Chinese Journal of Animal Nutrition >

2026 , Vol. 38 >Issue 2: 1212 - 1224

DOI: https://doi.org/10.12418/CJAN2026.097

RESEARCH PAPER

Effects of Rumen Fluid Transplantation and Feeding Compound Probiotics on Growth Performance, Serum Biochemical Indices and Fecal Microbiota Structure of Yaks

  • WANG Zhilong , 1, 2, 3, 4 ,
  • XU Shengchun 1, 2, 3, 4 ,
  • LIU Xinbing 1, 2, 3, 4 ,
  • ZHANG Jiyuan 1, 2, 3, 4 ,
  • WANG Jiaming 1, 2, 3, 4 ,
  • YE Long 1, 2, 3, 4 ,
  • YANG Yingkui 1, 2, 3, 4 ,
  • WANG Honglu 5 ,
  • SHI Haisheng 5 ,
  • CHAI Shatuo 1, 2, 3, 4 ,
  • LYU Jiaying 1, 2, 3, 4 ,
  • WANG Xun , 1, 2, 3, 4, * ,
  • WANG Shuxiang , 1, 2, 3, 4, *
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  • 1 Qinghai University, Xining 810016, China
  • 2 Qinghai Provincial Key Laboratory of Animal Nutrition and Feed Science in Plateau Herding, Xining 810016, China
  • 3 Qinghai Provincial Yak Engineering and Technology Research Center, Xining 810016, China
  • 4 National Yak Technology Innovation Center (Preparation), Xining 810016, China
  • 5 Huangyuan County Animal Husbandry and Veterinary Station, Xining 812100, China
* WANG Xun, associate professor, E-mail: ;
WANG Shuxiang, associate professor, E-mail:

Received date: 2025-07-08

  Online published: 2026-02-12

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Abstract

This experiment aimed to study the effects of rumen fluid transplantation and feeding compound probiotics on growth performance, serum biochemical indices and fecal microbiota structure of yaks. Thirty 8-month-old weaned yaks with similar body weight and good body condition were randomly divided into 3 groups with 10 yaks in each group. The control group (CK group) was fed a basal diet; the rumen transplantation group (RT group) was fed a basal diet, and conducted rumen fluid transplantation on the days 5, 3 and 1 before the start of the experiment; the compound probiotics group (CP group) was fed the basal diet supplemented with 0.2% compound probiotics. The experiment lasted for 60 days. The results showed as follows: 1) the final body weight, total weight gain and average daily gain of CP group were significantly higher than those of CK group (P<0.05), and the dry matter intake and feed to gain ratio of RT group and CP group were significantly lower than those of CK group (P<0.05). 2) The serum interleukin-1β (IL-1β) content of CP group was significantly lower than that of CK group (P<0.05), the serum catalase (CAT) activity of RT group and CP group was significantly higher than that of CK group (P<0.05), and the serum glutathione peroxidase (GSH-Px) activity of RT group was significantly higher than that of CK group (P<0.05). 3) The Chao1 index, Observed_features index, Shannon index and Simpson index of CP group were significantly higher than those of RT group and CK group (P<0.05). 4) At the phylum level, the fecal Firmicutes relative abundance of CP group and CK group was significantly higher than that of RT group (P<0.05). At the genus level, the fecal UCG-005 relative abundance of CK group was significantly higher than that of RT group (P<0.05), the fecal Monoglobus relative abundance of CP group was significantly higher than that of RT group (P<0.05), and the fecal Clostridium_sensu_stricto_1 relative abundance of RT group was significantly higher than that of CP group (P<0.05). 5) The KEGG functional prediction analysis revealed that compared with CK group, the translation, nucleotide metabolism and signaling molecules and interaction et al metabolic pathways were significantly enriched in RT group (P<0.05), and the nucleotide metabolism, amino acid metabolism and signaling molecules and interaction et al metabolic pathways were significantly enriched in CP group (P<0.05). In conclusion, the rumen fluid transplantation is more suitable for quickly improving the antioxidant capacity and feed utilization efficiency of yaks in the short term; while feeding compound probiotics is more beneficial for long-term maintenance of intestinal microbiota stability, enhancing anti-inflammatory response regulation ability, and promoting weight gain and improving feed utilization efficiency of yaks.

Key words： yaks; rumen fluid transplantation; compound probiotics; growth performance; inflammation; antioxidant; fecal microbiota

Cite this article

WANG Zhilong , XU Shengchun , LIU Xinbing , ZHANG Jiyuan , WANG Jiaming , YE Long , YANG Yingkui , WANG Honglu , SHI Haisheng , CHAI Shatuo , LYU Jiaying , WANG Xun , WANG Shuxiang . Effects of Rumen Fluid Transplantation and Feeding Compound Probiotics on Growth Performance, Serum Biochemical Indices and Fecal Microbiota Structure of Yaks[J]. Chinese Journal of Animal Nutrition, 2026 , 38(2) : 1212 -1224 . DOI: 10.12418/CJAN2026.097

牦牛(Bos grunniens)具备在高寒、低氧及粗饲料匮乏生态环境中生存和利用资源的独特适应能力[1]。在断奶期,犊牛面临从母乳到粗饲料的过渡,这一阶段瘤胃发育尚不完全,微生物群落的稳定性较差,容易引发肠道菌群失调、黏膜屏障损伤及系统性炎症反应,进而导致生长迟缓和死亡率升高[2]。因此,针对这一阶段的微生态调节显得尤为重要,瘤胃液移植和添加复合益生菌作为新兴的微生态干预措施,已被提出用于改善反刍动物的肠道健康[3]。近年来,瘤胃液移植技术得到了研究人员的广泛关注,其在重建瘤胃微生物区系方面取得了一定的效果[4]。Ribeiro等[5]研究表明,接种瘤胃提取物成功改变了牛瘤胃微生物组,增加了蛋白质的消化率和氮的保留率,并改变了挥发性脂肪酸(volatile fatty acids,VFA)组成。Zhong等[6]研究表明,对羔羊接种新鲜瘤胃液有利于改善过渡期羔羊的生长性能。Leo-Penu等[7]研究表明,对长期运输的肉牛进行瘤胃液移植,显著提高了干物质采食量,通过改善采食行为促进了营养摄入,为肌肉恢复提供了物质基础,间接缓解了运输应激对肉牛的负面影响。
益生菌分为单一益生菌和复合益生菌,与单一益生菌相比,复合益生菌更有优势,不同菌株相结合时可以发挥协同作用[8]。饲粮中添加枯草芽孢杆菌和地衣芽孢杆菌可以降低育猪的腹泻率,提高免疫力[9]。地衣芽孢杆菌具有调节肠道微生物群落、促进生长、抗炎和免疫刺激等作用[10]。此外,地衣芽孢杆菌还通过抗氧化激活宿主合成谷胱甘肽过氧化物酶(GSH-Px)和超氧化物歧化酶(SOD),增强机体抗氧化能力[11]。唐文雅等[12]研究发现,嗜酸乳杆菌通过代谢产生乳酸,降低肠道pH,抑制肠道致病菌大肠杆菌的增殖,降低腹泻率。虽然现有研究分别探讨了瘤胃液移植或添加益生菌对反刍动物的影响,但尚未有研究系统比较这2种微生态干预措施在断奶牦牛中的效果差异及作用机制。因此,本试验旨在研究瘤胃液移植与饲喂复合益生菌对牦牛生长性能、血清生化指标及粪便菌群结构的影响,进一步探讨这2种微生态干预措施对牦牛健康的作用机制,以及哪种微生态干预措施更适合改善断奶期牦牛的健康状况,为微生态调控技术在牦牛养殖中的应用提供理论依据。

1 材料与方法

1.1 伦理声明

本试验由青海大学动物伦理委员会批准,试验伦理批准编号为QHU20240917。

1.2 试验材料

复合益生菌购自某生物工程有限公司,其中枯草芽孢杆菌D165、嗜酸乳杆菌HC1356和地衣芽孢杆菌D132比例为1∶1∶1,有效活菌数≥1×1010 CFU/g。
选取5~10头3岁左右的成年健康公牦牛作为胃液供供体牦牛,瘤胃液供体牦牛和受体牦牛均来自同一牧场,并饲喂相同饲粮,以确保瘤胃微生物群落的一致性和移植效果的稳定性。在试验开始前第5、3、1天,采用口腔采集法在晨饲前用瘤胃液采集管收集新鲜瘤胃液。为最大限度保持瘤胃微生物的活性,采集的瘤胃液立即用4层纱布过滤去除大颗粒饲料残渣,置于保温容瓶中备用,并在采集后30 min内完成灌服操作[13-15]。

1.3 试验设计与饲养管理

从海拔约3 200 m的牧场选取30头8月龄的健康牦牛,随机分为3组,每组10头牛。对照组(CK组)饲喂基础饲粮,不进行任何微生态干预;瘤胃液移植组(RT组)饲喂基础饲粮,并在试验开始前第5、3、1天进行瘤胃液移植,瘤胃液移植方案参考Li等[16]的研究结果,每头牦牛灌服约340 mL左右,一共移植3次,确保每头牦牛移植1 L的新鲜瘤胃液。复合益生菌组(CP)在基础饲粮中添加0.2%的复合益生菌[17-18]。
本试验于2025年1月在青海省西宁市湟源县青海众汇农牧科技有限公司进行。基础饲粮组成及营养水平见表1,以全混合日粮(total mixed ration,TMR)形式饲喂,3组牦牛在牛栏中进行群饲,每天08:00进行饲喂,自由饮水。试验期60 d。
表1 基础饲粮组成及营养水平(风干基础)

Table 1 Composition and nutrient levels of the basal diet (air-dry basis) %

原料
Ingredients		 含量
Content		 营养水平
Nutrient levels2)		 含量
Content
玉米Corn 45.20 代谢能ME/(MJ/kg) 12.00
豆粕Soybean meal 20.00 干物质DM 86.00
棉籽粕Cottonseed meal 8.00 粗蛋白质CP 19.50
麦麸Wheat bran 10.00 粗脂肪EE 4.00
苜蓿粉Alfalfa powder 8.00 中性洗涤纤维NDF 30.00
碳酸钙CaCO3 1.20 酸性洗涤纤维ADF 17.00
磷酸氢钙CaHPO4 0.50 钙Ca 1.00
食盐NaCl 0.30 磷P 0.45
预混料Premix1) 4.20
大豆油Soybean oil 2.60
合计Total 100.00

1)预混料为每千克饲粮提供 The premix provided the following per kg of the diet: Cu 15 mg,Fe 70 mg,Zn 25 mg,Mn 35 mg,Se 0.3 mg,I 0.5 mg,Co 0.2 mg,VA 5 000 IU,VD 600 IU,VE 50 IU。

2)代谢能为计算值,其他为实测值。ME was a calculated value, while the others were measured values.

1.4 样品采集

试验期最后1天采集血液和粪便。通过颈静脉采血法,采集所有犊牛血液样品,静止放置2 h后在4 ℃、1 095×g条件下离心10 min,取上清液分装于2 mL的冻存管中,移至液氮中保存待测。每组随机选取5头牛,用无菌长臂手套取新鲜直肠粪便,移至液氮中保存待测。

1.5 指标测定

1.5.1 饲粮营养成分

粗蛋白质(CP)含量参考GB/T 6432—2018的方法进行测定,粗脂肪(EE)含量参考GB/T 6433—2006的方法进行测定,中性洗涤纤维(NDF)含量参考GB/T 20806—2022的方法进行测定,酸性洗涤纤维含量参考NY/T 1459—2022的方法进行测定,钙含量参照GB/T 6436—2018的方法进行测定,磷含量参照GB/T 6437—2018的方法进行测定。代谢能参考《中国饲料成分及营养价值表(2024年第35版)》[19]进行计算。

1.5.2 生长性能测定

试验期间每天记录各组牦牛的投喂饲料量和剩料量,计算干物质采食量(DMI),在试验开始和结束时,对所有牦牛进行空腹称重,计算总增重(TWG)、平均日增重(ADG)和料重比(F/G)。

1.5.3 血清生化指标测定

采用酶联免疫吸附测定法,使用酶标分析仪(Epoch,BioTek,美国)测定血清炎症细胞因子含量,炎症细胞因子包括:肿瘤坏死因子-α(TNF-α,货号H052-1-1)、白细胞介素-1β(IL-1β,货号H002-1-1)和白细胞介素-6(IL-6,货号H007-1-1);采用试剂盒测定血清抗氧化指标,抗氧化指标包括:丙二醛(MDA,货号G4302)、过氧化氢酶(CAT,货号G4307)、谷胱甘肽过氧化物酶(GSH-Px,货号G4310)、超氧化物歧化酶(SOD,货号G4306)和总抗氧化能力(T-AOC,货号G4313),试剂盒均购自武汉塞维尔生物有限公司,参照试剂盒说明书进行测定。

1.5.4 粪便微生物16S rDNA扩增子测序

粪便微生物16S rDNA测序由北京诺禾致源科技股份有限公司进行检测。使用DNA提取试剂盒(D5625-02,BioTek,美国)从瘤胃液样本中提取肠道微生物的总基因组DNA。使用2%琼脂凝胶电泳评估DNA的完整性。使用通用正向引物(5'-CCTAYGGGRBGCASCAG-3')和反向引物(5'-GGACTACNNGGGGTATCTAAT-3')扩增细菌16S rDNA基因的V3~V4区域。采用线性判别分析效应大小(LEfSe)进一步分析筛选出组间差异的菌属,线性判别分析(LDA)评分阈值设置为4.0。对牦牛粪便菌群进行KEGG功能预测,数据采用t-test进行检验分析。

1.6 数据统计分析

使用SPSS 27.0软件进行统计分析,试验数据经正态性检验(Shapiro-Wilk检验)及方差齐性检验后,组间比较采用单因素方差分析(one-way ANOVA),P<0.05时采用Dunn检验进行两两比较;对不符合正态分布或方差不齐的试验数据,组间比较采用Kruskal-Wallis H检验,P<0.05时采用Dunn检验进行两两比较。P<0.05表示显著差异,P>0.05表示差异不显著,数据以平均值和均值标准误(SEM)表示。

2 结果

2.1 瘤胃液移植与饲喂复合益生菌对牦牛生长性能的影响

由表2可知,RT组牦牛的终末体重、总增重和平均日增重与CK组没有显著差异(P>0.05),而CP组牦牛的终末体重、总增重和平均日增重显著高于CK组(P<0.05)。RT组和CP组牦牛的干物质采食量和料重比显著低于CK组(P<0.05)。
表2 瘤胃液移植与饲喂复合益生菌对牦牛生长性能的影响

Table 2 Effects of rumen fluid transplantation and feeding compound probiotics on growth performance of yaks

项目
Items		 组别Groups SEM P值
P-value
RT CP CK
初始体重IBW/kg 76.06 76.01 76.84 0.304 0.285
终末体重FBW/kg 103.05ab 104.55a 99.55b 0.870 0.049
总增重TWG/kg 26.99ab 28.54a 22.71b 0.964 0.032
平均日增重ADG/(g/d) 449.77ab 475.68a 378.50b 16.067 0.032
干物质采食量DMI/(kg/d) 3.86b 3.93b 4.38a 0.019 <0.001
料重比F/G 8.60b 8.27b 11.60a 0.113 <0.001

同行数据肩标不同小写字母表示差异显著(P<0.05),相同或无字母表示差异不显著(P>0.05)。下表同。

In the same row, values with different small letter superscripts mean significant difference (P<0.05), while with the same or no letter superscripts mean no significant difference (P>0.05). The same as below.

2.2 瘤胃液移植与饲喂复合益生菌对牦牛血清生化指标的影响

由表3可知,RT组的血清IL-1β含量与CK组没有显著差异(P>0.05),而CP组的血清IL-1β含量显著低于CK组(P<0.05);RT组和CP组的血清CAT活性显著高于CK组(P<0.05);RT组的血清GSH-Px活性显著高于CK组(P<0.05),而CP组的血清GSH-Px活性与CK组没有显著差异(P>0.05)。
表3 瘤胃液移植与饲喂复合益生菌对牦牛血清生化指标的影响

Table 3 Effects of rumen fluid transplantation and feeding compound probiotics on serum biochemical indices of yaks

项目
Items		 组别Groups SEM P值
P-value
RT CP CK
肿瘤坏死因子-α TNF-α/(pg/mL) 133.82 163.23 136.87 16.568 0.742
白细胞介素-1β IL-1β/(pg/mL) 13.11ab 9.00b 16.17a 1.071 0.013
白细胞介素-6 IL-6/(pg/mL) 36.14 64.08 53.00 10.764 0.505
丙二醛MDA/(nmol/mL) 6.42 3.67 3.19 0.616 0.168
过氧化氢酶CAT/(U/mL) 28.49a 21.77a 9.60b 3.266 0.013
谷胱甘肽过氧化物酶GSH-Px/(U/mL) 19.79a 8.51b 5.47b 2.094 0.004
超氧化物歧化酶SOD/(U/mL) 23.02 16.53 15.83 1.734 0.570
总抗氧化能力T-AOC/(U/mL) 36.00 32.42 19.86 13.041 0.870

2.3 瘤胃液移植与饲喂复合益生菌对牦牛粪便菌群多样性的影响

由图1-A可知,RT组、CP组和CK组牦牛粪便中共鉴别出10 477个扩增子序列变异(ASVs),RT组鉴别出2 948个ASVs,其中独有886个;CP组鉴别出4 050个ASVs,其中独有1 689个;CK组鉴别出3 479个ASVs,其中独有1 212个。由图1-B和1-C可知,RT组、CP组和CK组的覆盖率(goods_coverage)均大于0.99,稀释曲线曲度平缓,表明测序深度能够满足粪便菌群的分析需求;Alpha多样性指数分析显示,CP组的Chao1指数、Observed_features指数、Shannon指数和Simpson指数显著高于RT组和CK组(P<0.05)。由图1-D可知,基于Bray-Curtis距离进行的主坐标分析(PCoA)并结合Adonis检验,RT组、CP组和CK组之间的样本距离分离明显(P<0.05),说明不同处理对牦牛粪便菌群结构的影响显著。
图1 瘤胃液移植与饲喂复合益生菌对牦牛粪便菌群多样性的影响

A:韦恩图;B:菌群稀释曲线;C:Alpha多样性分析;D:主坐标分析(PCoA)三维和二维图。箱线图上标*表示差异显著(P<0.05),ns表示差异不显著(P>0.05)。

Fig.1 Effects of rumen fluid transplantation and feeding compound probiotics on fecal microbiota diversity of yaks

A: Venn diagram; B: bacterial dilution curve; C: Alpha diversity analysis; D: principal coordinate analysis (PCoA) three- and two-dimensional plots. The * in the box plot indicated significant difference (P<0.05), while ns indicated no significant difference (P>0.05).

2.4 瘤胃液移植与饲喂复合益生菌对牦牛粪便菌群门水平相对丰度的影响

由表4可知,在门水平上选取相对丰度前10的菌门,各组牦牛粪便的优势菌门均为拟杆菌门(Bacteroidota)和厚壁菌门(Firmicutes)。CP组和CK组的粪便厚壁菌门相对丰度显著高于RT组(P<0.05),嗜盐菌门(Halobacterota)仅在CP组中出现。
表4 瘤胃液移植与饲喂复合益生菌对牦牛粪便菌群门水平相对丰度的影响

Table 4 Effects of rumen fluid transplantation and feeding compound probiotics on relative abundances of fecal microbiota at phylum level of yaks %

项目
Items		 组别Groups SEM P值
P-value
RT CP CK
厚壁菌门Firmicutes 55.32b 63.15a 61.79a 1.393 0.046
拟杆菌门Bacteroidota 32.05 33.16 26.16 1.350 0.063
螺旋菌门Spirochaetota 9.95 2.10 9.70 1.748 0.107
变形菌门Proteobacteria 0.34 0.11 1.14 0.224 0.168
广古菌门Euryarchaeota 1.82 0.62 0.81 0.209 0.064
疣微菌门Verrucomicrobiota 0.04 0.38 0.04 0.019 0.183
放线菌门Actinobacteriota 0.18 0.11 0.12 0.036 0.993
耐久菌门Patescibacteria 0.25 0.14 0.15 0.036 0.623
纤维杆菌门Fibrobacterota 0.01 0.08 0.02 0.014 0.153
嗜盐菌门Halobacterota 0.00 0.27 0.00 0.008 0.117

2.5 瘤胃液移植与饲喂复合益生菌对牦牛粪便菌群属水平相对丰度的影响

由表5可知,在属水平上选取相对丰度前10的菌属,CK组的粪便UCG-005相对丰度显著高于RT组(P<0.05),CP组的粪便单球菌属(Monoglobus)相对丰度显著高于RT组(P<0.05),RT组的粪便狭义梭菌属1(Clostridium_sensu_stricto_1)相对丰度显著高于CP组(P<0.05)。
表5 瘤胃液移植与饲喂复合益生菌对牦牛粪便菌群属水平相对丰度的影响

Table 5 Effects of rumen fluid transplantation and feeding compound probiotics on relative abundances of fecal microbiota at genus level of yaks %

项目
Items		 组别Groups SEM P值
P-value
RT CP CK
密螺旋体属Treponema 9.78 2.02 9.68 1.762 0.113
理研菌科RC9肠道群Rikenellaceae_RC9_gut_group 12.91 11.67 10.80 0.847 0.629
UCG-005 10.08b 12.74ab 13.82a 0.695 0.032
另枝菌属Alistipes 3.76 4.89 3.96 0.350 0.264
拟杆菌属Bacteroides 3.98 2.98 2.14 0.331 0.065
单球菌属Monoglobus 1.27b 3.46a 1.78ab 0.377 0.049
厌氧孤菌属Anaerovibrio 1.87 0.12 1.34 0.372 0.083
罗博特氏菌属Romboutsia 2.47 1.13 1.14 0.283 0.073
狭义梭菌属1 Clostridium_sensu_stricto_1 3.12a 0.82b 2.13ab 0.394 0.042
克里斯滕森菌科R-7群Christensenellaceae_R-7_group 3.04 3.10 2.96 0.181 0.887

2.6 牦牛粪便菌群LFfSe分析

由图2可知,RT组在密螺旋体属(Treponema)、狭义梭菌属1上显著富集;CP组在单球菌属上显著富集;CK组在UCG_005上显著富集。
图2 牦牛粪便菌群LEfSe分析

g_Treponema:密螺旋体属;g_Clostridium_sensu_stricto_1:狭义梭菌属1;g_Monoglobus:单球菌属。

Fig.2 LEfSe analysis of fecal microbiota of yaks

2.7 牦牛粪便菌群KEGG功能预测

由图3可知,RT组相较于CK组显著富集于翻译,核苷酸代谢,折叠、排序与降解和信号分子与相互作用等代谢途径(P<0.05)。值得注意的是,CP组相较于CK组也同时显著富集于核苷酸代谢,折叠、排序与降解和信号分子与相互作用等代谢途径(P<0.05),此外还显著富集于氨基酸代谢等代谢途径(P<0.05)。
图3 牦牛粪便菌群KEGG功能预测

Means in groups:组间平均值;95% confidence intervals:95%置信区间;Difference between groups:组间差异。

Fig.3 KEGG functional prediction of fecal microbiota of yaks

3 讨论

3.1 瘤胃液移植与饲喂复合益生菌对牦牛生长性能的影响

动物的生长发育状况能够通过平均日增重得到直观的反映,而干物质采食量则是评估动物健康状况和营养摄取能力的关键指标[20-21]。在反刍动物的早期培育和健康管理中,通过微生物干预来调控其生长发育是一项重要的研究策略[22]。张鑫[23]研究发现,接种新鲜的瘤胃液和灭菌瘤胃液对犊牛的增重均没有显著影响。Zhang等[24]研究发现,多次接种瘤胃液对牦牛的增重没有显著影响,但提高了饲料转化效率。在本试验中,瘤胃液移植对牦牛的总增重和平均日增重没有显著影响,但显著降低了干物质采食量和料重比,说明瘤胃液移植提高了牦牛对饲料的利用率,这与以上的研究结果基本一致。茹作娟等[25]研究发现,饲粮中添加复合益生菌使犊牛的增重显著高于对照组,料重比显著低于对照组。Wang等[26]研究表明,饲喂复合益生菌提高了湖羊的平均日增重,降低了料重比,提高了饲料转化效率。在本试验中,饲喂复合益生菌显著提高了牦牛的总增重和平均日增重,显著降低了干物质采食量和料重比,进而提高了饲料转化效率,这与上述的研究结果基本一致。以上研究结果表明,瘤胃液移植与饲喂复合益生菌对犊牛的生长性能上的表现存在差异,前者对增重无显著影响,而后者更容易在降低干物质采食量的同时带来增重与饲料转化效率的双重改善。

3.2 瘤胃液移植与饲喂复合益生菌对牦牛血清生化指标的影响

细胞因子是由免疫细胞合成并分泌的微量多肽类信号分子,它们在细胞间传递信息,调节细胞的生理活动,增强机体的免疫应答能力[27]。TNF-α最初被鉴定为一种导致肿瘤坏死的介质,然而后续研究揭示了其在自身免疫性疾病病理生理过程中的关键作用。白细胞介素在免疫细胞的激活与调节中发挥着至关重要的作用,特别是IL-1β和IL-6,作为与动物机体炎症反应密切相关的促炎细胞因子,其含量的升高可能指示机体遭受氧化应激等不利因素影响,从而引发炎症反应[28]。刘波[29]研究发现,对断奶犊牛饲喂复合益生菌能够有效抑制促炎细胞因子的释放,增强机体免疫力。王海玉等[30]研究发现,喂复合益生菌能够提高羔羊的免疫力,降低炎症反应和腹泻率。在本试验中,CP组牦牛血清促炎细胞因子IL-1β含量显著低于CK组,这与以上研究结果基本一致。
氧化应激是指体内活性氧(ROS)和活性氮(RNS)等自由基的积累超过了抗氧化系统的清除能力,导致细胞和组织的损伤。过量的自由基能够引起脂质过氧化、DNA损伤以及蛋白质功能失调,从而影响机体的正常生理功能[31]。在断奶阶段,牦牛面临从母乳向粗饲料的剧烈过渡,此时肠道微生物群落的失衡常常导致肠道菌群紊乱、免疫系统抑制和氧化应激反应的增强。尤其是在断奶应激期,牦牛的免疫系统尚未完全成熟,肠道屏障功能较弱,容易受到外界因素的影响,进而引发氧化应激、肠道黏膜损伤及系统性炎症反应。研究表明,断奶应激不仅加剧了肠道微生态的紊乱,也促使活性氧种类(如过氧化氢、超氧化物等)增加,从而加重氧化损伤和免疫抑制[32]。CAT和GSH-Px是体内2种关键的抗氧化酶,它们通过清除过氧化氢和其他活性氧种类,有效减少氧化应激对细胞的损害,维持细胞的正常功能[33]。本试验中,RT组和CP组的血清CAT和GSH-Px活性升高,可能是由于瘤胃液移植和饲喂益生菌改善了牦牛肠道微生态环境,促进了抗氧化酶的合成和活性,从而增强了牦牛的抗氧化能力。尽管RT组和CP组的血清CAT活性显著高于CK组,但仅RT组的血清GSH-Px活性显著高于CK组,这可能与2种微生态干预措施的机制差异有关。瘤胃液移植通过引入成熟的微生物群落,可能在短期内更显著地改善了牦牛的抗氧化能力,而复合益生菌则可能需要更长时间才能显著提高某些抗氧化酶活性。这也反映了不同微生态干预措施对牦牛健康的影响机制和效果的差异,短期内通过瘤胃液移植能够快速恢复肠道微生物群的稳定性,从而更有效地缓解氧化应激和免疫抑制,而饲喂益生菌则可能通过长期的微生态调节实现逐步的健康改善。

3.3 瘤胃液移植与饲喂复合益生菌对牦牛粪便菌群多样性的影响

Observed_features指数和Chao1指数表示物种的丰富度,Shannon指数表示菌群的多样性,Simpson指数表示菌群的均匀度,覆盖率表示粪便样本测序的覆盖度,测序覆盖度越高,覆盖率越大。王海玉等[34]研究发现,饲粮中添加复合益生菌显著提高了肠道微生物Chao1指数和Observed_features指数。在本试验中,CP组牦牛粪便菌群中Chao1指数、Observed_features指数、Shannon指数和Simpson指数显著高于CK组,表明饲喂复合益生菌不仅增加了菌群的数量,还促进了菌群的均衡分布,有助于维持肠道生态系统的稳定性,这与以上研究结果基本一致。俞少博[35]研究发现,将新鲜瘤胃液移植给羔羊后,其瘤胃和结肠内的微生物多样性并未产生明显改变。在本试验中,与CK组相比,RT组牦牛粪便中的Alpha多样性指标未表现出显著变化,这与以上研究结果基本一致。

3.4 瘤胃液移植与饲喂复合益生菌对牦牛粪便菌群门水平和属水平相对丰度的影响

肠道微生物群落在反刍动物健康中扮演着至关重要的角色,不仅参与食物的消化和营养吸收,还对免疫系统、代谢功能和病原的防御起着重要作用[36]。在门水平上,拟杆菌门和厚壁菌门作为各组的优势菌门,反映了牦牛肠道微生物群落的核心组成,符合反刍动物肠道菌群的普遍特征[37]。厚壁菌门和拟杆菌门是反刍动物肠道中最常见的菌群,这些菌群在消化代谢、能量获取和肠道健康中具有至关重要的作用。厚壁菌门主要参与复杂碳水化合物和纤维素的降解,它们能够有效地分解植物性食物,生成短链脂肪酸,为宿主提供能量[38]。CP组和CK组的粪便厚壁菌门相对丰度显著高于RT组。这可能源于复合益生菌的持续补充促进了厚壁菌门相关菌群的定植和繁殖[39]。嗜盐菌门仅在CP组中出现,其存在可能与复合益生菌调节肠道微环境有关。嗜盐菌门通常存在于高盐或极端环境,其在肠道中的功能尚未明确[40]。
在属水平上,CK组的粪便UCG-005相对丰度显著高于RT组,UCG-005是反刍动物肠道中重要的纤维降解菌,能够有效利用复杂植物多糖,产生短链脂肪酸,支持宿主能量代谢和肠道屏障功能[41]。CP组的粪便单球菌属相对丰度显著高于RT组,表明饲喂复合益生菌促进了该属菌的定植。单球菌属具有利用果胶和半纤维素参与体内糖醛酸和戊糖的代谢途径[42];同时与健康群落有关,其失调可能与肠道屏障的破坏和促炎细胞因子含量增加有关[43]。益生菌通过改善肠道环境、提供代谢底物,增强了单球菌属的优势地位,从而促进肠道健康。RT组的粪便狭义梭菌属1相对丰度显著高于CP组。狭义梭菌属1可以与其他肠道微生物相互作用,在肠道中可以生成短链脂肪酸,有减轻炎症的效果[44]。此外,狭义梭菌属1可能使RT组牦牛处于能量代谢活跃状态,该菌属可通过发酵途径产生丁酸等有益代谢物[45]。进一步使用LFfSe对属水平上的差异菌属进行分析,除了上述的菌属在3组中显著富集外,还发现密螺旋体属显著富集于RT组中。密螺旋体属在反刍动物中常参与纤维降解和短链脂肪酸合成[46]。
整体看来,RT组牦牛通过引入成熟的瘤胃微生物群落,强化了生成短链脂肪酸的关键菌群如狭义梭菌属1和密螺旋体属,两者在厌氧环境下的有机物分解与能量代谢中具有重要地位,可直接促进乙酸、丁酸等短链脂肪酸生成[47]。短链脂肪酸能够上调抗氧化酶活性,以恢复脂质氧化水平,对维持肠道稳态方面具有关键作用[48]。在本试验中,RT组的血清CAT和GSH-Px活性显著上升,表明瘤胃液移植调节了菌群结构,从而降低了机体断奶后的氧化应激。但同时引发了菌群结构的短期波动,导致部分菌群如UCG-005和厚壁菌门的相对丰度下降,这可能是因为微生物群落的重组导致肠道环境的变化,从而使微生物群落逐渐发生变化。CP组牦牛通过饲喂复合益生菌调节肠道微生态,显著提高了粪便厚壁菌门和单球菌属等有益菌群的相对丰度,促进肠道代谢和免疫功能的协调发展,这与本试验中CP组的血清促炎细胞因子含量降低相吻合。CK组牦牛粪便菌群则维持了较高的UCG-005相对丰度,体现了自然肠道微生态的稳定状态。这些差异反映了2种微生态干预措施在改善牦牛肠道微生物功能和宿主健康上的不同优势和潜力。

3.5 瘤胃液移植与饲喂复合益生菌对牦牛粪便菌群KEGG功能代谢的影响

本研究采用16S rDNA扩增子测序结合PICRUSt预测进行功能分析。RT组和CP组都同时显著富集于核苷酸代谢途径,已有研究指出核苷酸代谢不仅为细胞提供DNA/RNA合成的原料,还通过影响还原型烟酰胺腺嘌呤二核苷酸磷酸及谷胱甘肽的生成调节细胞内活性氧稳态[49]。此外,氧化修饰的核苷酸能够激活核因子E2相关因子2信号通路,从而上调抗氧化酶表达,增强细胞清除自由基的能力[50]。在本研究中,RT组和CP组核苷酸代谢途径的增强可能通过调节氧化还原相关代谢途径提高抗氧化能力。这与本试验中RT组和CP组血清中抗氧化酶活性升高的结果相一致。氨基酸代谢是免疫细胞代谢重编程的核心,调控T细胞、巨噬细胞等免疫细胞的活化、分化及功能[51]。同时氨基酸不仅为免疫细胞提供能量和合成底物,还通过其代谢产物参与信号传导与氧化还原平衡维持[52]。复合益生菌可能通过促进氨基酸代谢活性,增强宿主抗氧化能力并调节免疫反应,从而改善氧化应激与炎症状态。本研究结果不能代表实际功能,此外每组5头牦牛的样本量处于微生物组学研究的最小统计边界,后续需要扩大样本量,这些作用机制需要进行代谢组学或者宏基因组学分析进一步来确定。

4 结论

瘤胃液移植更适用于短期内快速改善牦牛的抗氧化能力并提高饲料利用效率;而饲喂复合益生菌则更有利于长期维持牦牛的肠道菌群稳态,增强抗炎反应调节能力,同时促进牦牛增重和改善饲料利用效率。

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