Received date: 2025-04-18

  Online published: 2025-11-14

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摘要

半胱胺(CS)是一种在动物胃肠道和下丘脑产生的生物化合物,参与辅酶A和牛磺酸代谢。CS的巯基能与生长抑素特异性结合,间接地调节生长激素分泌,还可以作为细胞抗氧化剂。在反刍动物生产中,CS可以促进生长、缓解应激、增强抗氧化性能、减少瘤胃甲烷产生,是一种绿色安全高效的饲料添加剂。在绿色健康养殖的背景下,CS受到广泛关注。本文总结了CS的机体代谢途径、生理功能,阐述了CS对反刍动物主要生产性能的影响及应用前景。

关键词： 半胱胺; 反刍动物; 生产性能; 甲烷排放; 饲料添加剂

本文引用格式

吴铁成 , 王涛 , 刘俊阳 , 刘斌 . 半胱胺的生理功能及其在反刍动物生产中的应用[J]. 动物营养学报, 2025 , 37(11) : 7241 -7250 . DOI: 10.12418/CJAN2025.589

Abstract

Cysteamine (CS) is a biological compound produced in the gastrointestinal tract and hypothalamus of animals, which is involved in coenzyme A and taurine metabolism. The sulfhydryl group of CS can specifically bind to somatostatin, indirectly regulate the secretion of growth hormone, and can also act as a cellular antioxidant. In ruminant production, CS can promote animal growth, relieve stress, enhance antioxidant performance, and reduce rumen methane production. It is a green, safe and efficient ruminant feed additive. In the context of green and healthy breeding, the application of CS in ruminant production has received extensive attention. This paper summarized the metabolic pathways and physiological functions of CS, and combined with ruminant production, elaborated the influence of CS on the main production performance of ruminants and its application prospects.

Key words： cysteamine; ruminants; performance; methane emissions; feed additive

畜牧业的核心目标之一在于高效且可持续地为人类供应畜产品。反刍动物作为畜牧业中不可或缺的部分,以其独特的消化机制——反刍,在草食畜牧业中占据重要的地位。在全球气候变化和世界人口增长的背景下,反刍动物生产面临着诸多挑战,包括饲料资源的短缺、生产效率的提升以及温室气体的排放等问题[1]。半胱胺(cysteamine,CS)作为动物体内天然存在的生物活性物质,能够通过耗竭生长抑素(somatostatin,SS)来间接提高生长激素(growth hormone,GH)和胰岛素样生长因子-1(insulin-like growth factor-1, IGF-1)的含量,从而促进动物生长[2-3]。CS还可以提高家畜饲料转化率[4],改善胴体品质[5],参与体内重要的营养物质的代谢[6]以及降低瘤胃发酵过程中甲烷(CH4)的产生[7]。因此,CS作为一种饲料添加剂,在提高反刍动物生产效率、实现畜牧业可持续发展等方面有巨大的潜力。本文就CS在反刍动物中的应用及其作用方式的研究进展进行综述。

1 CS的理化性质及机体代谢途径

CS是一种含有巯基和氨基的化合物,其线性分子式为HS-CH2-CH2-NH2,相对分子质量为77.140 7,有特殊气味,不易溶于水,化学性能不稳定。因此,在饲料中直接添加CS具有一定的局限性,为了提高适口性、药代动力学特性以及实用性,在生产中通常制成包被半胱胺盐酸盐使用。CS是辅酶A(coenzyme A, CoA)和牛磺酸代谢途径的重要环节之一,CoA是由半胱氨酸和泛酸合成的,而内源性CS产生于CoA的降解过程中。牛磺酸内源性生物合成途径中,半胱氨酸通过半胱氨酸双加氧酶和半胱氨酸亚磺酸脱羧酶代谢,生成亚牛磺酸[8-9],此外,CS通过半胱胺双加氧酶也可生成亚牛磺酸,亚牛磺酸进一步被氧化成牛磺酸(图1)。因此,CS可替代部分半胱氨酸合成牛磺酸。美国食品和药物管理局(FDA)和欧洲药品管理局(EMA)批准CS可应用于人体和生产畜产品的哺乳动物。
图1 半胱胺在体内的代谢

Fig.1 Cysteamine metabolism in vivo

2 CS的生理功能

CS是一种含巯基和氨基的氨基硫醇,它在生物体内具有重要的生理调节功能,能够改善动物的生长性能、免疫功能、抗氧化能力和繁殖能力,并通过调节肠道微生物群落优化整体健康状态。

2.1 调节动物机体GH-IGF-1轴相关激素

动物生长发育是受多因素调控的复杂生理过程,其中神经内分泌系统通过生长轴发挥核心调控作用[10]。生长轴由下丘脑-垂体-靶器官构成的功能级联系统,而GH-IGF-1轴作为该系统的核心代谢调节网络,通过协调GH、IGF-1及其受体系统,在细胞增殖、组织分化以及机体代谢调控中发挥关键作用[11]。该轴的调控始于下丘脑通过双相分泌生长激素释放激素(growth hormone-releasing hormone, GHRH)和SS,前者促进垂体GH的合成与分泌,后者则通过负反馈机制抑制GH和GHRH的分泌[12]。SS作为该调控网络的关键负向调节因子,其分子结构与功能特征具有重要研究价值。Brazeau等[13]首次从绵羊下丘脑分离出14肽SS,其分子内Cys3-Cys14二硫键形成的环状结构对生物活性至关重要。CS能与SS分子内二硫键形成混合二硫键复合物,导致SS空间构象改变并显著降低其免疫活性。饲粮中每日添加7.5和15 mg/kg BW CS均显著提高了东北细毛羊肝脏和肌肉中IGF-1 mRNA的相对表达量[14]。而Shokrollahi等[3]在羔羊饲粮中添加CS后,血清GH含量较对照组极显著提高。类似效应在猪[15]、绵羊[16]和水牛[17]等物种中均得到验证。这些研究证实CS通过耗竭动物体内SS,来实现GH-IGF-1轴的调控,从而有效改善动物生产效率。

2.2 增强动物机体免疫性能

CS作为含硫生物活性分子,通过调节氧化还原稳态和免疫信号通路,在抗炎、抗菌及免疫稳态维持中发挥重要作用[18-20]。沈赞明等[21]试验表明,泌乳荷斯坦奶牛连续4周摄入包被半胱胺盐酸盐后(第1周2 000 U/d,后3周3 000 U/d),外周淋巴细胞转化率、白细胞介素-2和白细胞介素-6含量较对照组分别提高32%、28%和23%。在肠道免疫调控方面,饲粮补充CS后,猪空肠黏膜免疫球蛋白A(immunoglobulin A,IgA)、免疫球蛋白M(immunoglobulin M,IgM)和免疫球蛋白G(immunoglobulin G,IgG)含量较对照组极显著增加[22],猪肠道免疫因子的mRNA相对表达量显著增加,减轻敌草快引起的炎症反应[23]。在剂量效应方面,在体内低剂量CS(12.5 mg/kg BW)可增强伴刀豆球蛋白A和脂多糖诱导的小鼠脾淋巴细胞增殖,而高剂量(300和400 mg/kg BW)则呈现抑制作用[24]。因此,CS可能通过促进淋巴细胞有丝分裂及上调免疫活性分子基因表达来实现对病原微生物的防御。

2.3 缓解动物机体应激

CS作为多巴胺-β-羟化酶(dopamine β-hydroxylase,DβH)的竞争性抑制剂,可阻断多巴胺(dopamine,DA)向去甲肾上腺素(norepinephrine,NE)的转化。Terry等[25]研究发现,大鼠静脉注射300 mg/kg BW CS 4 h后,极显著降低NE和肾上腺素含量,同时DA蓄积量极显著增加,揭示了CS在神经内分泌应激反应中的关键作用。CS在奶牛[26]和红罗非鱼[27]的应用中也表现出抗应激作用。因此,CS可能通过调节神经内分泌,来缓解动物机体的生理应激状态。

2.4 提高动物机体抗氧化性能

氧化应激的核心机制在于活性氧(reactive oxygen species, ROS)的过量积累,这类含未配对电子的高活性分子可通过链式反应引发DNA氧化断裂、脂质过氧化及蛋白质羰基化等生物大分子损伤[28]。Zhou等[22]研究表明,在育肥猪饲粮中补充142 mg/kg CS,能显著提高空肠黏膜中谷胱甘肽(glutathione,GSH)含量和谷胱甘肽过氧化物酶(glutathione peroxidase,GSH-Px)活性,降低丙二醛(malondialdehyde,MDA)含量,可减少的氧化应激。CS治疗还降低了蛛网膜下腔出血后48 h大鼠脑皮层中的ROS、MDA含量和增加了GSH-Px活性[29]。研究发现,当CS的添加浓度过高时,在过渡金属的参与下,CS会被氧化生成H2O2,还会通过抑制GSH-Px的活性进一步加重氧化应激[30]。
CS通过多层次抗氧化防御系统的协同作用,有效维持细胞氧化还原稳态,作用机制可概括为以下3方面:其一,CS分子中的β-巯基具有强还原电势,可通过电子转移机制直接清除自由基,如羟基自由基(-OH)、过氧化氢(H2O2)等;其二,CS通过裂解胱氨酸二硫键生成半胱氨酸,从而提升GSH合成速率;其三,CS的代谢产物牛磺酸能够调节核因子E2相关因子2/抗氧化反应元件(Nrf2/ARE)信号通路,激活调控细胞内ARE的表达,启动抗氧化酶的表达,从而减轻氧化应激损伤[31-32]。

3 CS对反刍动物瘤胃代谢的影响

瘤胃是一个复杂的微生物生态系统,栖息着大量的细菌、真菌、原虫等微生物群落。CS作为一种饲料添加剂,能够通过改变反刍动物瘤胃微生物的组成和功能,提高饲料转化效率,减少温室气体排放,并优化瘤胃发酵过程,从而对反刍动物的生长和健康产生积极影响。

3.1 瘤胃发酵参数

瘤胃发酵参数(如pH、氨态氮、挥发性脂肪酸等)的动态变化不仅直接影响瘤胃微生物的健康状态和代谢活性,还对宿主的整体健康和生产性能具有重要意义[33-34]。研究表明,在藏绵羊断奶羔羊的精料中连续42 d每2 d添加300 mg/kg BW的半胱胺盐酸盐,能显著提高瘤胃液总挥发性脂肪酸(TVFA)含量[35]。进一步研究发现,在绵羔羊饲粮中连续64 d添加60 mg/kg BW包被CS(含量50%),可使瘤胃液中氨态氮、微生物蛋白(MCP)、丙酸和TVFA含量分别提高15.79%、7.43%、12.23%和15.64%[36]。体外发酵试验表明,CS也能调控瘤胃挥发性脂肪酸组成、促进MCP合成。因此,CS为调控瘤胃发酵提供了有效策略[37-38]。

3.2 瘤胃微生物多样性

瘤胃微生物的多样性与宿主健康、代谢能力密切相关[39-40]。研究表明,绒山羊饲粮补充120 mg/kg BW的半胱胺盐酸盐,显著提高了瘤胃中魏斯氏菌属(Weissella)和乳球菌属(Lactococcus)的相对丰度,但是对细菌α多样性影响不显著[2],这些细菌在体外[41-42]和体内[43]均表现出良好的抗氧化活性,表明其可能对宿主抗氧化性能具有潜在影响。Hu等[44]进一步发现,牦牛饲粮中每日补充5 g半胱胺盐酸盐,显著增加了瘤胃中普雷沃氏菌属1(Prevotella_1,纤维素分解菌)、丁酸弧菌属2(Butyribrio_2,丁酸合成菌)以及纤维杆菌属(Fibrobacter,纤维素分解菌)的丰度,显著降低了α多样性。类似的结果也在水牛的研究中得到验证[17]。这表明CS作为一种饲料添加剂,对瘤胃微生物的影响也不同,这可能和物种与添加剂量有关,适宜的剂量不仅能够调节瘤胃微生物群落结构,还能通过促进纤维素分解菌和丁酸合成菌的生长,增强了反刍动物对营养物质的可利用性,进而提高了营养物质的吸收效率。

3.3 CH4排放

反刍动物产生的CH4占全球温室气体排放量的16%,并导致饲料能量损失2%~12%[45]。瘤胃中CH4的主要生成途径是产甲烷古菌通过CO2和H2的还原反应生成CH4,这一过程占瘤胃CH4生成的82%,是CH4主要的生成途径[46]。因此,抑制瘤胃CH4生成对反刍动物产业的可持续发展具有重要意义。Barnett等[4]在绵羊试验中发现,添加80 mg/kg BW的CS可使CH4排放量从19.3降至17.1 g CH4/kg DMI。在牛的试验中连续56 d每日饲粮中添加80 mg/kg BW半胱胺盐酸盐,虽未显著降低CH4产量,但显著降低了CH4生成速率[47]。此外,水牛瘤胃体外发酵模型显示,当CS添加量为0.01%时,对CH4生成无显著影响,但当添加量增加至0.8%时,CH4产量显著减少[37]。这些差异可能与CS添加量、试验时间或不同物种的瘤胃微生物群落特性有关。
CS的CH4减排机制可概括为以下2方面:其一,CS通过减少瘤胃中原生动物种群的数量,直接抑制CH4生成[47];其二,CS通过促进丙酸生成,增加H2的消耗,从而间接抑制CH4生成[36]。这一双重效应不仅减少了CH4排放,还提高了饲料能量的利用效率,为反刍动物的低碳生产提供了潜在策略。

4 CS对反刍动物生产性能的影响

CS通过改善反刍动物瘤胃代谢、调节生长轴相关激素水平和增强抗氧化能力,显著提高了反刍动物的生产性能和饲料利用效率,例如降低了料重比[36]及增加日增重[4]、产奶量[48]以及山羊绒品质[49]。在大多数反刍动物研究中,CS在饲粮中的添加剂量在10~100 mg/kg BW时能获得较大的生产性能提升,见表1。CS提高反刍动物生产性能的作用方式见图2。
表1 CS对反刍动物生产性能的影响

Table 1 Effects of cysteamine on performance of ruminant

动物
Animals		 试验期
Trial
period/d		 类型及纯度
Type and
purity		 剂量
Dose/
(mg/kg
BW)		 添加频率
Add
frequency/
(d/次)		 显著影响的指标
Significant
indicators		 参考文献
References



绵羊
Sheep


35


半胱胺盐酸
盐/99%


80		 1 日增重提高28.26%,
料重比降低24.43%


[4]
3 日增重提高22.83%,
料重比降低14.50%
肉牛
Beef cattle		 63 包被半胱胺
盐酸盐/30%		 57 1 日增重提高15.68%,
料重比降低14.2%		 [50]
东北细毛羊
Northeast
fine-wool sheep		 85 半胱胺 125 1 日增重提高20.96%,
饲料转化率降低15.82%		 [51]
山羊
Goat		 60 包被半胱胺
盐酸盐/30%		 100 1 日增重提高41.18%,
料重比降低8.45%		 [52]
藏羊
Tibetan sheep		 40 半胱胺 13.5 1 日增重提高45.54%,
料重比降低31.27%		 [53]
牦牛
Yak		 28 包被半胱胺
盐酸盐/30%		 90 1 日增重提高54.17% [54]
黄牛
Yellow cattle		 42 包被半胱胺
盐酸盐/30%		 80 1 日增重提高16.07%,
料重比降低15.24%		 [46]
绵羊
Sheep		 64 包被半胱
胺/50%		 60 1 日增重提高19.92%,
料重比降低13.51%		 [47]
奶牛
Dairy cattle		 30 包被半胱胺
盐酸盐/30%		 5 1 乳脂率和乳蛋白率
分别提高9.60%、5.02%		 [26]
奶牛
Dairy cattle		 56 包被半胱胺
盐酸盐/30%		 75 g/头 1 产奶量和乳脂率分
别提高6.21%、9.70%		 [55]
绒山羊
Cashmere goat		 90 包被半胱胺
盐酸盐/30%		 120 1 产绒量、伸直长度分别
提高12.67%、20.67%		 [49]
东北细毛羊
Fine-wool sheep		 85 半胱胺 125 1 毛长比提高14.38% [51]
绒山羊
Fine-wool sheep		 60 半胱胺 100 7 绒伸直长度、毛伸直长度分别
提高46.56%、25.32%		 [56]
藏绵羊
Tibetan sheep		 35 包被半胱胺
亚酸盐		 10 1 精子活力提高4.55% [57]
哈萨克羔羊
卵母细胞
Kazakh lamb
oocytes		 - 半胱胺 100 μmol/L - 提高羔羊卵母细胞卵裂率、
囊胚率、成熟率、正常受精率、
卵裂率及胚泡率		 [58]
羊精液
Sheep semen		 - Se-NPs半胱胺 1 μg/mL - 显著增加精子的运动性、
活力、膜完整性		 [59]
水牛精液
Buffalo semen		 - 半胱胺 100 μmol/L - 显著提高解冻后的运动参数
指标和质膜完整		 [60]
图2 CS提高反刍动物生产性能的作用方式

Hypothalamus:下丘脑;Pituitary:垂体;Liver:肝脏;Rumen:瘤胃;Anti-oxidant properties:抗氧化特性;Immune properties:免疫特性;Anti-stress properties:抗应激特性;Body weight gain:增重;Feed efficiency:饲料转化效率;Milk production:产奶量;Wool and cashmere growth:绒毛生长性能;Reproductive performance:繁殖性能;CS:半胱胺 cysteamine;SS:生长抑素 somatostatin;GHRH:生长激素释放激素 growth hormone-releasing hormone;GH:生长激素 growth hormone;IGF-1:胰岛素样生长因子-1 insulin-like growth factor-1;NH3-N:氨态氮 ammonia nitrogen;TVFA:总挥发性脂肪酸 total volatile fatty acids;CH4:甲烷 methane;H2S:硫化氢 hydrogen sulfide;H+:氢离子 hydrogen ion;HS-:硫氢根离子 hydrosulfide ion。

Fig.2 Mode of action of cysteamine in improving performance of ruminants

5 小结与展望

CS作为一种饲料添加剂,在提高反刍动物生产效率、提升免疫力、增强抗氧化能力、降低瘤胃CH4排放、改善畜产品品质等方面有积极的作用,其优势主要有:1)CS能耗竭不同物种的SS;2)CS参与机体代谢,无残留风险;3)CS生产工艺相对简单,成本较低;4)CS可直接添加饲粮,便于实际生产应用。然而,CS也有潜在的风险和限制,高剂量CS可能引发机体氧化反应,导致炎症(如消化道溃疡),进而降低采食量和消化代谢率,限制其应用效果。因此,应考虑CS在不同生产系统中剂量效应及安全性评估。尽管CS的上述功能已被部分揭示,但跨代效应机制不明:现有研究多聚焦单一生理阶段,母源CS如何通过胎盘或乳汁介导代谢物跨代传递,进而影响子代生产性能的证据不足。在研究方法上,现有方法主要以单一候选指标测定为主,不能全面解释其作用机制。因此,后续的研究需要集中在CS的跨代效应机制以及整合多种“组学”技术获得的信息,来发现CS的作用机理、明确CS的需要量等,从而提高反刍动物生产性能,改善畜产品品质。

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