动物营养学报 >

2023 , Vol. 35 >Issue 9: 5619 - 5628

DOI: https://doi.org/10.12418/CJAN2023.518

研究论文

胍基乙酸对育肥猪生长性能、屠宰性能、血清生化指标、肌纤维特性及肌肉发育相关调节因子基因表达的影响

  • 段浩楠 , 1 ,
  • 武殿阁 1 ,
  • 申帅峰 1 ,
  • 王雨佳 1 ,
  • 闫晓栋 2 ,
  • 陈宝江 , 1, *
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  • 1 河北农业大学动物科技学院,保定 071001
  • 2 河北广瑞生物制品有限公司,石家庄 051500
* 陈宝江,教授,博士生导师,E-mail:

段浩楠(1997—),男,河北邯郸人,硕士研究生,研究方向为动物营养与饲料科学。E-mail:

Copy editor: 武海龙

收稿日期: 2023-04-04

  网络出版日期: 2023-09-13

基金资助

河北省科技攻关项目——仔猪专用功能性饲料添加剂产品研发与应用(21326606D)

企业横向项目——胍基乙酸在猪饲料中的应用研究

收起

Effects of Guanidinoacetic Acid on Growth Performance, Slaughter Performance, Serum Biochemical Indexes, Muscle Fiber Characteristics and Gene Expression of Regulatory Factors Related to Muscle Development of Finishing Pigs

  • DUAN Haonan , 1 ,
  • WU Diange 1 ,
  • SHEN Shuaifeng 1 ,
  • WANG Yujia 1 ,
  • YAN Xiaodong 2 ,
  • CHEN Baojiang , 1, *
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  • 1 College of Animal Science and Technology, Hebei Agricultural University, Baoding 071001, China
  • 2 Hebei Granray Bioproducts Co., Ltd., Shijiazhuang 051500, China
* professor, E-mail:

Received date: 2023-04-04

  Online published: 2023-09-13

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摘要

本试验旨在研究饲粮中添加胍基乙酸对育肥猪生长性能、屠宰性能、血清生化指标、肌纤维特性及肌肉发育相关调节因子基因表达的影响。选取体重[(86.00±1.00) kg]相近的杜长大三元杂交猪48头,随机分为3组,每组4个重复,每个重复4头猪(公母各占1/2)。对照组饲喂基础饲粮,T1组在基础饲粮中添加500 mg/kg胍基乙酸,T2组在基础饲粮中添加500 mg/kg胍基乙酸+200 mg/kg甜菜碱。预试期3 d,正试期38 d。结果表明:1)T1组和T2组的末重、平均日增重和平均日采食量显著高于对照组(P<0.05);T1组和T2组的料重比显著低于对照组(P<0.05),且T2组的料重比显著低于T1组(P<0.05)。2)与对照组相比,T2组的胴体重有升高的趋势(P=0.079)。T1组和T2组的屠宰率和眼肌面积显著高于对照组(P<0.05)。3)与对照组相比,T1组和T2组的第19天血清低密度脂蛋白含量有降低的趋势(P=0.084)。T1组和T2组的第38天血清尿素含量显著低于对照组(P<0.05);T1组和T2组的第38天血清总蛋白含量显著高于对照组(P<0.05),且T2组的第38天血清总蛋白含量显著低于T1组(P<0.05)。4)T1组和T2组的肌纤维横截面积显著高于对照组(P<0.05)。5)与对照组相比,T2组的肌肉生肌因子5(Myf5)基因相对表达量有升高的趋势(P=0.093)。T2组的肌肉肌细胞生成素(MyoG)基因相对表达量显著高于对照组(P<0.05)。综上所述,饲粮中添加胍基乙酸和甜菜碱能提高育肥猪生长性能、屠宰性能和肌纤维横截面积,促进肌纤维的增殖与分化,且联合添加胍基乙酸与甜菜碱的促生长效果优于单独添加胍基乙酸。

关键词： 胍基乙酸; 育肥猪; 生长性能; 肌纤维特性; 基因表达

本文引用格式

段浩楠 , 武殿阁 , 申帅峰 , 王雨佳 , 闫晓栋 , 陈宝江 . 胍基乙酸对育肥猪生长性能、屠宰性能、血清生化指标、肌纤维特性及肌肉发育相关调节因子基因表达的影响[J]. 动物营养学报, 2023 , 35(9) : 5619 -5628 . DOI: 10.12418/CJAN2023.518

Abstract

This experiment was conducted to investigate the effects of guanidinoacetic acid on growth performance, slaughter performance, serum biochemical indexes, muscle fiber characteristics and gene expression of regulatory factors related to muscle development of finishing pigs. Forty-eight Duroc-Landrace-White crossbred pigs with similar body weight [(86.00±1.00) kg] were selected and randomly divided into three groups with four replicates in each group and four pigs in each replicate (half male and half female). The control group was fed a basal diet, the T1 group was fed the basal diet supplemented with 500 mg/kg guanidinoacetic acid, and the T2 group was fed the basal diet supplemented with 500 mg/kg guanidinoacetic acid+200 mg/kg betaine. The pre-feeding period was 3 days and the formal period was 38 days. The results showed as follows: 1) the final body weight, average daily gain and average daily feed intake of T1 and T2 groups were significantly higher than those of the control group (P<0.05); the feed to gain ratio of T1 and T2 groups was significantly lower than that of the control group (P<0.05), and the feed to gain ratio of T2 group was significantly lower than that of T1 group (P<0.05). 2) Compared with the control group, the carcass weight of T2 group tended to be increased (P=0.079). The dressing percentage and eye muscle area of T1 and T2 groups were significantly higher than those of the control group (P<0.05). 3) Compared with the control group, the serum low-density lipoprotein content on day 19 of T1 and T2 groups tended to be decreased (P=0.084). The serum urea content on day 38 of T1 and T2 groups was significantly lower than that of the control group (P<0.05); the serum total protein content on day 38 of T1 and T2 groups was significantly higher than that of the control group (P<0.05), and the serum total protein content on day 38 of T2 group was significantly lower than that of T1 group (P<0.05). 4) The muscle fiber cross-sectional area of T1 and T2 groups was significantly higher than that of the control group (P<0.05). 5) Compared with the control group, the gene relative expression level of myogenic factor 5 (Myf5) in muscle of T2 group tended to be increased (P=0.093). The gene relative expression level of myogenin (MyoG) in muscle of T2 group was significantly higher than that of the control group (P<0.05). In conclusion, the addition of guanidinoacetic acid and betaine in the diet can improve the growth performance, slaughter performance and muscle fiber cross-sectional area of finishing pigs, and promote the proliferation and differentiation of muscle fiber, and the combined addition of guanidinoacetic acid and betaine to promote growth is better than the addition of guanidinoacetic acid alone.

Key words： guanidinoacetic acid; finishing pigs; growth performance; muscle fiber characteristics; gene expression

提高畜禽生长性能是畜牧业永恒的目标,也是当今社会和消费者的共同诉求。我国是世界上猪肉最大的生产国和消费国[1],随着经济的发展,消费者对猪肉瘦肉率的需求不断提高。因此,提高猪肉,特别是瘦肉产量是生猪养殖的主要趋势[2]。瘦肉率的提高与骨骼肌的发育密切相关,骨骼肌的基本功能单位是肌纤维,其发育与成肌细胞的增殖和分化密切相关[3]。肌纤维的发育受营养水平、外源性营养补充、运动等因素的调节。近年来,应用外源性营养补充调控瘦肉率的研究受到越来越多的关注[4]。
胍基乙酸(guanidine acetic acid,GAA)又名胍乙酸[5],是一种新型营养补充剂,是合成肌酸(creatine,Cr)及磷酸肌酸在体内的直接前体。补充胍基乙酸能提高机体肌酸、磷酸肌酸和ATP含量[6⇓-8],为动物生长和肌纤维发育提供了更多的能量。此外,胍基乙酸还具有提高动物肉品质、屠宰性能、抗氧化能力以及改善能量代谢等作用[9⇓⇓-12]。胍基乙酸由于安全无害、稳定性好、易于吸收、价格低廉,被应用于动物生产,并表现出良好效果[13-14]。胍基乙酸合成肌酸过程中需要甲基供体提供甲基,同时会生成同型半胱氨酸[15]。甜菜碱又称三甲基甘氨酸,能将自身甲基转移给同型半胱氨酸进而重新生成蛋氨酸,促进胍基乙酸合成肌酸[16]。目前胍基乙酸和甜菜碱在动物生产上联合应用主要集中于改善能量代谢、瘤胃发酵和肉品质[16-17],而对肌肉生长研究较少。因此,本试验通过研究单独添加胍基乙酸及联合添加胍基乙酸和甜菜碱对育肥猪生长性能和肌纤维发育的影响,为胍基乙酸在动物生产中提供科学依据。

1 材料与方法

1.1 试验材料

胍基乙酸(有效含量≥98%)和甜菜碱(有效含量≥98%)均由石家庄某生物制品有限公司提供。

1.2 试验设计

试验选择体重[(86.00±1.00) kg]相近的杜长大三元杂交育肥猪48头,随机分为3组,每组4个重复,每个重复4只猪(公母各占1/2)。对照组饲喂基础饲粮,T1组在基础饲粮中添加500 mg/kg胍基乙酸,T2组在基础饲粮中添加500 mg/kg胍基乙酸+200 mg/kg甜菜碱。预试期3 d;正试期从预试期结束至对照组育肥猪达到110 kg出栏体重,为38 d。基础饲粮参照NRC(2012)肥育猪的营养需要配制,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
玉米 Corn 60.00
小麦 Wheat 12.00
干酒糟及其可溶物 DDGS 5.00
豆粕 Soybean meal 18.00
石粉 Limestone 1.15
磷酸氢钙 CaHPO4 1.20
L-赖氨酸盐酸盐 L-Lys·HCl 0.30
食盐 NaCl 0.35
预混料 Premix1) 2.00
合计 Total 100.00
营养水平 Nutrient levels2)
消化能 DE/(MJ/kg) 13.87
粗蛋白质 CP 14.59
钙 Ca 0.56
总磷 TP 0.52
有效磷 AP 0.27
赖氨酸 Lys 0.82
蛋氨酸 Met 0.25
蛋氨酸+半胱氨酸 Met+Cys 0.50
苏氨酸 Thr 0.50
色氨酸 Try 0.14

1)预混料为每千克饲粮提供 The premix provided the following per kg of the diet:VA 3 000 IU,VB1 2.5 mg,VB2 6.5 mg,VB6 3.0 mg,VB12 0.025 mg,VD3 300 IU,VE 38.5 mg,VK3 1.35 mg,烟酸 nicotinic acid 25 mg,泛酸 pantothenic acid 15 mg,生物素 biotin 0.75 mg,Cu (as copper sulfate) 5 mg,Fe (as ferrous sulfate) 100 mg,Zn (as zinc sulfate) 80 mg,Mn (as manganese sulfate) 20 mg,I (as potassium iodide) 0.50 mg,Se (as sodium selenite) 0.30 mg。

2)消化能为计算值,其余为实测值。DE was a calculated value, while the others were measured values.

1.3 饲养管理

试验于邯郸市曲周县中道农牧有限公司进行。试验前做好消毒准备工作,试验正式开始后,每日饲喂2次,自由饮水。

1.4 指标测定

1.4.1 生长性能

分别于试验开始和结束时称重并记录,记录每天采食量,计算平均日增重(average daily gain,ADG)、平均日采食量(average daily feed intake,ADFI)和料重比(feed to gain ratio,F/G)。

1.4.2 屠宰性能

试验结束后,每个重复选取1头接近平均体重的育肥猪进行屠宰,参考《猪胴体性状测定技术规范》(NY/T 825—2004)[18]测定宰前活重、胴体重、屠宰率、平均背膘厚度、眼肌面积、胴体直长和胴体斜长。

1.4.3 血清生化指标

正式试验第19天和第38天,每个重复选取1头接近平均体重的育肥猪,前腔静脉采血5 mL,3 500 r/min离心10 min,取上清液后分装于离心管,-80 ℃保存。血清总蛋白、白蛋白、尿素、葡萄糖、总胆固醇、高密度脂蛋白、低密度脂蛋白、甘油三酯含量均采用江苏酶免实业有限公司试剂盒测定,按照试剂盒说明书进行操作。

1.4.4 肌纤维特性

取经10%中性甲醛溶液固定的背最长肌样品,依次经过脱水、修剪、包埋、切片、染色和封片后,拍摄图像并采集200倍图片,通过Motic Images Advanced 3.2进行分析,测量肌纤维直径和横截面积。

1.4.5 肌肉发育相关调节因子基因表达

屠宰后采集背最长肌肉样,立即放入液氮罐冻存,随后保存于-80 ℃冰箱。采用Trizon法提取背最长肌中总RNA后检测其浓度、纯度和完整性。检测合格后用PrimeScriptTM RT试剂盒(TaKaRa)逆转录合成cDNA,所有引物由上海派森诺生物科技股份有限公司合成,实时荧光定量PCR引物序列见表2。采用EDC-810型PCR仪(北京东胜创新生物科技有限公司)进行分析。反应体系:cDNA 1 μL,上、下游引物各0.4 μL,2×SYBR real-time PCR premixture 10 μL,补加RNase free ddH2O至20 μL。以甘油醛-3-磷酸脱氢酶(GAPDH)为内参基因,采用2-△△Ct法[19]计算肌肉发育相关调节因子[肌细胞生成素(MyoG)、生肌决定因子(MyoD)、生肌因子5(Myf5)和肌肉生长抑制素(MSTN)]基因相对表达量。
表2 实时荧光定量PCR引物序列

Table 2 Primer sequences of real-time quantitative PCR

基因名称
Gene names		 GenBank登录号
GenBank accession No.		 引物序列
Primer sequences (5'—3')
肌细胞生成素
MyoG		 NM_001012406 F:CAGGGGATCATCTGCTCACA
R:TTGGGCATGGTTTCATCTGG
生肌决定因子
MyoD		 NM_001002824 F:TCCGACGGCATGATGGATTA
R:GAGATGCTCTCCACGATGCT
生肌因子5
Myf5		 NM_001278775.1 F:AAGCGTGCAAGAGGAAATCC
R:CTCTGGTTGGGGTTAGTCGT
肌肉生长抑制素
MSTN		 NM_214435.2 F:CTTGACATGAACCCAGGCAC
R:TTCAGCCCATCTTCTCCTGG
甘油醛-3-磷酸脱氢酶
GAPDH		 XM_021091114.1 F:TCGGAGTGAACGGATTTGGC
R:TGCCGTGGGTGGAATCATAC

1.5 统计分析

采用SPSS 19.0软件进行单因素方差分析,采用Duncan氏法进行多重比较,试验结果用“平均值±标准误”表示,P<0.05表示差异显著,0.05≤P<0.10表示有趋势。

2 结果

2.1 胍基乙酸对育肥猪生长性能的影响

由表3可知,T1组和T2组的末重、平均日增重和平均日采食量显著高于对照组(P<0.05);T1组和T2组的料重比显著低于对照组(P<0.05),且T2组的料重比显著低于T1组(P<0.05)。
表3 胍基乙酸对育肥猪生长性能的影响

Table 3 Effects of guanidinoacetic acid on growth performance of finishing pigs

项目
Items		 组别 Groups P值
P-value
对照 Control T1 T2
初重 IBW/kg 86.41±0.22 86.35±0.24 85.97±0.46 0.604
末重 FBW/kg 113.13±0.69b 120.75±1.56a 123.06±0.41a <0.001
平均日增重 ADG/g 703.09±13.86b 905.40±39.61a 976.12±9.06a <0.001
平均日采食量 ADFI/kg 2.45±0.07b 2.86±0.11a 2.79±0.11a 0.030
料重比 F/G 3.48±0.11a 3.16±0.07b 2.86±0.10c 0.004

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 胍基乙酸对育肥猪屠宰性能的影响

由表4可知,与对照组相比,T2组的胴体重有升高的趋势(P=0.079)。T1组和T2组的屠宰率和眼肌面积显著高于对照组(P<0.05)。各组之间宰前活重、背膘厚度、胴体直长、胴体斜长均无显著差异(P>0.05)。
表4 胍基乙酸对育肥猪屠宰性能的影响

Table 4 Effects of guanidinoacetic acid on slaughter performance of finishing pigs

项目
Items		 组别 Groups P值
P-value
对照 Control T1 T2
宰前活重 Live weight before slaughter/kg 117.33±1.45 120.67±3.67 125.00±2.04 0.150
胴体重 Carcass weight/kg 87.67±1.17 91.17±2.67 94.87±1.68 0.079
屠宰率 Dressing percentage/% 74.71±0.26b 75.56±0.08a 75.90±0.30a 0.036
眼肌面积 Eye muscle area/cm2 44.40±1.64b 52.36±2.11a 51.32±2.35a 0.046
背膘厚度 Back fat depth/mm 21.19±1.43 23.23±2.49 22.69±1.25 0.717
胴体斜长 Carcass diagonal length/cm 96.00±1.15 94.00±1.00 97.25±2.21 0.459
胴体直长 Carcass straight length/cm 108.67±0.88 105.33±2.33 109.25±2.25 0.406

2.3 胍基乙酸对育肥猪血清生化指标的影响

由表5可知,与对照组相比,T1组和T2组的第19天血清低密度脂蛋白含量有降低的趋势(P=0.084)。T1组和T2组的第38天血清尿素含量显著低于对照组(P<0.05);T1组和T2组的第38天血清总蛋白含量显著高于对照组(P<0.05),且T2组的第38天血清总蛋白含量显著低于T1组(P<0.05)。
表5 胍基乙酸对育肥猪血清生化指标的影响

Table 5 Effects of guanidinoacetic acid on serum biochemical indexes of finishing pigs

项目
Items		 组别 Groups P值
P-value
对照 Control T1 T2
第19天 Day 19
葡萄糖 GLU/(mg/mL) 0.59±0.02 0.64±0.02 0.61±0.03 0.365
总胆固醇 TC/(μg/mL) 95.76±2.19 97.00±2.50 98.67±2.48 0.699
尿素氮 Urea/(mg/L) 20.20±0.87 21.64±0.46 21.08±1.12 0.576
总蛋白 TP/(mg/mL) 3.65±0.02 3.52±0.11 3.77±0.11 0.273
白蛋白 ALB/(mg/mL) 26.91±0.49 25.70±0.93 27.36±1.69 0.591
高密度脂蛋白 HDL/(mmol/L) 1.72±0.18 1.68±0.22 2.05±0.20 0.387
低密度脂蛋白 LDL/(mmol/L) 4.63±0.10 3.02±0.50 3.08±0.52 0.084
甘油三酯 TG/(μg/mL) 367.84±8.89 388.38±17.41 413.19±25.83 0.279
第38天 Day 38
葡萄糖 GLU/(mg/mL) 0.70±0.03 0.74±0.01 0.74±0.03 0.668
总胆固醇 TC/(μg/mL) 110.35±1.87 107.43±1.58 108.79±1.25 0.459
尿素氮 Urea/(mg/L) 25.75±0.49a 22.96±0.73b 22.77±0.81b 0.023
总蛋白 TP/(mg/mL) 3.68±0.10c 4.37±0.05a 4.02±0.03b <0.001
白蛋白 ALB/(mg/mL) 31.15±1.35 28.03±0.47 30.86±1.56 0.283
高密度脂蛋白 HDL/(mmol/L) 1.74±0.21 2.08±0.12 1.62±0.21 0.241
低密度脂蛋白 LDL/(mmol/L) 4.42±0.07 3.57±0.51 4.00±0.34 0.288
甘油三酯 TG/(μg/mL) 407.53±14.18 439.82±2.89 432.37±19.63 0.355

2.4 胍基乙酸对育肥猪肌纤维特性的影响

由表6可知,各组之间肌纤维直径无显著差异(P>0.05)。T1组和T2组的肌纤维横截面积显著高于对照组(P<0.05)。
表6 胍基乙酸对育肥猪肌纤维特性的影响

Table 6 Effects of guanidinoacetic acid on muscle fiber characteristics of finishing pigs

项目
Items		 组别 Groups P值
P-value
对照 Control T1 T2
肌纤维直径 Muscle fiber diameter/μm 63.10±1.93 68.72±1.66 66.30±2.30 0.187
肌纤维横截面积
Muscle fiber cross-sectional area/μm2		 4 073.31±85.81b 5 347.05±247.77a 5 173.07±277.52a 0.013

2.5 胍基乙酸对育肥猪肌肉发育相关调节因子表达的影响

由表7可知,T2组的肌肉MyoG基因相对表达量显著高于对照组(P<0.05)。各组之间肌肉MSTN、MyoD基因相对表达量无显著差异(P>0.05)。与对照组相比,T2组的肌肉Myf5基因相对表达量有升高的趋势(P=0.093)。
表7 胍基乙酸对育肥猪肌肉发育相关调节因子基因表达的影响

Table 7 Effects of guanidinoacetic acid on gene expression of regulatory factors related to muscle development of finishing pigs

项目
Items		 组别 Groups P值
P-value
对照 Control T1 T2
肌细胞生成素 MyoG 1.00±0.01b 1.04±0.03ab 1.06±0.02a 0.045
肌肉生长抑制素 MSTN 1.01±0.03 0.95±0.02 1.01±0.02 0.119
生肌因子5 Myf5 1.01±0.03 1.11±0.04 1.16±0.07 0.093
生肌决定因子 MyoD 1.02±0.06 1.11±0.03 1.12±0.04 0.222

3 讨论

3.1 胍基乙酸对育肥猪生长性能的影响

生长性能是衡量养猪经济效益的重要指标,提高生长性能对于提高养猪竞争力尤为关键。Jayaraman等[20]和Li等[21]研究发现,饲粮中添加胍基乙酸显著提高了育肥猪的平均日增重和饲料效率;De Souza等[22]研究发现,饲粮中添加0.06%胍基乙酸显著提高了肉鸡的平均日增重、平均日采食量和饲料效率;Lin等[23]研究发现,胍基乙酸能显著提高牛蛙的末重。本试验结果表明,试验组育肥猪的末重、平均日增重和平均日采食量均显著提高,料重比显著降低,与前人报道结果[21,24-25]一致。这是因为添加胍基乙酸可以减少体内甘氨酸和精氨酸的消耗[26],为动物生长提供了更多的蛋白质;同时,添加胍基乙酸可以提高机体肌酸和磷酸肌酸储备,磷酸肌酸可迅速转化为ATP为动物生长提供能量。此外,本试验结果表明,胍基乙酸基础上补充甜菜碱,料重比显著低于单独添加胍基乙酸组。这可能是因为胍基乙酸在合成肌酸过程中需要蛋氨酸提供甲基,同时会生成同型半胱氨酸,甜菜碱可与同型半胱氨酸在甜菜碱同型半胱氨酸甲基转移酶作用下重新生成蛋氨酸,促进了肌酸的合成[16-17],从而为育肥猪生长提供了更多的ATP。

3.2 胍基乙酸对育肥猪屠宰性能的影响

胴体重、屠宰率、胴体斜长、胴体直长是影响生猪生产的重要经济性状,眼肌面积和背膘厚度是胴体瘦肉率的评价指标。He等[27]研究发现,随着胍基乙酸添加量的增加,育肥猪胴体重和瘦肉百分比有升高趋势。本研究结果表明,试验组育肥猪屠宰率和眼肌面积均显著提高,且胍基乙酸基础上补充甜菜碱组胴体重与对照组相比有升高的趋势,与Jayaraman等[20]报道一致。这可能是因为胍基乙酸为肌肉生长提供了更多的肌酸、磷酸肌酸和ATP[27⇓-29],提高了肌肉产量。另外,有研究发现,生长激素能直接促进脂肪分解[30],减少胴体脂肪沉积,胍基乙酸还能促进生长激素的分泌[5],进而提高育肥猪眼肌面积。综上所述,饲粮中添加胍基乙酸和甜菜碱能提高育肥猪屠宰率,促进肌肉发育。

3.3 胍基乙酸对育肥猪血清生化指标的影响

总蛋白、白蛋白、尿素含量可反映机体蛋白质代谢和氨基酸平衡。Zhang等[31]研究发现,在羔羊饲粮中添加胍基乙酸后,血清总蛋白含量显著升高,血清尿素含量显著降低。Ren等[24]报道,羔羊饲粮中添加胍基乙酸和甜菜碱能显著提高血清总蛋白含量。本研究结果表明,试验组育肥猪第38天血清中总蛋白含量显著升高,尿素含量显著降低。这可能是因为胍基乙酸和甜菜碱能够提高胰岛素和胰岛素样生长因子-1(IGF-1)含量[32⇓-34],激活了哺乳动物雷帕霉素靶蛋白(mTOR)信号通路,促进了蛋白质的合成[35],同时胰岛素还能通过减少蛋白质的分解来提高机体氮沉积。尿素含量降低的原因可能是胍基乙酸增加了微生物蛋白合成[24],促进了非蛋白氮的利用。总胆固醇、高密度脂蛋白、低密度脂蛋白是衡量血脂代谢的重要指标,低密度脂蛋白含量的升高会导致脂肪合成增多。本研究结果表明,试验组育肥猪第19天血清低密度脂蛋白含量有降低的趋势,与张翼[36]等研究结果一致。这可能是因为胍基乙酸和甜菜碱通过影响脂肪酸合成酶(FAS)和肉碱棕榈酰转移酶-1(CPT-1)的基因表达[37],减少了脂肪的合成。

3.4 胍基乙酸对育肥猪肌纤维特性的影响

肌纤维的发育与肌纤维数量和肌纤维横截面积有关,猪的肌纤维数量在出生后不会发生改变,但会增粗和变长[38]。Lu等[39]报道,饲粮中添加胍基乙酸可显著提高育肥猪肌纤维横截面积。本试验研究结果表明,试验组肌纤维横截面积均显著高于对照组,与Li等[40]研究结果一致。这可能是因为补充胍基乙酸后,肌肉MyoG、Myf5等生肌因子的基因相对表达量提高[41],同时添加胍基乙酸能提高机体胰岛素样生长因子-1的含量,从而促进肌肉内蛋白的沉积,进而促进了肌肉的生长和肌纤维的肥大。

3.5 胍基乙酸对育肥猪肌肉发育相关调节因子基因表达的影响

骨骼肌的发育包括以下4个过程:中胚层干细胞分化产生成肌细胞[42]、成肌细胞分化融合产生肌管、肌纤维形成、肌纤维成熟[43]。肌细胞的增殖与分化是肌纤维发育过程中必不可少的过程,它决定了肌细胞的命运和肌肉的最终结构。骨骼肌的分化是一个复杂的过程[44],由肌源性调节因子(myogenic regulatory factors,MRFs)调控[45]。MRFs家族包括Myf5、生肌因子4(Mrf4)、MyoD和MyoG[46]。MyoD和Myf5基因主要在成肌细胞的形成过程中发挥作用。MyoG基因是肌肉发育的重要调节因子,主要负责驱动终末分化,不仅能促进成肌细胞的分化和融合[47],而且能有效激活该过程所需的靶基因[48-49]。有研究发现,MyoG基因相对表达量的下调能抑制成肌细胞的融合和肌肉细胞分化[50⇓-52];Davie等[48]报道,骨骼肌缺乏MyoG基因会导致小鼠在出生时死亡,这表明MyoG基因对于肌肉生长发育具有重要调节作用。Ibrahim等[53]研究发现,饲粮中添加胍基乙酸能显著提高鸭子肌肉中MyoG基因相对表达量。本试验研究发现,胍基乙酸基础上补充甜菜碱组肌肉MyoG基因相对表达量显著升高,并且Myf5基因相对表达量有升高的趋势。Lu等[39]报道,胍基乙酸显著提高了育肥猪背最长肌中MyoD、Myf5等基因相对表达量。MSTN是一种具有重要生理功能的负调节因子,也称为生长分化因子-8,主要作用是抑制肌细胞生长和分化[54-55]。Farshidfar等[56]研究发现,胍基乙酸及其代谢物肌酸能通过下调MSTN基因相对表达量来消除其对肌肉生长的抑制作用。Lu等[39]研究发现,胍基乙酸能显著降低育肥猪背最长肌中MSTN基因相对表达量。但本试验发现,胍基乙酸对MSTN基因相对表达量并无显著影响,造成不同结果的原因有待进一步研究。综上所述,饲粮中添加胍基乙酸和甜菜碱能提高背最长肌MyoG基因相对表达量,进而促进肌纤维的发育。

4 结论

饲粮中添加胍基乙酸和甜菜碱能提高育肥猪生长性能、屠宰性能和肌纤维横截面积,促进肌纤维的增殖与分化,且联合添加胍基乙酸与甜菜碱的促生长效果优于单独添加胍基乙酸。

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