Chinese Journal of Animal Nutrition >

2023 , Vol. 35 >Issue 4: 2336 - 2348

DOI: https://doi.org/10.12418/CJAN2023.220

RESEARCH PAPER

Effects of Threonine and Isoleucine on Slaughter Performance, Organ Indexes and Intestinal Morphological Structure, Digestive Enzyme Activities, Antioxidant and Immune Indexes of Wanxi White Geese

  • DING Yuancui ,
  • FANG Suting ,
  • XU Lianzhu ,
  • ZHANG Fengyan ,
  • HU Qianqian ,
  • LI Shenghe , * ,
  • REN Man , *
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  • Anhui Key Laboratory of Animal Nutritional Regulation and Health, College of Animal Science, Anhui Science and Technology University, Fengyang 233100, China
*LI Shenghe, professor, E-mail: ;
REN Man, associate professor, E-mail:

Received date: 2022-10-26

  Online published: 2023-04-12

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Abstract

The purpose of this experiment was to study the effects of threonine and isoleucine on slaughter performance, organ indexes and intestinal morphological structure, digestive enzymes activities, antioxidant and immune indexes of Wanxi white geese. A total of 180 thirty-day-old Wanxi white geese with similar body weight were randomly divided into 3 groups with 3 replicates per group and 20 geese per replicate. The control group was fed a basal diet, the threonine group was fed the basal diet supplemented with 2.0 g/kg L-threonine, and the isoleucine group was fed the basal diet supplemented with 2.5 g/kg L-isoleucine. The experiment lasted for 60 days. The results showed as follows: 1) there were no significant differences in slaughter performance and organ indexes at 90 days of age among all groups (P>0.05). 2) The villus height and crypt depth in duodenum and jejunum at 60 days of age of isoleucine group were significantly higher than those of the control group (P<0.05 or P<0.01), and the crypt depth in duodenum, jejunum and ileum at 60 days of age and villus height in duodenum and ileum at 90 days of age of threonine group were significantly higher than those of the control group (P<0.05 or P<0.01). 3) The duodenal trypsin activity at 60 and 90 days of age of threonine group and isoleucine group was significantly higher than that of the control group (P<0.01), and the duodenal maltase activity at 60 and 90 days of age and activities of amylase and lactase in jejunum at 90 days of age of threonine group were significantly higher than those of the control group (P<0.05 or P<0.01). 4) The jejunal total antioxidant capacity at 60 days of age of isoleucine group was significantly higher than that of control group and threonine group (P<0.05). The jejunal malondialdehyde (MDA) content at 60 days of age of threonine group was significantly lower than that of control group and isoleucine group (P<0.01), and the ileal MDA content at 90 days of age of isoleucine group was significantly lower than that of the control group (P<0.05). 5) The contents of immunoglobulin A, interleukin-4 and interferon-γ in jejunum at 90 days of age of isoleucine group were significantly lower than those of control group and threonine group (P<0.05), the contents of immunoglobulin G and immunoglobulin M in jejunum of threonine group were significantly higher than those of control group and isoleucine group (P<0.01), the jejunal tumor necrosis factor-α content of isoleucine group and threonine group was significantly lower than that of the control group (P<0.01). In conclusion, isoleucine and threonine have no significant effect on the slaughter performance and organ growth of Wanxi white geese in the growth period, but isoleucine and threonine improve the intestinal tissue structure and antioxidant function, as well as the intestinal immune function of Wanxi white geese during 30 to 60 days of age and 61 to 90 days of age, respectively.

Key words： Wanxi white geese; threonine; isoleucine; intestinal tissues; immune function

Cite this article

DING Yuancui , FANG Suting , XU Lianzhu , ZHANG Fengyan , HU Qianqian , LI Shenghe , REN Man . Effects of Threonine and Isoleucine on Slaughter Performance, Organ Indexes and Intestinal Morphological Structure, Digestive Enzyme Activities, Antioxidant and Immune Indexes of Wanxi White Geese[J]. Chinese Journal of Animal Nutrition, 2023 , 35(4) : 2336 -2348 . DOI: 10.12418/CJAN2023.220

2021年世界肉鹅出栏量约6.6亿只,商品鹅出栏量约5.7亿只,肉鹅产值达522.6亿元,水禽业总产值达1 892.2亿元[1]。随着粮食安全问题和环境污染压力的凸显,畜禽饲料产业面临新的挑战,也为新技术的开发和利用提供了机遇。由于鹅产业发展起步晚、基础理论缺乏,造成鹅饲料配制技术尚未完全成熟,氨基酸在鹅饲粮中的应用缺乏研究支撑。因此,研究单体氨基酸是肉鹅低蛋白质饲料配制的前提。本课题组前期研究发现,饲粮苏氨酸(threonine,Thr)水平与断奶仔猪肠道功能密切相关,适宜水平的苏氨酸可提高仔猪肠道黏膜屏障功能和改善肠道形态结构,降低断奶仔猪的腹泻[2];低蛋白质饲粮中补充异亮氨酸(isoleucine,Ile)可提高仔猪小肠免疫球蛋白和防御素含量,并抑制大肠杆菌感染及其引发的炎症反应,缓解仔猪腹泻,提高生长性能[3]。越来越多的研究也表明,饲粮中氨基酸水平可调节肠道屏障功能[4]、减少肠黏膜萎缩[5]、刺激局部免疫系统并维持微生物菌群平衡[6]。
苏氨酸是家禽继蛋氨酸和赖氨酸之后的第3限制性氨基酸。苏氨酸参与肠道黏膜中各种蛋白质(尤其是黏蛋白)的合成,对维持肠道黏膜功能具有重要作用,也可提高肠道上皮细胞的更新速率[7]。苏氨酸可以提高肉鸡早期的免疫力、抗氧化能力和肠道健康[8],提高猪生长性能、胴体品质和免疫功能[9-10]。异亮氨酸可以提高肠道的黏膜免疫功能,促进肠道中防御素分泌,对动物的生长、发育和代谢也起着重要作用[11],同时异亮氨酸可能通过增强动物和人类对葡萄糖摄取来增加肠道发育及健康[12]。水禽营养和饲料配制研究起步晚,基础研究缺乏,苏氨酸和异亮氨酸对鹅肠道结构和功能的研究鲜见报道。因此,本研究主要围绕氨基酸对皖西白鹅肠道结构和免疫功能的影响开展研究,为鹅的氨基酸生理功能研究提供理论基础,也为氨基酸在鹅饲料配方中的应用和低蛋白质饲粮的配制提供数据支持。

1 材料与方法

1.1 试验设计与饲养管理

选取30日龄体重相近的皖西白鹅180只随机分为3组,每组3个重复,每个重复20只,公母各占1/2。对照组饲喂基础饲粮,试验组分为苏氨酸组和异亮氨酸组,苏氨酸组在基础饲粮中添加2.0 g/kg的L-苏氨酸,异亮氨酸组在基础饲粮中添加2.5 g/kg的L-异亮氨酸。试验期60 d,分2个阶段,分别为30~60日龄和61~90日龄。基础饲粮以玉米、豆粕、稻谷、麦麸和玉米蛋白粉为主要成分,参照皖西白鹅生长阶段的营养需要量进行配制,营养水平参考《中国饲料成分及营养价值表(2019年第30版)》[13],试验阶段基础饲粮中苏氨酸和异亮氨酸需要量分别为0.56%、0.55%,基础饲粮组成及营养水平见表1。皖西白鹅饲养试验在安徽展羽生态农业开发有限公司试验场进行。试验前对鹅舍和器具进行消毒,试验鹅自由采食、饮水,严格按照鹅厂免疫程序进行免疫接种。
表1 基础饲粮组成及营养水平(风干基础)

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

项目
Items		 30~60日龄
30 to 60
days of age		 61~90日龄
61 to 90
days of age
原料Ingredients
玉米Corn 55.84 41.55
去皮豆粕
Dehulled soybean meal		 21.00 19.00
稻谷Paddy 12.00
麦麸Wheat bran 10.00 16.00
玉米蛋白粉Corn gluten meal 5.00
石粉Limestone 1.05 1.30
磷酸氢钙CaHPO4 0.90 0.90
食盐NaCl 0.40 0.40
矿物质预混料
Mineral premix1)		 0.05 0.05
维生素预混料
Vitamin premix2)		 0.02 0.02
稻壳Rice hull 5.00 8.00
L-赖氨酸盐酸盐L-Lys·HCl 0.05
L-苏氨酸L-Thr 0.12 0.20
DL-蛋氨酸DL-Met 0.06 0.07
氯化胆碱Choline chloride 0.50 0.50
碘化钾KI 0.01 0.01
合计Total 100.00 100.00
营养水平Nutrient levels3)
粗蛋白质CP 17.15 15.23
粗脂肪EE 2.98 9.05
代谢能ME/(MJ/kg) 11.70 10.51
粗纤维CF 4.34 6.71
项目
Items		 30~60日龄
30 to 60
days of age		 61~90日龄
61 to 90
days of age
钙Ca 0.80 0.80
总磷TP 0.67 0.65
有效磷AP 0.43 0.36
赖氨酸Lys 0.93 0.77
蛋氨酸Met 0.40 0.33
苏氨酸Thr 0.80 0.77
异亮氨酸Ile 0.70 0.58

1)矿物质预混料为每千克饲粮提供 The mineral premix provided the following per kg of diets: Zn 90 mg,Mn 80 mg,Fe 80 mg,Cu 20 mg,I 0.35 mg。

2)维生素预混料为每千克饲粮提供 The vitamin premix provided the following per kg of diets: VA 9 000 IU,VD3 33 000 IU,VE 24 IU,VB6 3 mg,VK3 1.8 mg,VB2 5 mg,VB12 0.1 mg,烟酸 nicotinic acid 40 mg,生物素 biotin 0.05 mg,D-泛酸 D-pantothenic acid 15 mg。

3)氨基酸为实测值,其余为计算值。Amino acids were measured values, while the others were calculated values.

1.2 指标测定

1.2.1 屠宰性能

90日龄时,每个重复随机选取2只鹅,禁食12 h后颈静脉放血致死,湿法拔毛沥干水分后称重,测定其屠宰性能,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)。

1.2.2 器官指数

60和90日龄时,每个重复随机选取2只鹅进行屠宰,解剖取出腺胃和肌胃(去除内容物)、肝脏(无胆囊)、胰腺、脾脏、法氏囊称重,按照以下公式计算器官指数:
器官指数(%)=100×器官鲜重(g)/宰前活重(g)。

1.2.3 肠道形态结构

试验结束时,取十二指肠、空肠、回肠组织,置于4%多聚甲醛固定液中,进行苏木精-伊红(HE)染色。使用正置荧光显微镜(BX63,日本Olympus公司)对切片进行拍照,每张切片选择5个视野,使用Image-Pro Plus 6.0软件(Media,美国Cybernetics公司)测量绒毛高度、隐窝深度,计算绒毛高度/隐窝深度(V/C)。

1.2.4 肠道消化酶活性和抗氧化指标

试验结束时,收集十二指肠、空肠、回肠黏膜,-80 ℃保存,采用试剂盒测定肠道黏膜淀粉酶(amylase)、麦芽糖酶(maltase)、乳糖酶(lactase)、胰蛋白酶(trypsin)、脂肪酶(lipase)、超氧化物歧化酶(SOD)、谷胱甘肽过氧化物酶(GSH-Px)活性和总抗氧化能力(T-AOC)以及丙二醛(MDA)含量,所用试剂盒均购自南京建成生物工程研究所。

1.2.5 血清和空肠免疫指标

90日龄时,每个重复随机选取2只鹅,翅静脉采血后4 000 r/min离心10min,取上清;各重复取相同部位的空肠组织制备组织液。采用酶联免疫吸附测定(ELISA)法和酶标仪(Multiskan,美国Thermo Fisher Scientific公司)检测血清和空肠免疫球蛋白A(IgA)、免疫球蛋白G(IgG)、免疫球蛋白M(IgM)、白细胞介素-2(IL-2)、白细胞介素-4(IL-4)、肿瘤坏死因子-α(TNF-α)、干扰素-γ(IFN-γ)含量,所用试剂盒购自上海鑫乐生物科技有限公司。

1.3 数据处理与分析

采用SPSS 25.0统计软件对数据进行单因素方差分析(one-way ANOVA),若差异显著,则采用Duncan氏法进行多重比较,以P<0.05为差异显著,P<0.01为差异极显著,结果采用平均值±标准误(mean±SE)表示。

2 结果

2.1 苏氨酸和异亮氨酸对90日龄皖西白鹅屠宰性能和器官指数的影响

由表2可知,对照组、苏氨酸组和异亮氨酸组之间90日龄皖西白鹅屠宰率、全净膛率、半净膛率、胸肌率和腿肌率均无显著差异(P>0.05)。
表2 苏氨酸和异亮氨酸对90日龄皖西白鹅屠宰性能的影响

Table 2 Effects of threonine and isoleucine on slaughter performance of Wanxi white geese at 90 days of age %

项目
Items		 对照组
CON group		 苏氨酸组
Thr group		 异亮氨酸组
Ile group		 P值
P-value
屠宰率Dressing percentage 77.08±1.07 77.16±0.95 76.95±0.36 0.87
全净膛率Eviscerated percentage 59.10±0.98 58.94±0.63 59.38±0.58 0.70
半净膛率Semi-eviscerated percentage 71.51±0.89 71.45±0.69 71.59±0.42 0.89
胸肌率Breast muscle percentage 6.51±0.45 6.14±0.18 6.64±0.35 0.36
腿肌率Leg muscle percentage 8.01±0.13 7.65±0.09 7.69±0.24 0.17

同行数据肩标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),相同小写字母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), and with different capital letter superscripts mean significant difference (P<0.01), while with the same small letter or no letter superscripts mean no significant difference (P>0.05). The same as below.

由表3可知,对照组、苏氨酸组和异亮氨酸组之间60、90日龄皖西白鹅肌胃指数、腺胃指数、肝脏指数、胰腺指数、脾脏指数和法氏囊指数均无显著差异(P>0.05)。
表3 苏氨酸和异亮氨酸对60、90日龄皖西白鹅器官指数的影响

Table 3 Effects of threonine and isoleucine on organ indexes of Wanxi white geese at 60 and 90 days of age %

项目
Items		 对照组
CON group		 苏氨酸组
Thr group		 异亮氨酸组
Ile group		 P值
P-value
60日龄60 days of age
肌胃指数Gizzard index 3.66±0.08 4.05±0.30 3.72±0.38 0.39
腺胃指数Gland stomach index 0.32±0.01 0.36±0.03 0.33±0.01 0.23
肝脏指数Liver index 1.69±0.18 2.02±0.17 1.76±0.20 0.27
胰腺指数Pancreas index 0.21±0.01 0.25±0.01 0.23±0.02 0.20
脾脏指数Spleen index 0.06±0.00 0.07±0.00 0.07±0.00 0.18
法氏囊指数Bursa of Fabricius index 0.04±0.00 0.04±0.00 0.04±0.01 0.67
90日龄90 days of age
肌胃指数Gizzard index 3.47±0.11 3.66±0.13 3.47±0.08 0.26
腺胃指数Gland stomach index 0.22±0.01 0.27±0.02 0.24±0.01 0.06
肝脏指数Liver index 1.53±0.09 1.60±0.06 1.57±0.16 0.70
胰腺指数Pancreas index 0.18±0.00 0.19±0.00 0.18±0.00 0.48
脾脏指数Spleen index 0.09±0.01 0.09±0.01 0.08±0.00 0.80
法氏囊指数Bursa of Fabricius index 0.05±0.01 0.05±0.00 0.04±0.00 0.39

2.3 苏氨酸和异亮氨酸对60、90日龄皖西白鹅肠道形态结构的影响

由表4、表5可知,异亮氨酸组60日龄十二指肠绒毛高度显著高于对照组和苏氨酸组(P<0.05),V/C极显著高于对照组和苏氨酸组(P<0.01)。苏氨酸组60日龄十二指肠隐窝深度极显著高于对照组和异亮氨酸组(P<0.01)。异亮氨酸组90日龄十二指肠绒毛高度极显著高于对照组(P<0.01)。苏氨酸组和异亮氨酸组90日龄十二指肠隐窝深度与对照组无显著差异(P>0.05),但苏氨酸组极显著高于异亮氨酸组(P<0.01)。苏氨酸组90日龄十二指肠V/C极显著低于对照组和异亮氨酸组(P<0.01)。
表4 苏氨酸和异亮氨酸对60日龄皖西白鹅肠道形态结构的影响

Table 4 Effects of threonine and isoleucine on intestinal morphological structure of Wanxi white geese at 60 days of age

项目
Items		 对照组
CON group		 苏氨酸组
Thr group		 异亮氨酸组
Ile group		 P值
P-value
十二指肠Duodenum
绒毛高度Villus height/μm 880.88±23.84b 864.35±20.26b 926.55±13.80a 0.03
隐窝深度Crypt depth/μm 297.62±9.07C 425.14±6.69A 386.18±11.28B <0.01
绒毛高度/隐窝深度V/C 2.43±0.08B 2.18±0.04C 2.71±0.06A <0.01
空肠Jejunum
绒毛高度Villus height/μm 882.26±11.84Bb 919.04±17.11ABab 966.32±26.39Aa <0.01
隐窝深度Crypt depth/μm 343.15±6.26Bb 441.87±10.65Aa 433.34±8.20Aa <0.01
绒毛高度/隐窝深度V/C 2.62±0.06Aa 2.16±0.08Bb 2.26±0.07Bb <0.01
回肠Ileum
绒毛高度Villus height/μm 912.86±19.98B 1 075.9±16.93A 779.10±30.74C <0.01
隐窝深度Crypt depth/μm 338.36±7.36B 371.35±7.43A 280.69±11.10C <0.01
绒毛高度/隐窝深度V/C 2.75±0.07 2.94±0.07 2.88±0.13 0.19
表5 苏氨酸和异亮氨酸对90日龄皖西白鹅肠道形态结构的影响

Table 5 Effects of threonine and isoleucine on intestinal morphological structure of Wanxi white geese at 90 days of age

项目
Items		 对照组
CON group		 苏氨酸组
Thr group		 异亮氨酸组
Ile group		 P值
P-value
十二指肠Duodenum
绒毛高度Villus height/μm 789.21±23.08Bc 867.74±19.48ABb 963.60±32.95Aa <0.01
隐窝深度Crypt depth/μm 369.40±7.21ABab 389.41±6.95Aa 359.66±7.45Bb <0.01
绒毛高度/隐窝深度V/C 2.65±0.10Aa 1.97±0.04Bb 2.68±0.10Aa <0.01
空肠Jejunum
绒毛高度Villus height/μm 928.23±34.12Aa 921.69±29.71Aa 551.81±31.26Bb <0.01
隐窝深度Crypt depth/μm 386.67±7.56Aa 395.05±13.92Aa 265.46±11.77Bb <0.01
绒毛高度/隐窝深度V/C 2.44±0.09Aa 2.52±0.11Aa 2.09±0.08Bb <0.01
回肠Ileum
绒毛高度Villus height/μm 638.25±17.44b 705.83±29.65a 702.33±17.09a 0.03
隐窝深度Crypt depth/μm 350.95±9.25b 341.40±8.16b 386.17±12.93a 0.01
绒毛高度/隐窝深度V/C 1.86±0.05b 2.12±0.10a 1.88±0.06a 0.03
异亮氨酸组60日龄空肠绒毛高度极显著高于对照组(P<0.01)。苏氨酸组和异亮氨酸组60日龄空肠隐窝深度极显著高于对照组(P<0.01),但V/C极显著低于对照组(P<0.01)。对照组和苏氨酸组90日龄空肠绒毛高度、隐窝深度和V/C极显著高于异亮氨酸组(P<0.01)。
苏氨酸组60日龄回肠绒毛高度和隐窝深度极显著高于对照组和异亮氨酸组(P<0.01)。苏氨酸组和异亮氨酸组90日龄回肠绒毛高度和V/C显著高于对照组(P<0.05),但异亮氨酸组隐窝深度显著高于对照组和苏氨酸组(P<0.05)。

2.4 苏氨酸和异亮氨酸对60、90日龄皖西白鹅肠道消化酶活性的影响

由表6、表7可知,各组之间60日龄十二指肠、空肠、回肠淀粉酶活性无显著差异(P>0.05),苏氨酸组和异亮氨酸组90日龄空肠淀粉酶活性均显著高于对照组(P<0.05)。苏氨酸组60日龄十二指肠麦芽糖酶活性显著高于对照组和异亮氨酸组(P<0.05),异亮氨酸组回肠麦芽糖酶活性极显著高于对照组和苏氨酸组(P<0.01);苏氨酸组90日龄十二指肠麦芽糖酶活性显著高于对照组(P<0.05),苏氨酸组回肠麦芽糖酶活性极显著高于对照组和异亮氨酸组(P<0.01)。异亮氨酸组和苏氨酸组60日龄空肠乳糖酶活性与对照组差异不显著(P>0.05),但苏氨酸组显著高于异亮氨酸组(P<0.05);苏氨酸组90日龄空肠乳糖酶活性极显著高于对照组和异亮氨酸组(P<0.01)。苏氨酸组和异亮氨酸组60、90日龄十二指肠胰蛋白酶活性均极显著高于对照组(P<0.01),空肠和回肠胰蛋白酶活性与对照组差异不显著(P>0.05)。各组之间60日龄十二指肠、空肠、回肠脂肪酶活性无显著差异(P>0.05),异亮氨酸组90日龄十二指肠脂肪酶活性极显著低于对照组和苏氨酸组(P<0.01)。
表6 苏氨酸和异亮氨酸对60日龄皖西白鹅肠道消化酶活性的影响

Table 6 Effects of threonine and isoleucine on intestinal digestive enzyme activities of Wanxi white geese at 60 days of age U/g prot

项目
Items		 对照组
CON group		 苏氨酸组
Thr group		 异亮氨酸组
Ile group		 P值
P-value
十二指肠Duodenum
淀粉酶Amylase 0.07±0.00 0.08±0.00 0.07±0.00 0.17
麦芽糖酶Maltase 125.78±6.54b 145.58±3.97a 125.79±3.37b 0.02
乳糖酶Lactase 3.04±0.08 3.29±0.13 4.30±0.65 0.06
胰蛋白酶Trypsin 1 127.33±62.48Bb 1 384.88±75.05Aa 1 570.66±38.69Aa <0.01
脂肪酶Lipase 0.75±0.05 0.78±0.09 0.63±0.08 0.22
空肠Jejunum
淀粉酶Amylase 0.06±0.00 0.08±0.01 0.06±0.00 0.17
麦芽糖酶Maltase 273.81±17.80 275.02±17.24 286.11±5.69 0.58
乳糖酶Lactase 12.19±0.03ab 12.88±0.53a 10.64±0.32b 0.02
胰蛋白酶Trypsin 1 317.33±337.50 1 106.22±561.50 1 676.22±143.50 0.35
脂肪酶Lipase 0.94±0.16 1.22±0.13 1.00±0.26 0.37
回肠Ileum
淀粉酶Amylase 0.11±0.00 0.15±0.05 0.20±0.04 0.20
麦芽糖酶Maltase 189.35±3.82Bb 194.12±24.86Bb 337.62±34.47Aa <0.01
乳糖酶Lactase 2.24±0.67 2.98±0.42 2.52±1.01 0.52
胰蛋白酶Trypsin 509.55±16.36 474.00±22.47 523.11±27.63 0.18
脂肪酶Lipase 1.37±0.20 1.17±0.03 1.21±0.07 0.36
表7 苏氨酸和异亮氨酸对90日龄皖西白鹅肠道消化酶活性的影响

Table 7 Effects of threonine and isoleucine on intestinal digestive enzyme activities of Wanxi white geese at 90 days of age U/g prot

项目
Items		 对照组
CON group		 苏氨酸组
Thr group		 异亮氨酸组
Ile group		 P值
P-value
十二指肠Duodenum
淀粉酶Amylase 0.07±0.00 0.08±0.00 0.07±0.00 0.17
麦芽糖酶Maltase 141.18±7.40b 217.36±11.78a 186.06±13.87ab 0.03
乳糖酶Lactase 2.35±0.71 2.49±0.14 3.63±0.67 0.17
胰蛋白酶Trypsin 1 127.33±62.48Bb 1 384.88±75.05Aa 1 570.66±38.69Aa <0.01
脂肪酶Lipase 1.28±0.04Aa 1.27±0.02Aa 0.99±0.02Bb <0.01
空肠Jejunum
淀粉酶Amylase 0.04±0.00b 0.09±0.01a 0.09±0.00a 0.02
麦芽糖酶Maltase 306.44±24.86 239.55±18.32 297.72±32.34 0.12
乳糖酶Lactase 4.26±0.69Bb 12.01±0.73Aa 4.53±0.15Bb <0.01
胰蛋白酶Trypsin 1 625.55±238.80 2 128.00±890.66 1 507.33±113.52 0.45
脂肪酶Lipase 1.38±0.29 1.22±0.04 0.77±0.14 0.07
回肠Ileum
淀粉酶Amylase 0.16±0.02 0.17±0.02 0.18±0.03 0.75
麦芽糖酶Maltase 292.75±20.30Bb 584.59±39.45Aa 365.29±40.61Bb <0.01
乳糖酶Lactase 5.53±1.43 6.14±0.81 5.62±0.95 0.71
胰蛋白酶Trypsin 789.55±92.02 874.00±285.02 793.77±195.18 0.78
脂肪酶Lipase 2.24±0.41 2.10±0.28 1.41±0.07 0.14

2.5 苏氨酸和异亮氨酸对60、90日龄皖西白鹅肠道抗氧化指标的影响

由表8、表9可知,各组之间60、90日龄十二指肠、空肠、回肠SOD和GSH-Px活性均无显著差异(P>0.05)。异亮氨酸组60日龄空肠T-AOC显著高于对照组和苏氨酸组(P<0.05),各组之间90日龄十二指肠、空肠、回肠T-AOC无显著差异(P>0.05)。苏氨酸组60日龄空肠MDA含量极显著低于对照组和异亮氨酸组(P<0.01),异亮氨酸组90日龄回肠MDA含量显著低于对照组(P<0.05)。
表8 苏氨酸和异亮氨酸对60日龄皖西白鹅肠道抗氧化指标的影响

Table 8 Effects of threonine and isoleucine on intestinal antioxidant indexes of Wanxi white geese at 60 days of age

项目
Items		 对照组
CON group		 苏氨酸组
Thr group		 异亮氨酸组
Ile group		 P值
P-value
十二指肠Duodenum
超氧化物歧化酶SOD/(U/mg prot) 24.28±2.23 28.60±3.42 19.43±2.66 0.06
谷胱甘肽过氧化物酶GSH-Px/(U/g prot) 86.37±7.66 90.96±13.22 90.24±4.63 0.74
总抗氧化能力T-AOC/(U/g prot) 0.95±0.03 0.96±0.00 0.96±0.00 0.82
丙二醛MDA/(nmol/mg prot) 3.86±0.80 2.61±0.71 2.05±0.04 0.09
空肠Jejunum
超氧化物歧化酶SOD/(U/mg prot) 26.60±4.29 32.90±1.10 35.80±5.30 0.16
谷胱甘肽过氧化物酶GSH-Px/(U/g prot) 145.50±2.13 148.20±3.92 148.89±5.02 0.57
总抗氧化能力T-AOC/(U/g prot) 0.53±0.04b 0.60±0.03b 0.82±0.06a 0.01
丙二醛MDA/(nmol/mg prot)) 6.79±0.17Aa 0.85±0.19Bb 5.08±0.89Aa <0.01
回肠Ileum
超氧化物歧化酶SOD/(U/mg prot) 30.01±1.85 30.23±4.08 32.52±0.62 0.53
谷胱甘肽过氧化物酶GSH-Px/(U/g prot) 5.72±0.54 6.63±0.86 6.85±1.80 0.47
总抗氧化能力T-AOC/(U/g prot) 0.73±0.12 0.53±0.03 0.63±0.04 0.20
丙二醛MDA/(nmol/mg prot) 3.90±0.71 3.69±0.60 1.58±0.12 0.06
表9 苏氨酸和异亮氨酸对90日龄皖西白鹅肠道抗氧化指标的影响

Table 9 Effects of threonine and isoleucine on intestinal antioxidant indexes of Wanxi white geese at 90 days of age

项目
Items		 对照组
CON group		 苏氨酸组
Thr group		 异亮氨酸组
Ile group		 P值
P-value
十二指肠Duodenum
超氧化物歧化酶SOD/(U/mg prot) 24.28±2.23 28.60±3.42 19.43±2.66 0.06
谷胱甘肽过氧化物酶GSH-Px/(U/g prot) 103.22±4.94 116.05±6.31 109.15±13.35 0.36
总抗氧化能力T-AOC/(U/g prot) 0.86±0.03 0.92±0.01 0.93±0.01 0.05
丙二醛MDA/(nmol/mg prot) 5.18±0.46 4.55±1.14 4.87±0.17 0.57
空肠Jejunum
超氧化物歧化酶SOD/(U/mg prot) 20.74±0.29 20.67±1.32 20.22±1.46 0.76
谷胱甘肽过氧化物酶GSH-Px/(U/g prot) 225.73±26.64 234.52±4.26 228.61±16.36 0.75
总抗氧化能力T-AOC/(U/g prot) 0.70±0.06 0.83±0.08 0.54±0.06 0.05
丙二醛MDA/(nmol/mg prot) 4.48±1.15 3.06±0.84 2.65±0.96 0.25
回肠Ileum
超氧化物歧化酶SOD/(U/mg prot) 32.52±4.99 32.81±0.65 33.74±0.66 0.78
谷胱甘肽过氧化物酶GSH-Px/(U/g prot) 15.38±4.67 16.36±1.19 20.54±0.89 0.25
总抗氧化能力T-AOC/(U/g prot) 0.59±0.11 0.70±0.00 0.82±0.09 0.10
丙二醛MDA/(nmol/mg prot) 0.89±0.14a 0.78±0.13ab 0.39±0.03b 0.03

2.6 苏氨酸和异亮氨酸对90日龄皖西白鹅血清和空肠免疫指标的影响

由表10可知,对照组、苏氨酸组和异亮氨酸组之间90日龄血清IgA、IgG、IgM、IL-2、IL-4、IFN-γ、TNF-α含量均无显著差异(P>0.05)。异亮氨酸组90日龄空肠IgA含量显著低于对照组和苏氨酸组(P<0.05),苏氨酸组空肠IgG、IgM含量极显著高于对照组和异亮氨酸组(P<0.01),对照组、苏氨酸组和异亮氨酸组之间空肠IL-2含量无显著差异(P>0.05),异亮氨酸组空肠IL-4和IFN-γ含量显著低于对照组和苏氨酸组(P<0.05),异亮氨酸组和苏氨酸组空肠TNF-α含量极显著低于对照组(P<0.01)。
表10 苏氨酸和异亮氨酸对90日龄皖西白鹅血清和空肠免疫指标的影响

Table 10 Effects of threonine and isoleucine on serum and jejunal immune indexes of Wanxi white geese at 90 days of age

项目
Items		 对照组
CON group		 苏氨酸组
Thr group		 异亮氨酸组
Ile group		 P值
P-value
血清Serum
免疫球蛋白A IgA/(mg/mL) 21.27±0.94 24.62±2.76 21.27±0.10 0.09
免疫球蛋白G IgG/(mg/mL) 15.33±3.75 16.38±4.79 8.99±0.35 0.08
免疫球蛋白M IgM/(mg/mL) 9.63±0.42 10.26±0.13 9.79±0.30 0.22
白细胞介素-2 IL-2/(ng/L) 156.02±22.18 171.47±8.49 166.48±16.96 0.55
白细胞介素-4 IL-4/(ng/L) 937.33±287.00 564.15±235.68 494.11±345.66 0.34
干扰素-γ IFN-γ/(ng/L) 260.07±6.50 218.77±26.30 242.92±34.62 0.30
肿瘤坏死因子-α TNF-α/(ng/L) 803.36±244.55 601.91±152.04 598.13±172.20 0.49
空肠Jejunum
免疫球蛋白A IgA/(mg/g prot) 34.15±0.55a 32.62±1.07a 26.39±2.27b 0.01
免疫球蛋白G IgG/(mg/g prot) 1.74±0.25Bb 3.82±0.17Aa 1.16±0.16Bb <0.01
免疫球蛋白M IgM/(mg/g prot) 1.31±0.35Bb 4.49±1.06Aa 1.52±0.15Bb <0.01
白细胞介素-2 IL-2/(ng/kg prot) 79.62±14.32 87.74±5.98 80.06±9.54 0.61
白细胞介素-4 IL-4/(ng/kg prot) 63.83±9.09a 46.58±10.90a 35.60±4.14b 0.04
干扰素-γ IFN-γ/(ng/kg prot) 85.50±11.50a 81.94±4.47a 49.17±5.75b 0.01
肿瘤坏死因子-α TNF-α/(ng/kg prot) 116.80±21.72Aa 46.02±5.26Bb 36.96±7.06Bb <0.01

3 讨论

3.1 苏氨酸和异亮氨酸对皖西白鹅消化吸收功能的影响

消化吸收功能是畜禽消化道整体健康状况的具体体现,屠宰性能、器官指数、肠道形态、消化酶活性等指标主要与肉鹅消化吸收功能及生长性能相关。畜禽屠宰性能各项指标在一定程度上体现在了畜产品的经济价值,器官指数是反映器官生长发育的重要指标之一。前人研究发现,苏氨酸对鹅和鸭的屠宰性能和器官指数无显著影响[14-15]。本课题组前期研究表明,饲粮添加苏氨酸和异亮氨酸均显著提高了60~90日龄皖西白鹅饲料转化效率[16];本试验中,饲粮添加苏氨酸和异亮氨酸对皖西白鹅屠宰性能和器官指数无显著影响,与前人研究结果[17-18]一致,主要原因可能与添加浓度和生长阶段有关,后续将开展进一步的研究。
肠道的形态结构与其消化吸收密切相关,绒毛高度、隐窝深度及V/C是常用评价肠道形态的指标。饲粮苏氨酸缺乏会引发畜禽的肠道形态变化,包括小肠绒毛高度降低、V/C下降、绒毛面积变小[19⇓-21]。Ren等[22]发现在低蛋白质饲粮(粗蛋白质水平17%)中添加支链氨基酸(BCAA)使仔猪十二指肠绒毛高度增加。本试验研究发现,饲粮添加异亮氨酸提高了60日龄十二指肠绒毛高度和V/C及回肠绒毛高度、隐窝深度;饲粮添加苏氨酸提高了90日龄十二指肠隐窝深度、V/C,空肠绒毛高度、隐窝深度、V/C以及回肠绒毛高度和V/C,与前人研究结果[23]一致,说明异亮氨酸和苏氨酸均可改善生长期皖西白鹅的小肠形态,增加肠道消化吸收面积,提高消化吸收效率,与生长性能的结果[16]一致。
除了肠道形态,消化酶活性也是评估畜禽消化吸收机能的重要指标,其参与食物的化学消化,营养、免疫和应激等均可影响消化酶的分泌和活性[24-25],功能性氨基酸可增加消化酶的合成进而影响其功能的发挥[26]。脂肪酶是由胰腺分泌至十二指肠的基本酶类,它可将脂肪水解成为游离的脂肪酸、甘油和甘油单酯;二糖不能被小肠直接吸收利用,因此小肠二糖酶对碳水化合物的利用极其重要,禽类黏膜二糖酶主要以蔗糖酶、麦芽糖酶和异麦芽糖酶为主[27-28]。报道显示,苏氨酸可提高蛋鸡空肠的淀粉酶和脂肪酶活性[29],对水产动物肠道消化酶活性影响显著[30-31],而异亮氨酸可提高水产动物肠道淀粉酶活性[32]。本试验发现,饲粮添加苏氨酸和异亮氨酸可以促进十二指肠胰蛋白酶吸收利用,添加苏氨酸对空肠乳糖酶活性有显著影响,添加异亮氨酸对空肠麦芽糖酶活性有显著影响。
苏氨酸和异亮氨酸一方面作为蛋白质合成的底物,可促进消化系统内相关功能蛋白的表达和功能酶的合成[33-34],因此肠细胞生长速度和消化酶活性增加;另一方面作为肠道上皮细胞的能量供体,苏氨酸和异亮氨酸增加均可提高肠道上皮细胞的增殖。以上2个方面作用使小肠绒毛高度增加,提高了消化吸收能力,最终提高了畜禽的生长性能。

3.2 苏氨酸和异亮氨酸对皖西白鹅抗氧化能力的影响

在动物体中有氧细胞代谢通常会产生各种活性氧(ROS)和活性氮(RNS),并在炎症情况下产生累积。活性氧在体内会损伤细胞大分子(蛋白质、脂质和DNA)诱发细胞氧化应激,破坏黏膜上皮的完整性,导致肠道屏障功能受损及动物发病率增加[33]。MDA含量、T-AOC和SOD、GSH-Px活性的变化不仅反映了清除氧自由基的能力,也间接反映了自由基对细胞膜损伤的程度[35]。Azzam等[36]发现饲粮中添加苏氨酸提高了蛋鸡SOD和GSH-Px活性。Zhao等[37]发现异亮氨酸(12.5~15.0 g/kg)通过增加SOD、GSH-Px、谷胱甘肽还原酶、谷胱甘肽S-转移酶的活性和表达以增加抗氧化能力并减少氧化损伤。本试验发现,饲粮添加苏氨酸可降低空肠MDA含量;添加苏氨酸可提高空肠T-AOC,降低回肠MDA含量。

3.3 苏氨酸和异亮氨酸对皖西白鹅免疫功能的影响

苏氨酸可以改善紧密连接、细胞迁移、肠道中的抗氧化反应和黏膜屏障功能[38];同样,支链氨基酸也通过促进肠道发育、营养转运蛋白和免疫相关功能的调节剂来改善肠道健康[39]。体内外试验证明,异亮氨酸可调节免疫器官、免疫细胞和免疫反应物质(如免疫球蛋白、细胞因子和宿主防御肽)[40]。支链氨基酸调节免疫功能,包括增加肠道免疫球蛋白、先天性和适应性免疫应答、促炎细胞因子和树突状细胞功能[41-42]。苏氨酸是免疫球蛋白的组成成分之一,饲粮添加苏氨酸可提高鸡和鸭的免疫球蛋白表达,降低IFN-γ含量,调控肠道微生物菌群[8,29,43]。本试验发现,饲粮添加苏氨酸可以提高空肠IgG、IgM含量,添加异亮氨酸可以降低空肠IL-4、IFN-γ含量,添加苏氨酸和异亮氨酸均可以降低空肠TNF-α含量。促炎细胞因子IFN-γ和TNF-α可通过改变肠道紧密连接中的膜微脂组成破坏肠上皮紧密连接的屏障功能[44]。苏氨酸通过复杂的信号网络影响肠道免疫系统[45],也被认为是细胞生长和增殖不可或缺的营养物质。支链氨基酸在蛋白质合成以及肠道健康和免疫起关键作用,磷脂酰肌醇-3-激酶(PI3K)/蛋白激酶B(AKT)/哺乳动物雷帕霉素靶蛋白(mTOR)作为主要的信号通路介导支链氨基酸发挥免疫调节功能[46]。

4 结论

异亮氨酸和苏氨酸对生长期皖西白鹅屠宰性能和器官生长无显著影响,但异亮氨酸和苏氨酸分别提高了30~60日龄、61~90日龄皖西白鹅肠道组织结构和抗氧化功能,同时也提高了小肠免疫功能。

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