Chinese Journal of Animal Nutrition >

2023 , Vol. 35 >Issue 9: 5736 - 5745

DOI: https://doi.org/10.12418/CJAN2023.528

ORESEARCH PAPER

Effects of Choline Supplementation on Growth Performance, Slaughter Performance, Serum Biochemical Indices and Liver Lipid Metabolism of Magang Geese during 28 to 70 Days of Age

  • DENG Yuyang , 1 ,
  • WHNG Heng 1 ,
  • ZHAN Yuchun 1 ,
  • LI Ye 1 ,
  • XIE Shisheng 1 ,
  • LI Sheng 2 ,
  • WANG Wence 1 ,
  • ZHU Yongwen , 1, * ,
  • YANG Lin , 1, *
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  • 1 Guangdong Province Key Laboratory of Animal Nutrition Regulation, College of Animal Science, South China Agricultural University, Guangzhou 510642, China
  • 2 Jinqian Feed (Dongguan) Co., Ltd., Dongguan 523147, China
* ZHU Yongwen, associate professor, E-mail: ;
YANG Lin, professor, E-mail:

Received date: 2023-03-28

  Online published: 2023-09-13

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Abstract

The objective of this experiment was to investigate the effects of choline supplementation on the growth performance, slaughter performance, serum biochemical indices and liver lipid metabolism of Magang geese during 28 to 70 days of age. A total of 432 healthy 28-day-old Magang gander geese of similar body weight were randomly divided into six groups, each with six replicates and twelve geese per replicate. The geese in the control group were fed a basal diet (corn-corn starch-corn gluten meal type semi-purified diet, choline content was 385 mg/kg). Others in the five experimental groups received a basal diet supplemented with 300, 600, 900, 1 200 and 1 500 mg/kg choline (choline was added to the basal diet by replacing the bentonite carrier in the premix in equal amount), respectively. The experiment lasted for 42 days (from 28 to 70 days of age). The results showed as follows: 1) dietary choline supplemental level had no significant effects on the average final weight, average daily gain and average daily feed intake of geese aged 28 to 70 days (P>0.05). But the feed/gain ratio in the 300 mg/kg choline supplementation group was significantly lower than that in the 1 200 and 1 500 mg/kg choline supplementation groups (P<0.05). 2) Dietary choline supplemental level had no significant effects on the eviscerated percentage, semi-eviscerated percentage and dressing percentage of 70-day-old Magang geese (P>0.05), and the breast muscle percentage in 300 mg/kg choline supplementation group was significantly higher than that in the control group and 1 500 mg/kg choline supplementation group (P<0.05). 3) Dietary choline supplemental level had no significant effects on the serum glucose (GLU) content and alanine transaminase (ALT) activity of 70-day-old Magang geese (P>0.05), while the serum aspartate aminotransferase (AST) activity was significantly lower in the 300 and 1 200 mg/kg choline supplementation groups than that in the control group (P<0.05). Moreover, the 900 mg/kg choline supplementation group had significantly lower serum and liver triglyceride (TG) content than that in the control group (P<0.05); the 600 mg/kg choline supplementation group had significantly higher serum high density lipoprotein cholesterol (HDL-C) content than that in the other groups (P<0.05). The HDL-C content in the 600 mg/kg choline group was significantly lower than that in the control group and 300 mg/kg choline supplementation group (P<0.05), while the liver low density lipoprotein cholesterol (LDL-C) content was significantly higher than that in the control group and 300 mg/kg choline supplementation group (P<0.05). 4) The mRNA relative expression level of hepatic acetyl coenzyme A carboxylase (ACC) in the 900 mg/kg choline supplementation group was significantly lower than that in the control group (P<0.05), the mRNA relative expression level of hepatic carnitine palmitoyltransferase 1-A (CPT1-A) was significantly higher than that in the remaining groups (P<0.05), and the mRNA relative expression level of peroxisome proliferator-activated receptor α (PPARα) was significantly higher than in the control group, 300 and 1 500 mg/kg choline supplementation groups (P<0.05). In summary, from 28 to 70 days of age, adding 300 mg/kg choline to a corn-corn starch-corn gluten meal type semi-purified diet (choline content is 385 mg/kg) can reduce the feed/gain ratio and increase the breast muscle percentage of Magang geese; a supplemental level of 900 mg/kg choline can cause the up-regulation of PPARα and CPT1-A mRNA expression levels and decrease the content of serum and liver TG of 70-day-old Magang geese. Take performance into account, it is recommended that the choline supplemental level in the diet of Magang goose during 28 to 70 days of age is 300 mg/kg, that is the actual content of choline in the diet is 685 mg/kg.

Key words： Magang geese; choline; growth performance; slaughter performance; serum biochemical indices; lipid metabolism

Cite this article

DENG Yuyang , WHNG Heng , ZHAN Yuchun , LI Ye , XIE Shisheng , LI Sheng , WANG Wence , ZHU Yongwen , YANG Lin . Effects of Choline Supplementation on Growth Performance, Slaughter Performance, Serum Biochemical Indices and Liver Lipid Metabolism of Magang Geese during 28 to 70 Days of Age[J]. Chinese Journal of Animal Nutrition, 2023 , 35(9) : 5736 -5745 . DOI: 10.12418/CJAN2023.528

胆碱又名胆素,最早是Strecker在1849年从动物胆汁中分离得到,广泛分布在植物和动物细胞中,以游离态或复合物的形式存在,如磷酸胆碱、甘油磷酸胆碱、鞘磷脂或者是卵磷脂[1]。由于胆碱在体内具有作为神经递质乙酰胆碱的前体、细胞膜磷脂的结构成分以及提供甲基等多种功能,是畜禽以及水产动物生长发育必需的维生素[1]。胆碱缺乏时,家禽主要表现为生长速度和饲料转化率下降、易患脂肪肝等[2-3]。饲粮添加胆碱可提高家禽生产性能和营养物质利用率,降低料重比(F/G)[4]。此外,胆碱在调节脂质代谢方面发挥着重要作用。研究表明,在29~48周龄肉种鸡饲粮中添加760 mg/kg胆碱可显著降低肝脏脂质含量和血清甘油三酯(TG)含量[5]。闻治国[6]通过肉鸭平均日增重(ADG)拟合胆碱需要量,推荐1~21日龄北京鸭的饲粮胆碱含量为1 300 mg/kg,而21~42日龄时则是1 150 mg/kg。现今关于胆碱在家禽方面的研究主要集中在鸡、鸭上,而对于肉鹅胆碱需要量的研究相对缺乏。张文旭等[7]综合生长性能、屠宰性能和养分利用率,推荐1~4周龄青农灰鹅饲粮胆碱添加量为1 800 mg/kg。NRC(2004)推荐4~8周龄肉鹅的胆碱需要量为1 000 mg/kg。然而,该数据是借鉴肉鸡的数据得出的,且年代较为久远。因此。本试验以马冈鹅为研究对象,研究不同胆碱添加水平对28~70马冈鹅生长性能、屠宰性能、血清生化与脂质代谢指标及肝脏脂质代谢的影响,以期确定马冈鹅生长中后期饲粮中胆碱的适宜添加水平,为完善我国肉鹅营养需要标准及合理配制马冈鹅饲粮提供参考。

1 材料与方法

1.1 试验设计及饲粮配制

选用432只同批出雏、体重相近且健康的28日龄马冈鹅公鹅,采用单因素随机分组试验设计分为6组,以12只试验鹅为1个重复,每组设有6个重复。对照组饲喂不添加胆碱的基础饲粮(胆碱含量为385 mg/kg,离子色谱法测得),其余5个试验组在基础饲粮中分别添加300、600、900、1 200、1 500 mg/kg的胆碱,胆碱是通过等量取代预混料中的膨润土载体加入到基础饲粮中。试验所用基础饲粮以NRC(1994)[8]中的家禽营养需要推荐量为参考,并考虑现今我国马冈鹅生产实际进行配制,为玉米-玉米淀粉-玉米蛋白粉型半纯合饲粮,其组成及营养水平见表1。饲养试验在揭阳市惠来县华南农大·金钱饲料科研合作基地中开展。试验鹅采用笼养方式,自由采食和饮水。试验期为43 d(28~70日龄)。
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 40.00
玉米淀粉 Corn starch 22.43
玉米蛋白粉Corn gluten meal (CP 60%) 14.00
芝麻粕 Sesame meal 3.00
稻壳粉 Rice husk powder 7.00
小麦麸 Wheat bran 4.00
大豆油 Soybean oil 3.00
石粉 Limestone 0.95
磷酸氢钙 CaHPO4 1.90
食盐 NaCl 0.45
L-赖氨酸盐酸盐 L-Lys·HCl 0.60
DL-蛋氨酸 DL-Met 0.06
L-精氨酸盐酸盐 L-Arg·HCl 0.40
L-色氨酸 L-Trp 0.08
苏氨酸 Thr 0.13
磷酸二氢钾 KH2PO4 0.10
氯化钾 KCl 0.30
七水合硫酸镁 MgSO4·7H2O 0.60
预混料 Premix1) 1.00
合计 Total 100.00
营养水平 Nutrient levels2)
代谢能 ME/(MJ/kg) 12.30
粗蛋白质 CP 14.78
粗纤维 CF 4.52
钙 Ca 0.99
总磷 TP 0.70
非植酸磷 AP 0.51
赖氨酸 Lys 0.77
蛋氨酸 Met 0.38
苏氨酸 Thr 0.60
蛋氨酸+半胱氨酸 Met+Cys 0.80
胆碱 Choline/(mg/kg) 385.00

1)预混料为每千克饲粮提供 The premix provided the following per kg of the diet: VA 9 000 IU,VD3 1 500 IU,VE 7.5 IU,VK 4 mg,VB1 0.6 mg,VB2 4.8 mg,生物素 biotin 1.5 mg,VB12 0.9 mg,泛酸钙 calcium pantothenate 7.5 mg,叶酸 folic acid 0.15 mg,烟酸 nicotinic acid 20 mg,Cu (as copper sulfate) 8 mg,Fe (as ferrous sulfate) 80 mg,Zn (as zinc sulfate) 90 mg,Mn (as manganese sulfate) 70 mg,Se (as sodium selenite) 0.3 mg,I (as potassium iodide) 0.4 mg,Mo (as sodium molybdate) 8.3 mg,Co (as cobalt chloride) 1.7 mg。

2)粗蛋白质和胆碱为实测值,其余为计算值。CP and choline were measured values, while the others were calculated values.

1.2 样品采集与制备

70日龄试验结束时,每个重复选取1只试验鹅,颈静脉采血10 mL,静置30 min后,3 000 r/min离心10 min分离血清,-20 ℃保存,用于血清生化和脂质代谢指标的测定。将采血后的试验鹅屠宰,分离出部分肝脏,放入冻存管并迅速置于液氮,样品保存在-80 ℃冰箱,用于后续肝脏相关指标的测定。剪取 0.1 g肝脏组织,在4 ℃生理盐水中漂洗后用滤纸吸干,称重后加入9倍重量体积的冰生理盐水,利用匀浆机研磨充分,3 000 r/min离心10 min,分离上清液并分装,置于-80 ℃冰箱保存,用于部分肝脏脂质代谢指标的测定。

1.3 测定指标及方法

1.3.1 生长性能

在28日龄时正式开始试验,试验期间以重复为单位称量、记录各组鹅的耗料重,若有死鹅、病鹅,及时淘汰并记录,在70日龄试验结束时对每一栏舍试验鹅称重,之后计算ADG、平均日采食量(ADFI)和F/G。

1.3.2 屠宰性能

在70日龄试验结束时,从每个重复选取1只试验鹅,颈静脉放血致死,湿法拔毛沥水后称重,测定其屠宰性能,包括屠宰率、半净膛率、全净膛率、胸肌率、腿肌率和腹脂率。试验操作及指标的测定方法参照《家禽生产性能名词术语和度量统计方法》(NY/T 823—2020)进行。

1.3.3 血清生化与脂质代谢指标以及肝脏脂质代谢指标

参考南京建成生物工程研究所试剂盒操作说明书检测血清、肝脏的如下全部或部分指标:葡萄糖(GLU)、甘油三酯(TG)、总胆固醇(T-CHO)、高密度脂蛋白胆固醇(HDL-C)、低密度脂蛋白胆固醇(LDL-C)、游离脂肪酸(FFA)含量及谷丙转氨酶(ALT)、谷草转氨酶(AST)活性。
肝脏总脂质含量:肝脏总脂质含量的测定参照Folch等[9]的方法,并略有改动。称取肝脏样品1 g左右,加入体积比为2:1的氯仿-甲醇溶液和50 μL抗氧化剂丁基羟基甲苯,在冰浴下匀浆,静置过滤,滤纸用氯仿-甲醇溶液充分冲洗,然后在滤液中加入10 mL 1.6%的氯化钙(CaCl2),摇匀静置1 h;离心后吸走上清液,将下层液体转入已经称重的容器中,在液氮下干燥至恒重,称重。
肝脏总磷脂含量:将上述脂质重新溶于10 mL氯仿-甲醇溶液中,取4 mL液体,参考Rizki等[10]的操作,采用抗坏血酸-钼蓝比色法测定肝脏总磷脂含量。

1.3.4 肝脏脂质代谢相关基因的表达

依据Trizol法,采用EZB试剂盒从肝脏样品中提取总RNA,然后使用核酸浓度分析仪(ND-2000,Gene Company,美国)测定RNA的浓度,并检测其在260和280 nm处的吸光度比值(OD260 nm/OD280 nm)。按照TaKaRa PrimeScriptTM RT Reagent Kit with gDNA Eraser Kit提供的方法进行逆转录,合成cDNA。
以β-肌动蛋白(β-actin)作为内参基因,引物由上海生工生物技术有限公司合成,引物序列见表2。使用Quantifast ST-BR Green PCR Kit,采用SYBR Green Ⅰ染料法,在Bio-Rad CFX-96型荧光定量PCR仪上进行实时荧光定量PCR。每个实时荧光定量PCR设2个重复。PCR反应体系为10 μL:SYBR Green Master Mix(2×)5 μL,上、下游引物各0.2 μL,cDNA模板1 μL,ddH2O 3.6 μL。PCR反应程序为:95 ℃预变性10 min,95 ℃变性10 s,56 ℃退火延伸30 s,总计40个循环。反应结束后采用2-ΔΔCt法计算目的基因乙酰辅酶A羧化酶(ACC)、过氧化物酶增殖物激活受体α(PPARα)和肉碱棕榈酰转移酶1-A(CPT1-A)mRNA相对表达量。
表2 引物序列

Table 2 Primer sequences

基因
Genes		 GenBank登录号
GenBank accession No.		 引物序列
Primer sequences (5'—3')
β-肌动蛋白
β-actin		 XM_013174886.1 F:GCCCTCTTCCAGCCATCTTTCTTG
R:CCACCAGACAGCACCGTGTTG
乙酰辅酶A羧化酶
ACC		 XM_040532346 F:CACAGATCCAGAGCACAGCACTTC
R:GGCAGGCAGTATCCGTTCATCAC
过氧化物酶增殖物激活受体α
PPARα		 XM_048046893 F:CCTTTCACCAGCATCCAGTCCTTC
R:TGTACTCCGTAATGGTAGCCTGAGG
肉碱棕榈酰转移酶1-A
CPT1-A		 XM_048058717 F:CCGCCATCTGTTCTGCCTCTATG
R:GCCAGCATCTCAGGGTTCTTCTTC

1.4 数据统计及分析

数据使用Excel 2013进行初步处理,后续使用SAS 9.2软件对试验数据进行单因素方差分析,差异显著时采用Duncan氏法进行组间多重比较,基因表达所得数据使用Graph Prism 8.0进行柱形图的绘制。试验结果以平均值和均值标准误(SEM)表示,以P<0.05表示差异显著。

2 结果

2.1 胆碱添加水平对28~70日龄马冈鹅生长性能的影响

由表3可知,胆碱添加水平对28~70日龄马冈鹅的平均末重、ADG和ADFI均没有显著影响(P>0.05);300 mg/kg添加组的F/G显著低于1 200和1 500 mg/kg添加组(P<0.05)。
表3 胆碱添加水平对28~70日龄马冈鹅生长性能的影响

Table 3 Effects of choline supplemental level on growth performance of Magang geese from 28 to 70 days of age

项目
Items		 胆碱添加水平
Choline supplemental levels/(mg/kg)		 均值
标准误
SEM		 P值
P-value
0 300 600 900 1 200 1 500
平均初重 AIBW/g 1 240 1 287 1 253 1 285 1 268 1 293 22.11 0.52
平均末重 AFBW/g 3 512 3 685 3 515 3 566 3 639 3 612 91.49 0.72
平均日增重 ADG/g 54.08 57.12 56.11 54.28 56.44 55.17 1.76 0.79
平均日采食量 ADFI/g 182.58 206.62 196.54 206.20 211.45 207.21 9.76 0.49
料重比 F/G 3.80abc 3.40c 3.66bc 3.82abc 4.11a 3.87ab 0.14 0.04

同行数据无字母标注或标有相同字母表示差异不显著(P>0.05),肩标不同小写字母表示差异显著(P<0.05)。下表同。

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

2.2 胆碱添加水平对70日龄马冈鹅屠宰性能的影响

由表4可知,胆碱添加水平对70日龄马冈鹅屠宰率、半净膛率、全净膛率、腿肌率、腹脂率均无显著影响(P>0.05);300 mg/kg添加组的胸肌率显著高于对照组和1 200 mg/kg添加组(P<0.05)。
表4 胆碱添加水平对70日龄马冈鹅屠宰性能的影响

Table 4 Effects of choline supplemental level on slaughter performance of Magang geese aged 70 days%

项目
Items		 胆碱添加水平
Choline supplemental levels/(mg/kg)		 均值标准误
SEM		 P值
P-value
0 300 600 900 1 200 1 500
屠宰率 Dressing percentage 89.07 89.20 89.56 90.21 88.22 89.51 0.72 0.53
半净膛率
Semi-eviscerated percentage		 77.63 77.62 80.11 78.55 79.51 77.18 0.83 0.08
全净膛率
Eviscerated percentage		 69.29 68.97 71.51 69.90 70.80 67.78 0.90 0.08
胸肌率
Breast muscle percentage		 8.68c 11.21a 10.06abc 10.34ab 9.14bc 10.66ab 0.51 0.02
腿肌率
Leg muscle percentage		 12.24 12.09 11.96 11.75 11.64 13.38 0.54 0.27
腹脂率
Abdominal fat percentage		 5.62 4.90 4.49 5.46 5.22 4.55 0.33 0.12

2.3 胆碱添加水平对70日龄马冈鹅血清生化与脂质代谢指标的影响

由表5可知,胆碱添加水平对血清GLU含量、ALT活性无显著影响(P>0.05);300、600、900和1 200 mg/kg添加组血清AST活性显著低于对照组(P<0.05)。对于脂质代谢指标,胆碱添加水平对70日龄马冈鹅血清T-CHO含量无显著影响(P>0.05);900 mg/kg添加组血清TG含量显著低于对照组(P<0.05);600 mg/kg添加组血清HDL-C含量显著高于其余各组(P<0.05);300 mg/kg添加组血清FFA含量显著高于对照组(P<0.05)。
表5 胆碱添加水平对70日龄马冈鹅血清生化与脂质代谢指标的影响

Table 5 Effects of choline supplemental level on serum biochemical and lipid metabolism indices of Magang geese aged 70 days

项目
Items		 胆碱添加水平
Choline supplemental levels/(mg/kg)		 均值标准误
SEM		 P值
P-value
0 300 600 900 1 200 1 500
葡萄糖 GLU/(mmol/L) 10.97 9.68 9.60 9.72 13.84 9.75 1.13 0.08
谷草转氨酶 AST/(U/L) 12.84a 3.03b 3.80b 5.75b 5.02b 14.98a 2.42 <0.01
谷丙转氨酶 ALT/(U/L) 5.87 5.56 8.26 6.88 7.80 9.23 1.27 0.31
游离脂肪酸 FFA/(mmol/L) 0.58c 1.22a 0.80abc 0.91abc 1.12ab 0.70bc 0.16 0.04
甘油三酯 TG/(mmol/L) 0.88a 0.76a 0.60ab 0.47b 0.75a 0.83a 0.09 0.03
总胆固醇 T-CHO/(mmol/L) 6.32 5.34 5.42 5.01 5.50 6.10 0.45 0.37
高密度脂蛋白胆固醇
HDL-C/(mmol/L)		 2.16b 2.57b 3.17a 2.60b 2.29b 2.21b 0.18 <0.01
低密度脂蛋白胆固醇
LDL-C/(mmol/L)		 1.74 2.25 2.21 2.14 1.82 2.59 0.20 0.06

2.4 胆碱添加水平对70日龄马冈鹅肝脏脂质代谢指标的影响

由表6可知,胆碱添加水平对70日龄马冈鹅肝脏总磷脂含量无显著影响(P>0.05);1 500 mg/kg添加组肝脏总脂质含量显著低于对照组和1 200 mg/kg添加组(P<0.05);900 mg/kg添加组肝脏TG含量显著低于除1500mg/kg添加组外的其余各组(P<0.05);300、900 mg/kg添加组肝脏T-CHO含量显著低于1 200 mg/kg添加组(P<0.05);600 mg/kg胆碱添加组肝脏HDL-C含量显著低于对照组和300 mg/kg添加组(P<0.05),而LDL-C含量显著高于对照组和300 mg/kg胆碱组(P<0.05)。
表6 胆碱添加水平对70日龄马冈鹅肝脏脂质代谢指标的影响

Table 6 Effects of choline supplemental level on liver lipid metabolism indices of Magang geese aged 70 days

项目
Items		 胆碱添加水平
Choline supplemental levels/(mg/kg)		 均值标准误
SEM		 P值
P-value
0 300 600 900 1 200 1 500
总磷脂 TPL/(mg/g肝脏) 61.81 53.45 52.96 58.15 56.43 57.76 2.78 0.35
总脂质 TL/(mg/g肝脏) 80.57a 55.24ab 64.35ab 67.58ab 70.18a 41.47b 8.79 0.04
甘油三酯 TG/(mmol/g prot) 0.17ab 0.16ab 0.20a 0.10c 0.19ab 0.13bc 0.02 <0.01
总胆固醇
T-CHO/(mmol/g prot)		 0.32ab 0.25b 0.35ab 0.20b 0.51a 0.37ab 0.07 0.04
高密度脂蛋白胆固醇
HDL-C/(mmol/g prot)		 2.97a 2.83a 2.07b 2.56ab 2.38ab 2.68ab 0.20 0.04
低密度脂蛋白胆固醇
LDL-C/(mmol/g prot)		 1.21b 1.12b 2.32a 0.91b 0.92b 1.38b 0.30 0.04

2.5 胆碱添加水平对70日龄马冈鹅肝脏脂质代谢相关基因表达的影响

由图1可知,胆碱添加水平对70日龄马冈鹅肝脏中ACC、CPT1-A和PPARα mRNA相对表达量均有显著影响(P<0.05),其中900 mg/kg添加组肝脏ACC mRNA相对表达量显著低于对照组和1 200 mg/kg添加组(P<0.05),PPARα mRNA相对表达量显著高于对照组和300、1 500 mg/kg添加组(P<0.05),而CPT1-A mRNA相对表达量则显著高于其余各组(P<0.05)。
图1 胆碱添加水平对70日龄马冈鹅肝脏脂质代谢相关基因表达的影响

数据柱标注不同小写字母表示差异显著(P<0.05),相同字母或无字母表示差异不显著(P>0.05)。

Fig.1 Effects of choline supplemental level on gene expression involved in lipid metabolism in liver of Magang geese aged 70 days

Date columns 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).

3 讨论

3.1 饲粮胆碱添加水平对28~70日龄马冈鹅生长性能和屠宰性能的影响

胆碱能够促进机体生长发育,在猪、反刍动物和水产动物上均已得到了证实[11⇓-13]。Workel等[14]的研究表明,饲粮中添加800 mg/kg胆碱显著提高了肉鸡的ADG,降低了F/G。Rahnama等[15]也得出了相似的结果。蒋雪樱等[16]研究表明,在22~42日龄肉鸡饲粮添加500 mg/kg甜菜碱显著提高了爱拔益加(AA)肉鸡的胸肌和腿肌的重量。本试验结果显示,饲粮添加300 mg/kg胆碱显著降低了28~70日龄马冈鹅的F/G,而胆碱添加水平对28~70日龄马冈鹅的ADG、ADFI和平均末重均无显著影响。在屠宰性能方面,本试验发现胆碱添加水平对70日龄马冈鹅的屠宰率、半净膛率和全净膛率均无显著影响。而张文旭等[7]的研究发现,在15周龄青农灰鹅饲粮中添加1 800 mg/kg胆碱能够显著提高肉鹅的屠宰率和全净膛率,与本试验结果不一致,这可能是鹅的品种与生长阶段不同所导致。此外,添加300 mg/kg胆碱显著提高了70日龄马冈鹅的胸肌率,与蒋雪樱等[16]的结果一致,这可能是由于胆碱在体内为蛋氨酸合成提供甲基,并提高了其有效利用率,使马冈鹅生长速度加快,提高了蛋白质的周转沉积[17]。

3.2 饲粮胆碱添加水平对70日龄马冈鹅血清生化与脂质代谢指标以及肝脏脂质代谢的影响

AST、ALT与动物肝脏功能密切相关,其活性可反映肝脏功能是否正常[18]。当肝脏功能衰退时,ALT和AST进入血液中增加,血液中这2种酶的活性也随之升高。因此,肝脏功能正常与否与可以通过测定血液中AST和ALT的活性进行评估。而肝脏出现脂肪性病变时,脂质增加会导致血糖水平升高[19]。本试验中,饲粮添加胆碱可降低血清AST活性,表明胆碱可能对肝脏功能有一定的改善作用。Yusuf等[20]在21~42日龄肉鸡饲粮中添加3 g/kg甜菜碱也得出同样的结果。在北京鸭的研究也表明,胆碱缺乏显著提高了血清中AST和ALT的活性,而对GLU含量无显著影响[6]。TG、T-CHO、LDL-C和HDL-C是体内脂质水平的重要标志,其含量反映了体内脂质的吸收和代谢。HDL-C能促进肝外胆固醇转运到肝脏代谢,维持胆固醇稳定,而LDL-C是TG的主要运输者,禽类脂肪沉积主要取决于血清TG含量。闻治国[6]的研究表明,饲粮胆碱含量对21日龄北京鸭血清T-CHO和HDL-C含量无显著影响,而饲粮胆碱含量为1 414 mg/kg时显著提高了血清FFA和LDL-C含量。Song等[21]研究表明,饲粮添加2 955 mg/kg胆碱显著升高了蛋鸭血清高密度脂蛋白(HDL)、极低密度脂蛋白(VLDL)和TG含量。本试验中,饲粮胆碱添加水平对70日龄马冈鹅血清T-CHO含量无显著影响,600 mg/kg胆碱添加水平显著降低了血清TG含量,300 mg/kg胆碱添加水平显著提高了血清HDL-C和FFA含量,这与前人研究结果大致相同。血清中TG和T-CHO含量下降说明机体脂肪分解代谢增强,这与本试验前文关于腹脂率降低的研究结果一致。
肝脏是脂质代谢的主要场所,在调节脂质代谢和维持机体脂质稳定方面有着重要作用[22]。肝脏总脂质和TG含量随饲粮胆碱添加水平的升高呈现逐渐下降的趋势,这在小鼠[10]和肉鸡[23]的研究中己经证实。本试验结果表明,添加900 mg/kg胆碱显著降低了70日龄马冈鹅肝脏中TG含量,而添加600 mg/kg胆碱显著降低了肝脏HDL-C含量和升高了肝脏LDL-C含量。这可能是因为胆碱磷酸化促进了肝脏中VLDL和HDL的合成,VLDL从脂肪组织和脂蛋白中动员TG以降低血清TG含量。TG的降解增加了血清FFA的含量。此外,HDL与胆固醇结合形成HDL-C,降低T-CHO含量,从而减少或重新分配体内脂肪。
除FFA和TG摄取外,肝脏脂质代谢还涉及脂肪酸的从头合成和氧化。ACC是催化脂肪酸合成反应的限速酶,调控脂肪合成[22],CPT1-A和PPARα在脂肪酸β氧化中起关键作用,前者是该过程的关键酶,后者则是促进脂肪酸β氧化的靶基因[23]。本试验结果表明,对照组肝脏中ACC mRNA相对表达量显著高于1 200 mg/kg添加组,与El Sayed等[24]和姚宏等[25]分别在肉鸭和肉鸡上报道的甜菜碱能降低肝脏ACC基因表达的结果一致。由此可知,胆碱通过下调ACC的表达抑制机体脂肪合成。随着饲粮中胆碱添加水平的增加,肝脏中PPARα mRNA相对表达量呈现先增加后下降的趋势,且900 mg/kg添加组肝脏中PPARα mRNA相对表达量显著高于对照组和300、1 500 mg/kg添加组。而900 mg/kg添加组肝脏中CPT1-A mRNA相对表达量显著高于各组。张彩霞[26]报道,饲粮中添加0.06%胆碱显著提高了獭兔肝脏中CPT1-A mRNA相对表达量。PPARα和CPT1-A的表达上调可促进脂肪酸氧化反应,降低肝脏TG和胆固醇含量,调节脂质代谢。研究表明,胆碱代谢产生甲基供体可通过降低PPARα启动子甲基化上调PPARα及其靶基因CPT1-A表达,降低肝脏TG累积[27]。

4 结论

本试验条件下,在玉米-玉米淀粉-玉米蛋白粉型半纯合饲粮(胆碱含量385 mg/kg)中添加300 mg/kg胆碱可降低28~70日龄的马冈鹅F/G,提高70日龄马冈鹅的胸肌率;添加900 mg/kg胆碱可降低70日龄马冈鹅血清和肝脏中TG含量,提高肝脏中PPARα和CPT1-A mRNA相对表达量。若考虑生产性能,建议28~70日龄马冈鹅饲粮中胆碱添加量为300 mg/kg(实际含量为685 mg/kg)。

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