Chinese Journal of Animal Nutrition >

2025 , Vol. 37 >Issue 7: 4692 - 4701

DOI: https://doi.org/10.12418/CJAN2025.383

RESEARCH PAPER

Effects of Dietary Supplementation with Creatine Pyruvate and Coated Gamma-Aminobutyric Acid on Growth Performance, Rumen Fermentation Parameters and Microflora of Heat-Stressed Goats

  • ZHAO Xingrui ,
  • WANG Peng ,
  • WAN Zibo ,
  • YI Zhonghua ,
  • HE Yong ,
  • KE Di ,
  • SONG Xiaozhen , *
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  • Jiangxi Key Laboratory of Animal Nutrition and Feed, Jiangxi Agricultural University, Nanchang 330045, China
* professor, E-mail:

Received date: 2024-12-19

  Online published: 2025-07-12

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Abstract

This experiment was conducted to study the effects of dietary supplementation with creatine pyruvate (CrPyr) and coated gamma-aminobutyric acid (GABA) on growth performance, rumen fermentation parameters and microflora of heat-stressed goats. A single-factor experimental design was adopted. Fifty-four 4-month-old Nubian black goats in good body condition and with similar body weight [(22.61±0.47) kg] were selected and randomly divided into 3 groups with 3 replicates in each group and 6 goats in each replicate. Goats in control group were fed a basal diet, goats in CrPyr group were fed the basal diet supplemented with 0.400% CrPyr, and goats in GABA group were fed the basal diet supplemented with 0.016% coated GABA. All experimental goats were raised in a high-temperature and high-humidity environment in summer. The pre-trial period lasted for 10 days and the formal trial period lasted for 60 days. The results showed as follows: 1) compared with the control group, the average daily gain and dry matter intake in the CrPyr group and the GABA group from 31 to 60 days were extremely significantly increased (P<0.01), and the feed-to-gain ratio was significantly decreased (P<0.05). 2) Compared with the control group, the pH of rumen fluid in the CrPyr group and the GABA group was significantly decreased (P<0.05), the content of ammonia nitrogen was extremely significantly increased (P<0.01), and the contents of isobutyric acid, butyric acid and isovaleric acid were significantly increased (P<0.05). The contents of acetic acid and total volatile fatty acids in the rumen fluid in the GABA group were significantly higher than those in the control group (P<0.05). 3) Compared with the control group, the Shannon index of rumen microorganisms in the CrPyr group and the GABA group was significantly increased (P<0.05); the relative abundance of Firmicutes in rumen microorganisms was significantly increased (P<0.05), the relative abundance of Verrucomicrobiota was extremely significantly increased (P<0.01), the relative abundance of Bacteroidota was significantly decreased (P<0.05), and the relative abundance of Prevotella was significantly decreased (P<0.05). The relative abundance of Rikenellaceae_RC9_gut_group in rumen microorganisms in the CrPyr group was significantly higher than that in the control group (P<0.05). In conclusion, dietary supplementation with 0.400% CrPyr or 0.016% coated GABA can increase the average daily gain and dry matter intake of goats under heat stress, increase the contents of some volatile fatty acids in rumen fluid and change the rumen microbiota.

Key words： heat stress; goats; creatine pyruvate; gamma-aminobutyric acid; growth performance; rumen fermentation; rumen microorganisms

Cite this article

ZHAO Xingrui , WANG Peng , WAN Zibo , YI Zhonghua , HE Yong , KE Di , SONG Xiaozhen . Effects of Dietary Supplementation with Creatine Pyruvate and Coated Gamma-Aminobutyric Acid on Growth Performance, Rumen Fermentation Parameters and Microflora of Heat-Stressed Goats[J]. Chinese Journal of Animal Nutrition, 2025 , 37(7) : 4692 -4701 . DOI: 10.12418/CJAN2025.383

热应激是指当环境温度高于动物等热区上限时机体所产生的非特异性应答反应的总和。热应激时机体的热平衡被打破,会导致动物自身产生一系列复杂的变化以维持内环境稳态[1]。
南方地区夏季气温高且湿度大,这种高温高湿环境易诱发肉羊的热应激反应。随着肉羊养殖集约化程度的不断提高,养殖密度逐渐增加,热应激对肉羊的负面影响也日益加重。有研究表明,热应激会导致绵羊的呼吸频率和心率加快,直肠温度升高,采食量下降,瘤胃功能紊乱[2-3]。为了缓解热应激对肉羊养殖的不利影响,可在饲粮中添加抗氧化剂、矿物质、维生素等添加剂,以减轻热应激对羊只造成的损伤[4]。
丙酮酸肌酸(creatine pyruvate,CrPyr)是由丙酮酸和肌酸构成的复合物,兼具丙酮酸和肌酸的生物学功能,同时弥补了丙酮酸易导致机体内环境紊乱以及肌酸溶解性差、吸收率低的缺点[5]。丙酮酸肌酸可以促进ATP的合成与释放,还可以调节蛋白质代谢相关激素(如胰岛素、生长激素等)的水平,从而促进蛋白质的合成及抑制肌肉中蛋白质的分解。丙酮酸肌酸中的肌酸部分是动物体重要的供能物质,同时还具有抗氧化和抗炎症作用,可以减轻动物的氧化应激和炎症反应。研究表明,饲粮中添加丙酮酸肌酸能够提高应激肉牛的瘤胃微生物蛋白含量,并促进瘤胃菌群平衡的恢复[6-7]。γ-氨基丁酸(gamma-aminobutyric acid,GABA)是一种广泛存在于动物体内的功能性非蛋白质类氨基酸[8]。它能够增强神经元细胞膜对氯离子的通透性,产生抑制效应,防止神经细胞过度兴奋,从而维持神经系统的平衡;还能够促进垂体释放生长激素,有利于动物生长;并能使毛细血管扩张,降低血压。有研究表明,在种鸡蛋中注射γ-氨基丁酸可以提高肉鸡孵化后的平均日采食量,增强肉鸡肝脏的抗氧化功能[9]。Bi等[10]研究发现,饲粮中添加γ-氨基丁酸可缓解育肥猪的运输应激,降低宰后猪肉的滴水损失,改善肉品质。Chen等[11]研究发现,饲粮中添加包被γ-氨基丁酸能使热应激肉牛的平均日增重和平均日采食量增加,提高了肉牛在热应激条件下的生产性能。目前关于丙酮酸肌酸和γ-氨基丁酸能否缓解山羊热应激、提高生长性能及促进瘤胃发酵的研究报道较少。因此,本试验以热应激条件下的山羊为研究对象,探究饲粮中添加丙酮酸肌酸和包被γ-氨基丁酸对其生长性能、瘤胃发酵参数及瘤胃微生物区系的影响,以期为丙酮酸肌酸和包被γ-氨基丁酸在山羊生产中的应用提供理论依据。

1 材料与方法

1.1 试验材料

丙酮酸肌酸购自上海某医药科技有限公司,其中丙酮酸含量≥30.0%,肌酸含量≥58.0%;包被γ-氨基丁酸由河南某生物技术有限公司赠送,其中γ-氨基丁酸含量>40.0%。

1.2 试验设计

本试验通过江西农业大学实验动物护理和使用委员会的审查和批准(批准编号:JXAULL-20200040)。
采用单因素试验设计,选取54只体况良好、体重[(22.61±0.47) kg]相近的4月龄努比亚黑山羊,随机分为3组,每组3个重复,每个重复6只。对照组饲喂基础饲粮,丙酮酸肌酸组在基础饲粮中添加0.400%的丙酮酸肌酸,γ-氨基丁酸组在基础饲粮中添加0.016%的包被γ-氨基丁酸。试验羊均饲养在夏季高温高湿环境中。试验期70 d,其中预试期10 d,正试期60 d。基础饲粮参考《肉羊营养需要量》(NY/T 816—2021)设计,其组成及营养水平见表1。
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)%

项目 Items 含量 Content
原料 Ingredients
花生秧 Peanut vine 60.00
玉米 Corn 20.00
豆粕 Soybean meal 10.50
麦麸 Wheat bran 7.00
碳酸氢钙 Ca(HCO3)2 0.50
磷酸氢钙 CaHPO4 0.50
石粉 Limestone 0.50
氯化钠 NaCl 0.20
预混料 Premix1) 0.80
合计 Total 100.00
营养水平 Nutrient levels2)
干物质 DM 88.24
代谢能 ME/(MJ/kg) 10.99
粗蛋白质 CP 11.68
粗脂肪 EE 1.99
中性洗涤纤维 NDF 50.62
酸性洗涤纤维 ADF 34.46
粗灰分 Ash 10.64
总磷 TP 0.53
钙 Ca 0.90

1)预混料为每千克饲粮提供The premix provided the following per kg of the diet:VA 4 510 IU,VD 450 IU,VE 72 IU,Fe 125 mg,Cu 300 mg,Mn 50 mg,Zn 45 mg,Co 25 mg,I 77.5 mg,Se 13 mg。
2)代谢能为计算值,其余为实测值。ME was a calculated value, while the others were measured values.

1.3 饲养管理

每组试验羊分为3栏饲养,每栏6只。每日08:00和16:00各喂料1次,先精后粗,羊只自由采食和饮水。粪便每天清扫2次,羊舍每周消毒2~3次。

1.4 样品采集及指标测定

1.4.1 饲粮常规营养成分

于正试期第30天和第60天采集精料和粗料各500 g,制成风干样品后,按照精粗比4:6混合均匀,参考国家标准或行业标准测定饲粮中干物质(GB/T 6435—2014)、粗蛋白质(GB/T 6432—2018)、粗脂肪(GB/T 6433—2006)、中性洗涤纤维(GB/T 20806—2022)、酸性洗涤纤维(NY/T 1459—2022)、粗灰分(GB/T 6438—2007)、总磷(GB/T 6437—2018)、钙(GB/T 6436—2018)的含量;代谢能依据中国饲料数据库中各原料营养价值数据计算,计算公式为:
代谢能=Σ(各原料代谢能值×各原料在饲粮中的比例)。

1.4.2 生长性能

于正试期第1天、第30天和第60天晨饲前对试验羊称重。正试期间每天记录试验羊的采食量。计算第1~30天、第31~60天的干物质采食量、平均日增重和料重比。

1.4.3 瘤胃发酵参数

于正试期第60天晨饲前,每组选取6只(每栏2只)体重适中的山羊,通过口腔抽取瘤胃内容物,通过4层纱布过滤后,取部分滤液立即采用高精度pH计(REXPHS-3C,上海仪电科学仪器股份有限公司)测定pH,其余瘤胃液于-80 ℃冰箱中保存。氨态氮(NH3-N)含量采用比色法[12]测定;微生物蛋白(MCP)含量采用考马斯亮蓝法[13]测定;挥发性脂肪酸(VFA)含量采用气相色谱仪(安捷伦8860)测定[14]。

1.4.4 瘤胃微生物

将正试期第60天采集的瘤胃液样品送至北京诺禾致源科技股份有限公司进行检测。采用16S rRNA基因测序技术测定瘤胃微生物多样性,提取样本的总DNA,按照指定测序区域合成带有barcode的特异性产物,进行PCR扩增与产物纯化、定量和均一化后,根据测序平台的要求构建测序文库,对构建的测序文库进行质量控制,包括检测文库的浓度、纯度和片段大小分布等,确保文库的质量符合测序要求。利用MiSeq PE300平台对文库进行测序,对获得的原始测序数据(raw data)进行质量控制和过滤后,使用QIIME2软件对优化后的序列进行操作分类单元(OTU)聚类分析和物种分类注释,解析微生物多样性。

1.5 数据统计与分析

试验数据用Excel 2021进行初步整理,然后利用SPSS 27.0软件进行单因素方差分析(one-way ANOVA),并采用LSD法进行多重比较。结果用平均值和均值标准误(SEM)表示,P<0.01表示差异极显著,P<0.05表示差异显著,P>0.05表示差异不显著。

2 结果与分析

2.1 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊生长性能的影响

由表2可知,与对照组相比,丙酮酸肌酸组和γ-氨基丁酸组第1~30天的干物质采食量、平均日增重和料重比均无显著变化(P>0.05);第31~60天的干物质采食量和平均日增重均极显著提高(P<0.01),料重比显著降低(P<0.05)。
表2 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊生长性能的影响

Table 2 Effects of CrPyr and coated GABA on growth performance of heat-stressed goats

项目
Items		 对照组
Control group		 丙酮酸肌酸组
CrPyr group		 γ-氨基丁酸组
GABA group		 SEM P值
P-value
干物质采食量 DMI/(g/d)
第1~30天Days 1 to 30 878.13 881.00 884.34 11.192 0.981
第31~60天 Days 31 to 60 859.93Bb 987.95Aa 963.87Aa 15.358 0.007
平均日增重 ADG/(g/d)
第1~30天Days 1 to 30 121.89 111.78 141.56 6.065 0.134
第31~60天 Days 31 to 60 81.89Bb 143.45Aa 117.67Aa 4.363 0.004
料重比 F/G
第1~30天Days 1 to 30 7.20 7.98 6.25 0.353 0.128
第31~60天 Days 31 to 60 10.77a 6.91b 8.27b 0.686 0.037

同行数据肩标无字母或相同小写字母表示差异不显著(P>0.05),不同小写字母表示差异显著(P<0.05),不同大写字母表示差异极显著(P<0.01)。下表同。

In the same row, values with no letter or the same small letter superscripts mean no significant difference (P>0.05), while with different small letter superscripts mean significant difference (P<0.05), and with different capital letter superscripts mean extremely significant difference (P<0.01). The same as below.

2.2 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊瘤胃发酵参数的影响

由表3可知,与对照组相比,丙酮酸肌酸组和γ-氨基丁酸组瘤胃液pH显著降低(P<0.05),氨态氮含量极显著提高(P<0.01),异丁酸、丁酸和异戊酸含量显著提高(P<0.05)。γ-氨基丁酸组瘤胃液乙酸和总挥发性脂肪酸含量显著高于对照组(P<0.05)。
表3 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊瘤胃发酵参数的影响

Table 3 Effects of CrPyr and coated GABA on rumen fermentation parameters of heat-stressed goats

项目
Items		 对照组
Control group		 丙酮酸肌酸组
CrPyr group		 γ-氨基丁酸组
GABA group		 SEM P值
P-value
pH 6.93a 6.88b 6.70b 0.042 0.013
氨态氮 NH3-N/(mg/dL) 5.58Bb 6.31Aa 6.28Aa 0.109 0.004
微生物蛋白 MCP/(mg/mL) 0.37 0.36 0.37 0.002 0.228
乙酸 Acetic acid/(mmol/L) 19.59b 24.78ab 30.38a 1.543 0.014
丙酸 Propionic acid/(mmol/L) 4.18 4.84 5.21 0.278 0.347
异丁酸 Isobutyric acid/(mmol/L) 0.37b 0.48a 0.49a 0.022 0.044
丁酸 Butyric acid/(mmol/L) 3.34b 4.34a 4.93a 0.236 0.012
异戊酸 Isovaleric acid/(mmol/L) 0.69b 0.86a 0.97a 0.037 0.013
戊酸 Valeric acid/(mmol/L) 0.36 0.36 0.40 0.024 0.749
乙酸/丙酸 Acetic acid/propionic acid 5.04 5.25 5.81 0.241 0.438
总挥发性脂肪酸 TVFA/(mmol/L) 28.54b 35.68ab 42.38a 2.001 0.011

2.3 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊瘤胃微生物区系的影响

2.3.1 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊瘤胃微生物Alpha多样性的影响

由表4可知,与对照组相比,丙酮酸肌酸组和γ-氨基丁酸组瘤胃微生物Shannon指数显著提高(P<0.05),瘤胃微生物Ace、Chao、Simpson和Sobs指数无显著变化(P>0.05)。各组瘤胃微生物覆盖率(Coverage)均大于99.9%,说明测序样品取样充分,能够准确反映山羊瘤胃微生物的种类与结构组成。
表4 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊瘤胃微生物Alpha多样性的影响

Table 4 Effects of CrPyr and coated GABA on Alpha diversity of rumen microorganisms of heat-stressed goats

项目
Items		 对照组
Control group		 丙酮酸肌酸组
CrPyr group		 γ-氨基丁酸组
GABA group		 SEM P值
P-value
Ace指数 Ace index 1 349.37 1 417.42 1 640.39 73.418 0.248
Chao指数 Chao index 1 327.26 1 418.89 1 631.76 73.187 0.231
覆盖率 Coverage/% 99.98 99.98 99.98 0.000 0.294
Shannon指数 Shannon index 4.77b 5.31a 5.54a 0.144 0.047
Simpson指数 Simpson index 0.04 0.01 0.01 0.014 0.100
Sobs指数 Sobs index 1 085.67 1 147.67 1 326.00 58.221 0.232

2.3.2 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊瘤胃微生物门水平相对丰度的影响

由表5可知,厚壁菌门(Firmicutes)和拟杆菌门(Bacteroidota)是各组瘤胃微生物的优势菌门,两者合计占总细菌数量的93%~94%。与对照组相比,丙酮酸肌酸组和γ-氨基丁酸组瘤胃微生物中厚壁菌门(Firmicutes)相对丰度显著提高(P<0.05),疣微菌门(Verrucomicrobiota)相对丰度极显著提高(P<0.01),拟杆菌门(Bacteroidota)相对丰度显著降低(P<0.05)。
表5 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊瘤胃微生物门水平相对丰度的影响

Table 5 Effects of CrPyr and coated GABA on relative abundances of rumen microorganisms at phylum level of heat-stressed goats%

项目
Items		 对照组
Control group		 丙酮酸肌酸组
CrPyr group		 γ-氨基丁酸组
GABA group		 SEM P值
P-value
厚壁菌门 Firmicutes 38.26b 49.66a 52.36a 2.467 0.040
拟杆菌门 Bacteroidota 54.56a 43.75b 41.01b 1.558 0.011
酸杆菌门 Acidobacteria 5.39 3.31 3.70 1.090 0.731
变形菌门 Proteobacteria 0.51 0.64 0.69 0.064 0.539
螺旋体门 Spirochaetota 0.53 0.76 0.36 0.141 0.517
髌骨细菌门 Patescibacteria 0.28 0.58 0.52 0.067 0.202
脱硫杆菌门 Desulfobacterota 0.13b 0.26ab 0.39a 0.043 0.039
未分类细菌 Unclassified bacteria 0.13b 0.37a 0.24a 0.011 0.031
疣微菌门 Verrucomicrobiota 0.09Bb 0.18Aa 0.36Aa 0.044 0.006
互养菌门 Synergistota 0.07 0.16 0.21 0.068 0.701
其他 Others 0.16 0.39 0.24 0.052 0.258

2.3.3 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊瘤胃微生物属水平相对丰度的影响

由表6可知,与对照组相比,丙酮酸肌酸组瘤胃微生物中理研菌科RC9肠道群(Rikenellaceae_RC9_gut_group)相对丰度显著提高(P<0.05);丙酮酸肌酸组和γ-氨基酸组瘤胃微生物中普雷沃氏菌属(Prevotella)相对丰度显著降低(P<0.05)。
表6 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊瘤胃微生物属水平相对丰度的影响

Table 6 Effects of CrPyr and coated GABA on relative abundances of rumen microorganisms at genus level of heat-stressed goats%

项目
Items		 对照组
Control group		 丙酮酸肌酸组
CrPyr group		 γ-氨基丁酸组
GABA group		 SEM P值
P-value
理研菌科RC9肠道群
Rikenellaceae_RC9_gut_group		 12.48b 16.62a 12.18b 0.208 0.018
普雷沃氏菌属 Prevotella 22.54a 8.11b 7.83b 0.348 0.027
未知属F082科 Norank_f_F082 6.21 6.80 8.35 0.902 0.631
克里斯滕森菌科R-7群
Christensenellaceae_R-7_group		 3.31 5.56 6.26 0.650 0.163
毛螺菌科NK3A20群
Lachnospiraceae_NK3A20_group		 2.80 4.13 5.63 0.503 0.057
未知属拟杆菌目RF16群
Norank_f_Bacteroidales_RF16_group		 3.87 3.77 3.38 0.590 0.940
双歧杆菌属 Bifidobacterium 4.79 2.20 2.33 0.888 0.585
未分类UCG-003 Unclassified UCG-003 2.00 4.01 2.84 0.635 0.459
瘤胃球菌科NK4A214群
Ruminococcaceae_NK4A214_group		 2.23 2.52 4.04 0.446 0.231
未分类UCG-004 Unclassified UCG-004 2.22 4.63 1.26 0.775 0.202
其他 Others 37.56 41.66 45.91 1.441 0.053

3 讨论

3.1 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊生长性能的影响

热应激可引起肉羊体温和呼吸频率异常升高,对粗饲料的利用效率下降,生长速度减缓[15]。研究发现,丙酮酸肌酸可促进能量的产生,维持动物的耐力,增强动物对应激的抵抗能力。丙酮酸肌酸在动物体内可分解为丙酮酸和肌酸,丙酮酸是动物能量代谢的关键物质,会影响糖代谢与脂代谢;肌酸则是动物体内重要的供能和储能物质,因此丙酮酸肌酸对动物生长有着重要影响[16-17]。本试验结果表明,饲粮中添加丙酮酸肌酸可极显著提高试验后期第31~60天山羊的平均日增重和干物质采食量,显著降低料重比,提高了热应激条件下山羊的生长性能。γ-氨基丁酸是动物体内主要的抑制性神经递质之一,能够减缓动物的呼吸频率,缓解应激对动物的伤害。它还可以促进动物生长激素的分泌,使动物的采食神经中枢兴奋,从而提高动物的采食量和生长速度[18]。研究表明,饲粮中添加γ-氨基丁酸可以改善热应激条件下肉鸡的采食量和体重,且改善效果与γ-氨基丁酸的添加剂量及饲喂时长相关[19]。吴欣等[20]研究发现,饲粮中添加γ-氨基丁酸能够显著提高绵羊的平均日采食量,极显著降低料重比,增加平均日增重,本研究结果与之相符。

3.2 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊瘤胃发酵参数的影响

瘤胃液pH是评估瘤胃发酵状态的重要指标,瘤胃液pH受多种因素影响,如饲粮结构、唾液分泌量、瘤胃挥发性脂肪酸含量和瘤胃吸收速度等[21]。反刍动物瘤胃液的pH通常为6.0~7.5,本试验中,对照组、丙酮酸肌酸组和γ-氨基丁酸组山羊瘤胃液pH均在正常范围之内。氨态氮是瘤胃微生物利用的主要氮源,瘤胃液中氨态氮含量过高或过低均会对瘤胃微生物发酵产生负面影响[22]。本试验结果表明,饲粮中添加丙酮酸肌酸或包被γ-氨基丁酸均极显著提高了山羊瘤胃液氨态氮含量,这可能与瘤胃微生物的组成有关。有研究表明,瘤胃微生物中的普雷沃氏菌更倾向于利用氨态氮和肽作为氮源,而不是游离氨基酸[23]。本试验中,丙酮酸肌酸组和γ-氨基丁酸组瘤胃普雷沃氏菌属相对丰度显著低于对照组,而瘤胃液氨态氮含量极显著高于对照组,与上述研究结果相符。Li等[24]研究发现,在饲粮中添加丙酮酸肌酸可以提高肉牛瘤胃微生物蛋白的合成量和抗氧化活性,有助于缓解氧化应激,改善瘤胃发酵功能。饲粮中的营养物质在瘤胃中发酵后,主要以挥发性脂肪酸和微生物蛋白的形式被动物利用。其中,挥发性脂肪酸是反刍动物主要的能源物质,微生物蛋白是反刍动物主要的蛋白质来源[25]。Wang等[26]研究发现,饲粮中添加γ-氨基丁酸可以优化瘤胃的发酵环境,增加瘤胃中乙酸和丁酸的比例,提高瘤胃发酵中挥发性脂肪酸的产量。本试验结果表明,饲粮中添加丙酮酸肌酸或包被γ-氨基丁酸均显著提高了瘤胃液中丁酸和异丁酸的含量。丁酸作为瘤胃发酵的主要产物之一,可以被瘤胃上皮优先吸收,进而促进瘤胃上皮细胞的增殖和分化,提升瘤胃对营养物质的吸收能力[27]。

3.3 丙酮酸肌酸和包被γ-氨基丁酸对热应激山羊瘤胃微生物区系的影响

瘤胃微生物区系是由细菌、真菌、古菌以及原虫等微生物组成的复杂生态系统,瘤胃菌群的稳定对瘤胃发酵和营养物质吸收有着重要影响[28]。饲粮中添加丙酮酸肌酸或γ-氨基丁酸都可以提高动物对应激的耐受性,减少应激对瘤胃微生物的不利影响。在微生物Alpha多样性分析中,Coverage用于评估样本中物种丰富度的覆盖情况。本试验中,各组瘤胃微生物Coverage均大于99.9%,表明测序数据充足。Shannon指数与微生物多样性呈正相关,本试验中,丙酮酸肌酸组和γ-氨基丁酸组瘤胃微生物Shannon指数均显著高于对照组,表明饲粮中添加丙酮酸肌酸或包被γ-氨基丁酸均能提高热应激山羊瘤胃微生物多样性。研究发现,反刍动物生长阶段瘤胃微生物中的优势菌门为厚壁菌门和拟杆菌门,本研究结果与其一致[29]。厚壁菌门是瘤胃利用粗饲料过程中的关键菌群,主要参与纤维素和半纤维素的降解利用;拟杆菌门在反刍动物瘤胃发酵过程中主要参与非纤维植物成分的分解和吸收[30]。有研究表明,厚壁菌门相对丰度的增加和拟杆菌门相对丰度的减少可以促进山羊生长[31]。本试验结果表明,饲粮中添加丙酮酸肌酸或包被γ-氨基丁酸均显著提高了热应激山羊瘤胃中厚壁菌门相对丰度,显著降低拟杆菌门相对丰度,从而能够促进山羊对粗饲料的消化和吸收。本试验中,饲粮中添加丙酮酸肌酸显著提高了瘤胃中理研菌科RC9肠道群相对丰度。研究表明,理研菌科RC9肠道群可以中和细胞的活性氧,保护细胞免受氧化应激,从而降低细胞的炎症反应[32]。理研菌科RC9肠道群还被证明在碳水化合物和蛋白质降解上有着重要的作用[33]。研究发现,普雷沃氏菌属有助于蛋白质的降解,能够提高瘤胃的消化吸收效率[34]。本试验结果表明,普雷沃氏菌属在对照组、丙酮酸肌酸组和γ-氨基丁酸组的瘤胃微生物中均为优势菌属,而且丙酮酸肌酸组和γ-氨基丁酸组瘤胃雷沃氏菌属相对丰度显著低于对照组,这可能与普雷沃氏菌属的耐受性有关。综上所述,饲粮中添加丙酮酸肌酸或包被γ-氨基丁酸可以提高热应激山羊瘤胃厚壁菌门相对丰度,促进瘤胃发酵。

4 结论

饲粮中添加0.400%丙酮酸肌酸或0.016%包被γ-氨基丁酸均能提高热应激条件下山羊的平均日增重和干物质采食量,提高瘤胃液中部分挥发性脂肪酸含量,改变瘤胃微生物区系。

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