Chinese Journal of Animal Nutrition >

2023 , Vol. 35 >Issue 1: 505 - 513

DOI: https://doi.org/10.3969/j.issn.1006-267x.2023.01.048

RESEARCH PAPER

Effects of Dietary Tributyrin on Growth Performance, Serum Biochemical Indices, Immune Function and Antioxidant Capacity of Juvenile Yellow Catfish (Pelteobagrus fulvidraco)

  • HOU Dongqiang , 1, 2 ,
  • LI Peijia 1, 2 ,
  • ZHAO Hongxia , 1, ** ,
  • CHEN Xiaoying 1 ,
  • ZHENG Chuntian 1 ,
  • CAO Junming 2
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  • 1 Guangdong Key Laboratory of Animal Breeding and Nutrition, Key Laboratory of Animal Nutrition and Feed Science in South China of Ministry of Agriculture and Rural Affairs, Institute of Animal Science, Guangdong Academy of Agricultural Sciences, Guangzhou 510640, China
  • 2 College of Fisheries, Guangdong Ocean University, Zhanjiang 524088, China
** professor, E-mail:

* Contributed equally

Received date: 2022-05-31

  Online published: 2023-01-11

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Abstract

This experiment was conducted to investigate the effects of dietary tributyrin on growth performance, serum biochemical indices, immunity function and antioxidant capacity of juvenile yellow catfish (Pelteobagrus fulvidraco). A total of 480 juvenile yellow catfish with an initial body weight of (5.53±0.02) g were selected and randomly divided into 4 groups with 3 replicates in each group, and 40 fish in each replicate. Four isonitrogenous and isolipid experimental diets were fed with 0 (G0), 500 (G1), 1 000 (G2) and 2 000 mg/kg (G3) triglyceride, respectively. The experiment lasted for 56 days. The results showed as follows: 1) weight gain rate and feed efficiency rate in G2 group were significantly higher than those in other groups (P<0.05). The fatness in G1 group was significantly lower than that in G0 and G3 groups (P<0.05), but there was no significant difference between G1 group and G2 group (P>0.05). 2) The serum glucose content in G3 group was significantly higher than that in G1 and G2 groups (P<0.05), and there were no significant differences in the serum contents of total protein, albumin, globulin, triglyceride, urea nitrogen and the activities of alanine aminotransferase and aspartate aminotransferase among all groups (P>0.05). 3) Compared with G0 group, serum lysozyme activity in G1, G2 and G3 groups was significantly increased (P<0.05), serum alkaline phosphatase activity in G1 and G2 groups was significantly increased (P<0.05), and serum immunoglobulin M content in G2 group was significantly higher than that in G3 group (P<0.05). There was no significant difference in serum complement C3 content among all groups (P>0.05). 4) Compared with G0 group, serum total antioxidant capacity in G1 group was significantly increased (P<0.05), and serum superoxide dismutase activity in G1, G2 and G3 groups was significantly increased (P<0.05). The serum catalase activity in G3 group was significantly higher than that in G0, G1 and G2 groups (P<0.05), but there were no significant differences in serum glutathione peroxidase activity, glutathione and malondialdehyde contents among all groups (P>0.05). In conclusion, dietary supplementation of 500 to 2 000 mg/kg triglyceride can significantly improve non-specific immune and antioxidant capacity of yellow catfish, and dietary supplementation of 1 000 mg/kg triglyceride can significantly improve weight gain rate and feed efficiency of yellow catfish. Combined with the comprehensive evaluation of growth performance, serum immunity and antioxidant indexes, the optimal supplemental level of triglyceride in yellow catfish diet is 1 000 mg/kg.

Key words： yellow catfish; triglyceride tributyrate; growth performance; serum biochemical indices; immune; antioxidant

Cite this article

HOU Dongqiang , LI Peijia , ZHAO Hongxia , CHEN Xiaoying , ZHENG Chuntian , CAO Junming . Effects of Dietary Tributyrin on Growth Performance, Serum Biochemical Indices, Immune Function and Antioxidant Capacity of Juvenile Yellow Catfish (Pelteobagrus fulvidraco)[J]. Chinese Journal of Animal Nutrition, 2023 , 35(1) : 505 -513 . DOI: 10.3969/j.issn.1006-267x.2023.01.048

有机酸或有机酸盐作为饲料添加剂,能够促进动物的生长性能、增强免疫功能和改善消化性能[1-2]。甲酸、乙酸、丁酸等有机酸化剂常温下容易蒸发产生异味,从而影响动物摄食。三丁酸甘油酯(tributyrin,TB)是一种非挥发性的自然稳定物质,由甘油及附着在甘油主链上的3个丁酸分子组成,其分子式为C15H26O6[3]。三丁酸甘油酯能够在肠道中的脂肪酶作用下水解生成甘油和丁酸,甘油和丁酸可进入细胞参与营养代谢并为小肠上皮细胞提供能量[4-5]。研究表明,饲料中添加0.1%三丁酸甘油酯可提高黄姑鱼(Nibea albiflora)生长性能[6];鲤鱼(Cyprinus carpio)饲料中添加2.0~4.0 g/kg三丁酸甘油酯显著改变肠道形态、微生物群分布的能力,从而提高鲤鱼的生长性能[7];添加443.5 mg/kg三丁酸甘油酯可改善草鱼(Ctenopharyngodon idella)肠道健康,从而提高草鱼的生长性能[8];饲料中添加0.06%三丁酸甘油酯可显著增加团头鲂(Megalobrama amblycephala)免疫、抗氧化能力、增重率和饲料利用率,显著提高团头鲂的生长性能[5]。以上研究结果证明,饲料添加三丁酸甘油酯能够显著促进水产动物的生长性能、免疫功能和肠道健康。
黄颡鱼(Pelteobagrus fulvidraco),鲶形目、鲿科、黄颡鱼属,是我国重要的经济杂食性淡水鱼,肉质鲜美,营养价值高。随着我国水产养殖规模化水平不断提高,黄颡鱼的健康状况和病害防治也逐渐引起人们的重视。本实验室前期研究表明,饲料中添加丁酸钠能够显著提高黄颡鱼生长性能、营养物质的沉积和抗氨氮应激能力[9-10]。然而,目前在黄颡鱼中尚未见到三丁酸甘油酯的相关报道。因此,本试验通过在饲料中分别添加不同水平的三丁酸甘油酯,研究其对黄颡鱼生长性能、血清生化指标、免疫功能及抗氧化能力的影响,评估三丁酸甘油酯在实用饲料中的适宜添加水平,旨在为三丁酸甘油酯在黄颡鱼的配合饲料中应用提供理论依据和技术支撑。

1 材料与方法

1.1 试验饲料

基础饲料组成及营养水平见表1。在基础饲料中分别补充0、500、1 000、2 000 mg/kg的三丁酸甘油酯(纯度>90%)配制成4种等氮(42%)等脂(9%)试验饲料,分别记作G0、G1、G2、G3。按配方准确称取饲料原料,并将饲料原料经粉碎通过60目筛并逐级混匀,然后加入鱼油、豆油,加水搅拌均匀,使用双螺杆挤压机(SLX-80型,华南理工大学科技实业总厂生产)将原料挤压成条,挤压机工作运转部位轴承温度保持45~50 ℃,亦可通过注入机器流水,挤压机工作时度则可控制在45 ℃以下。通过制粒机(G-500,华南理工大学科技实业总厂生产)破碎条状饲料,制成1.5 mm的颗粒饲料,在55 ℃下烘干,自然冷却后放入密封袋中,置于-20 ℃冰箱中保存备用。
表1 基础饲料组成及营养水平(干物质基础)

Table 1 Composition and nutrient levels of the basal diet (DM basis)%

项目Items 含量Content
原料Ingredients
鱼粉Fish meal 25.00
豆粕Soybean meal 30.00
菜籽粕Rapeseed meal 9.00
玉米蛋白粉Corn gluten meal 6.00
面粉Wheat flour 22.30
鱼油Fish oil 2.50
豆油Soybean oil 2.50
维生素预混料Vitamin premix1) 0.10
矿物质预混料Mineral premix2) 0.50
磷酸二氢钙Ca(H2PO4)2 1.50
维生素C磷酸酯
Vitamin C phosphate(35%)		 0.10
氯化胆碱Choline chloride 0.30
微晶纤维素Microcrystalline cellulose 0.20
合计Total 100.00
营养水平Nutrient levels3)
粗蛋白质Crude protein 42.6
粗脂肪Crude lipid 9.0
粗灰分Ash 8.3
水分Moisture 7.8

1)每千克维生素预混料含有 One kilogram of vitamin premix contained the following: VA 3 200 000 IU,VB1 4 g,VB2 8 g,VB6 4.8 g,VB12 16 mg,VD3 1 600 000 IU,VE 16 g,VK 4 g,泛酸钙 calcium pantothenate 16 g,叶酸 folic acid 1.28 g,烟酸 nicotinic acid 28 g,肌醇 inositol 40 g,生物素 biotin 64 mg。

2)每千克矿物质预混料含有 One kilogram of mineral premix contained the following: MgSO4·H2O 12 g,Ca(IO3)2 9 g,KCl 36 g,Met-Cu 1.5 g,ZnSO4·H2O 10 g,FeSO4·H2O 1 g,Met-Co 250 mg,NaSeO3 0.003 6 g。

3)营养水平为实测值。Nutrient levels were measured values.

1.2 试验设计及饲养管理

黄颡鱼苗购自广州市某水产养殖场,在室内循环水养殖系统中,用基础饲料驯化2周。选取480尾黄颡鱼幼鱼[(5.53±0.02) g],分别置于12个350 L的玻璃纤维圆柱形水箱中开展正式试验,每箱放鱼40尾。将4种试验饲料随机分配到4个组,每组3个重复。试验期间,每天分别于08:30和18:30投喂饲料,采取表观饱食投喂。养殖试验条件:水温27~32 ℃,pH 7.5~7.9,溶氧含量>6.0 mg/L,亚硝酸盐含量<0.01 mg/L,氨氮含量<0.20 mg/L,自然光照。试验期为56 d,每天观察黄颡鱼的摄食情况、健康状态,并记录。

1.3 采集样品

养殖试验结束后,禁食24 h,分别对各水箱的鱼进行称重和计数。取样前先用MS-222溶液(120 mg/L)麻醉处理。每箱随机取6尾鱼进行解剖,测量其体重、体长、肝脏重和内脏重。每箱取6尾麻醉鱼尾静脉采血,在4 ℃下以4 000 r/min离心15 min,收集血清保存于-80 ℃冰箱。

1.4 指标测定

1.4.1 生长性能和形态指标计算

增重率(%)=100×(Gt-G0)/G0;
成活率(%)=100×Nt/N0;
饲料效率(%)=100×(Gt-G0)/Gf;
摄食量[g/(尾·d)]=Gf/[(N0+Nt)×t/2];
肥满度(%)=100×Gt/ L t 3;
肝体比(%)=100×Gg/Gq;
脏体比(%)=100×Gn/Gq。
式中:G0表示鱼体初始体质量(g);Gt表示鱼体终末体质量(g);N0表示初始鱼尾数;Nt表示终末鱼尾数;t表示试验天数(d);Gf表示饲料摄入量(g);Lt表示试验鱼终末体长(cm);Gg表示肝脏重(g);Gn表示内脏重(g);Gq表示全鱼重(g)。

1.4.2 营养成分测定

采用AOAC(2019)[11]方法测定饲料的水分、粗蛋白质、粗脂肪和粗灰分含量。在105 ℃下将样品干燥至恒重后测定水分含量;采用凯氏定氮法测定粗蛋白质含量;采用索氏抽提法测定粗脂肪含量;在600 ℃的马弗炉中燃烧6 h后检测粗灰分含量。

1.4.3 血清生化指标测定

血清总蛋白、白蛋白、球蛋白、甘油三酯、尿素氮、葡萄糖含量及谷丙转氨酶和谷草转氨酶活性采用全自动化学分析仪(日立7600-110,日本)测定。

1.4.4 血清免疫和抗氧化指标测定

采用南京建成生物工程研究所试剂盒测定血清溶菌酶(LZM)、碱性磷酸酶(AKP)、过氧化氢酶(CAT)、谷胱甘肽过氧化物酶(GSH-Px)、超氧化物歧化酶(SOD)活性,总抗氧化能力(T-AOC)及谷胱甘肽(GSH)和丙二醛(MAD)含量。根据说明书(R&D Systems, Minneapolis,美国),采用酶联免疫吸附试验(ELISA)测定血清补体3 (C3)、免疫球蛋白M(IgM)含量。

1.5 数据统计

采用SPSS 19.0软件进行数据的统计分析,采用one-way ANOVA程序进行单因素方差分析,采用Duncan氏法进行组间的多重比较。P<0.05表示差异显著。试验数据使用平均值±标准误(n=3)表示。

2 结果与分析

2.1 生长性能和形体指标

由表2可知,G2组增重率和饲料效率显著高于其他各组(P<0.05)。G1组肥满度显著低于G0和G3组(P<0.05)。各组间的摄食量、肝体比和脏体比无显著差异(P>0.05)。
表2 饲料中三丁酸甘油酯水平对黄颡鱼生长性能和形态指标的影响

Table 2 Effects of dietary tributyrin levels on growth performance and physical indexes of yellow catfish (Pelteobagrus fulvidraco)

项目
Items		 组别Groups P值
P-value
G0 G1 G2 G3
增重率WGR/% 256.55±8.00a 265.02±33.28a 358.59±18.59b 279.50±14.87a 0.024
饲料效率FE/% 71.46±5.04a 76.55±2.95a 90.65±3.70b 73.96±3.41a 0.031
摄食量FI/[(g/(尾·d)] 0.36±0.02 0.34±0.02 0.39±0.02 0.37±0.02 0.393
肥满度CF/(g/cm3) 1.53±0.03b 1.43±0.02a 1.46±0.02ab 1.51±0.02b 0.019
肝体比HSI/% 1.29±0.06 1.28±0.05 1.42±0.05 1.32±0.04 0.190
脏体比VSI/% 6.87±0.31 6.76±0.27 7.51±0.27 6.92±0.46 0.399

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

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

2.2 血清生化指标

由表3可知,G3组血清葡萄糖含量显著高于G1和G2组(P<0.05)。各组间的血清总蛋白、白蛋白、球蛋白、甘油三酯、尿素氮含量及谷丙转氨酶、谷草转氨酶活性无显著差异(P>0.05)。
表3 饲料中三丁酸甘油酯水平对黄颡鱼血清生化指标的影响

Table 3 Effects of dietary tributyrin levels on serum metabolite indices of yellow catfish (Pelteobagrus fulvidraco)

项目
Items		 组别Groups P值
P-value
G0 G1 G2 G3
总蛋白TP/(g/L) 32.80±0.36 32.77±1.60 33.80±0.52 33.23±0.15 0.816
白蛋白ALB/(g/L) 12.03±0.44 11.93±0.29 12.70±0.06 12.40±0.35 0.364
球蛋白GLB/(g/L) 20.76±0.24 20.83±1.31 21.10±0.49 20.83±0.41 0.989
甘油三酯TG/(mmol/L) 4.32±0.44 4.10±0.45 4.40±0.11 4.42±0.29 0.907
葡萄糖GLU/(mmol/L) 3.73±0.42a 4.12±0.37a 4.73±0.28ab 5.61±0.26b 0.020
尿素氮UN/(mmol/L) 0.77±0.09 0.73±0.03 0.83±0.03 0.80±0.06 0.659
谷草转氨酶AST/(U/L) 560.04±22.72 498.43±1.53 492.98±39.05 554.11±14.42 0.159
谷丙转氨酶ALT/(U/L) 18.67±2.91 19.33±3.18 13.67±0.88 18.33±0.67 0.324

2.3 血清免疫指标

由表4可知,与G0组相比,G1、G2和G3组的血清LZM活性显著升高(P<0.05),G1和G2组血清AKP活性显著升高(P<0.05)。G2组的血清IgM含量显著高于G3组(P<0.05)。各组间的血清C3含量无显著差异(P>0.05)。
表4 饲料中三丁酸甘油酯水平对黄颡鱼血清免疫指标的影响

Table 4 Effects of dietary tributyrin levels on serum immune indices of yellow catfish (Pelteobagrus fulvidraco)

项目
Items		 组别Groups P值
P-value
G0 G1 G2 G3
溶菌酶LZM/(U/mL) 27.85±8.94a 67.11±4.84b 65.76±11.10b 58.92±6.08b 0.029
碱性磷酸酶AKP/(U/L) 4.80±0.28a 6.76±0.71b 6.33±0.23b 5.69±0.18ab 0.043
补体3 C3/(μg/mL) 1 351.09±74.60 1 151.71±18.35 1 161.55±107.22 1 227.54±138.95 0.474
免疫球蛋白M IgM/(μg/mL) 4.83±0.21ab 4.91±0.18ab 5.20±0.25b 4.30±0.04a 0.036

2.4 血清抗氧化指标

由表5可知,与G0组相比,G1组血清T-AOC升高(P<0.05),G1、G2和G3组的血清SOD活性显著升高(P<0.05)。G3组血清CAT活性显著高于G0、G1和G2组(P<0.05)。各组间的血清GSH-Px活性及GSH和MDA含量无显著差异(P>0.05)。
表5 饲料中三丁酸甘油酯水平对黄颡鱼血清抗氧化指标的影响

Table 5 Effects of dietary tributyrin levels on serum antioxidant indices of yellow catfish (Pelteobagrus fulvidraco)

项目
Items		 组别Groups P值
P-value
G0 G1 G2 G3
总抗氧化能力T-AOC/(U/mL) 3.31±0.36a 5.01±0.54b 3.83±0.14ab 4.09±0.42ab 0.042
超氧化物歧化酶SOD/(U/mL) 154.14±6.93a 180.33±1.49b 203.96±8.96c 186.73±1.51bc 0.004
过氧化氢酶CAT/(U/mL) 59.83±4.45a 65.38±4.91a 64.08±4.18a 92.70±3.68b 0.002
谷胱甘肽过氧化物酶
GSH-Px/(U/mL)		 257.36±12.30 228.85±31.01 304.78±18.50 249.27±14.73 0.248
谷胱甘肽GSH/(mg/L) 35.57±2.30 41.69±4.21 40.03±1.72 41.22±4.72 0.606
丙二醛MDA/(nmol/mL) 13.02±0.25 12.63±0.19 12.61±0.36 11.92±0.81 0.468

3 讨论

3.1 三丁酸甘油酯对黄颡鱼生长性能和形体指标的影响

三丁酸甘油酯在胰脂肪酶的水解下缓慢释放丁酸,被肠道上皮细胞吸收,增加肠道绒毛高度,为肠道细胞增殖分化提供能源物质[12]。在黄姑鱼[6]、草鱼[8]、鲤鱼[7]、杂交石斑鱼(Epinephelus fuscoguttatus♀×Epinephelus lanceolatus♂)[13]、大黄鱼(Larimichthys crocea)[14]、乌醴(Channa argus)[15]等的研究表明,饲料中添加适宜水平的三丁酸甘油酯能够显著提高增重率和降低饲料系数。本试验投喂黄颡鱼饲料为颗粒料,制粒温度保持在45~50 ℃,在饲料加工过程中可充分保护三丁酸甘油酯不被破坏;在实际生产中黄颡鱼饲料多为膨化料,制作膨化饲料时温度在130~160 ℃,原料在高温阶段时间很短(5~10 s),而三丁酸甘油酯在190 ℃的温度下分子才会变形,所以膨化过程中亦可保证其活性。本研究中,饲料添加3种不同水平的三丁酸甘油酯均可提高黄颡鱼的增重率和降低饲料效率,增重率和饲料效率呈现先升高后下降趋势,在1 000 mg/kg组达到最高,显著高于其他各组。与本试验结果一致,饲料添加一定剂量三丁酸甘油酯(0.5、1.0、1.5 g/kg)显著提高凡纳滨对虾(Litopenaeus vannamei)生长性能,当添加量大于1.5 g/kg时,凡纳滨对虾生长性能有所降低[16]。Volatiana等[17]发现饲料中添加6 g/kg的三丁酸甘油酯时,随着丁酸甘油酯添加量的增加,黑鲷(Acanthopagrus schlegelii)幼鱼的生长性能逐渐提高,超过此添加量,对脂肪酶和淀粉酶活性的影响显著降低,生长性能有所下降。三丁酸甘油酯在肠道中可分解为甘油和丁酸,丁酸可提高肠道消化酶活性,改善肠道形态结构提高消化率,适量添加三丁酸甘油酯可提高动物生长性能[16-17],过量添加可能会导致肠道pH过低,损害肠道菌群,降低肠道对营养物质的消化能力,抑制动物生长[18]。丁酸甘油酯独特的脂肪香味可提高饲料的适口性,饲料酸化后可增加动物唾液分泌量,增强动物食欲,增加动物的摄食量[19]。本试验条件下,1 000 mg/kg组和2 000 mg/kg组摄食量高于对照组,500 mg/kg组摄食量低于对照组,1 000 mg/kg组摄食量高于其他各组,但各组间无显著差异。肥满度、肝体比、肠体比、脏体比和肠长比是反映鱼体、肝脏、肠道、内脏发育状况和衡量脏器功能的重要指标[20]。本研究结果表明,500 mg/kg组肥满度显著低于0和2 000 mg/kg组,各组间的摄食量、肝体比和脏体比无显著差异。三丁酸甘油酯促进动物生长的机制有多种:一是丁酸可降低动物肠道pH,增加消化酶的活性,增强肠道消化吸收能力,从而提高动物生长性能[19];二是短链脂肪酸可促生长相关因子的表达,如生长激素(GH)和胰岛素样生长因子1(IGF1)[21]。本研究表明,添加水平为1 000 mg/kg的三丁酸甘油酯可增加黄颡鱼摄食量,显著提高黄颡鱼增重率和降低饲料系数,从而提高黄颡鱼生长性能。

3.2 三丁酸甘油酯对黄颡鱼血清生化指标的影响

动物机体中物质代谢、健康状况等可通过血清生化指标判断,血液中的葡萄糖是机体的重要能量来源,本研究中,随着三丁酸甘油酯添加水平的增加,血清中葡萄糖含量逐渐升高,2 000 mg/kg组达到最高值,在金头鲷[22-23](Sparus aurata)、斑鱼[16](Channa argus)得到相同的结果。血清总蛋白含量反映了机体蛋白质吸收代谢程度,也是维持血液渗透压重要成分。本试验条件下,血清总蛋白、白蛋白和球蛋白含量与对照组相比无显著差异,说明三丁酸甘油酯对黄颡鱼血清中蛋白质含量无显著影响。高剂量添加三丁酸甘油酯增加脂肪沉积[24],添加少量三丁酸甘油酯则可有效降低肝脏和血清脂肪含量,在高水平大豆油饲料中,添加4 g/kg三丁酸甘油酯可显著降低大黄鱼脂肪沉积,减少脂肪生成相关基因的表达[21]。而本研究发现,饲料中添加3种不同水平的三丁酸甘油酯对黄颡鱼脂肪沉积无显著影响,推测可能和鱼的种类和添加剂量偏低有关。谷草转氨酶和谷丙转氨酶是判断肝脏健康状况和蛋白质代谢的重要指标,谷丙转氨酶和谷草转氨酶正常情况下应存在于肝细胞内,当肝细胞损伤时,谷丙转氨酶和谷丙转氨酶才会进入血液。本研究表明,500、1 000、2 000 mg/kg组黄颡鱼血清谷草转氨酶活性低于对照组,但各组间无显著差异;各组间血清谷丙转氨酶活性无显著差异,说明饲料中添加三丁酸甘油酯对黄颡鱼肝脏功能无显著影响,在胡子鲶[25](Clarias fuscus)中研究结果与本试验相同。尿素氮是蛋白质代谢最终产物,也是判断机体蛋白质代谢能力强弱的关键指标。血清中尿素氮含量在1 000 mg/kg组为最高,但各组间无显著差异,说明三丁酸甘油酯对黄颡鱼蛋白质代谢无显著影响。

3.3 三丁酸甘油酯对黄颡鱼血清免疫功能的影响

溶菌酶由T细胞、B细胞、淋巴细胞、噬菌体等细胞产生,鱼血液中高水平的LZM与吞噬细胞释放LZM产量增加有关[26],本试验中,与对照组相比,500、1 000和2 000 mg/kg组的血清LZM活性显著升高,其中500 mg/kg组活性最高,随添加水平增多而呈下降趋势,推测饲料中添加三丁酸甘油酯可能激活吞噬细胞等产生LZM。同样,Xie等[7]发现在鲤鱼高植物蛋白质饲料中添加2 g/kg三丁酸甘油酯可显著提高血清LZM活性;Zhou等[10]在草鱼饲料中添加200~400 mg/kg的三丁酸甘油酯亦得到相同的结论。AKP是一种碱性磷酸水解酶,是甲壳类和鱼类重要的非特异性指标,也是反映水生动物健康状况的重要指标[27];C3是由肝细胞产生、参与体液免疫反应的大分子球白质,可以通过补体经典激活途径和旁路激活途径参与免疫应答,可直接溶解或中和病毒[28],本试验中,500和1 000 mg/kg组血清AKP活性显著升高,各组间血清C3含量无显著差异。IgM是B细胞分泌的一种参与体液免疫的抗体,IgM可与病原体结合发生非特异性免疫反应。本试验结果发现,与0 mg/kg组相比,500、1 000、2 000 mg/kg组血清IgM含量都有所增加,其中1 000 mg/kg组的血清IgM含量显著高于2 000 mg/kg组。本研究结果表明,饲料中添加适量的三丁酸甘油酯可增加黄颡鱼非特异性免疫。

3.4 三丁酸甘油酯对黄颡鱼血清抗氧化能力的影响

鱼类在正常或压力条件下会产生自由基和活性氧,为避免这些化合物对机体产生损伤,生物体可产生抗氧化酶如SOD、GSH-Px、CAT及MDA和T-AOC。T-AOC反映了机体整体抗氧化综合能力,SOD可清除机体产生的自由基,维持机体抗氧化能力和抗氧化平衡[29],本试验中,500 mg/kg组黄颡鱼血清T-AOC显著高于0 mg/kg组,但1 000、2 000 mg/kg组T-AOC逐渐降低,说明添加500 mg/kg三丁酸甘油酯可显著提高黄颡鱼综合抗氧化能力,过量添加三丁酸甘油酯可能对黄颡鱼抗氧化能力产生负面影响,500、1 000、2 000 mg/kg组黄颡鱼血清SOD的活性显著提高,1 000 mg/kg组血清SOD活性最高,表明饲料中添加适量的三丁酸甘油酯可显著增加黄颡鱼抗氧化能力,在草鱼[8,29]、团头鲂[5]上也有相同的结果。机体在应激或压力环境下易产生对细胞毒害作用的过氧化氢,而CAT能将过氧化氢分解为水和氧气,避免过氧化氢对细胞的损害,本试验中,2 000 mg/kg组血清CAT活性显著高于其他各组,表明饲料添加适量三丁酸甘油酯可提高CAT活性,增强机体抗氧化能力。三丁酸甘油酯可增加ATP的含量,激活磷脂酰肌醇3-激酶(PI3K)和蛋白激酶B(AKT)信号传导,进一步激活细胞内重要的多功能转录因子——乳红系衍生的核因子2相关因子(Nrf2)信号通路,提高抗氧化酶基因的表达,从而增强抗氧化酶活性[30]。本试验中,各组间的血清GSH-Px活性及GSH和MAD含量无显著差异,这与草鱼[8]、乌醴[15]、团头鲂[5]的研究结果不同,推测原因可能和鱼的种类和三丁酸甘油酯的添加水平有关,具体原因有待进一步研究。研究结果表明,饲料中添加三丁酸甘油酯有助于黄颡鱼提高抗氧化能力,不同添加水平对黄颡鱼抗氧化指标也会产生不同的影响。

4 结论

本研究表明,饲料中添加不同水平的三丁酸甘油酯可改善黄颡鱼的生长性能、饲料效率,增强非特异性免疫力和抗氧化能力,三丁酸甘油酯可能是鱼类潜在的免疫刺激剂,其最适宜添加水平为1 000 mg/kg。

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