EFFECTS OF DIETARY MULBERRY LEAF EXTRACT AND 1-DEOXYNOJIRIMYCIN ON GROWTH, DIGESTION AND IMMUNITY CAPACITY, AND INTESTINAL MICROORGANISM OF CHINESE GIANT SALAMANDER (ANDRIAS DAVIDIANUS)
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摘要: 为探究桑叶物提取物(MLE)和单体1-脱氧野尻霉素(DNJ)对大鲵(Andrias davidianus)生长、消化、免疫能力和肠道菌群影响的差异, 研究以基础饲料为对照组, 分别添加0.75%的MLE和0.04‰的DNJ制成3种等氮等脂的饲料, 饲喂初始均重为(47.81±0.23) g的大鲵90d。结果: MLE组和DNJ组增重率和饲料效率显著提高(P<0.05); 肠道脂肪酶和胰蛋白酶活力显著提高(P<0.05), 肠道丙二醛、血浆内毒素显著降低(P<0.05), 肠黏膜绒毛数量显著增加(P<0.05), 肠黏膜上皮细胞间紧密连接更加紧密; 与对照组相比, MLE组雷帕霉素靶蛋白(Target of rapamycin, TOR)、白细胞介素-10 (Interleukin, IL-10)、多聚免疫球蛋白受体(Polymeric immunoglobulin receptor, pIgR)、类Toll受体2(Toll-like receptor 2, TLR2)和髓样分化初级反应基因88 (Myeloid differentiation primary response gene88, MyD88) mRNA表达水平显著上调(P<0.05), DNJ组pIgR mRNA表达水平显著上调(P<0.05); 肠道菌群中瘤胃球菌科细菌丰度降低, Romboutsia丰度增加, 肠道微生物多样性较对照组提高(P<0.05)。与DNJ组相比, MLE组和DNJ组增重率和饲料效率无显著差异(P>0.05), MLE组胃蛋白酶、H+-K+-ATP酶和Na+-K+-ATP酶活力显著提高, 肠道总抗氧化能力显著提高(P<0.05), 肠黏膜上皮细胞紧密连接程度提高, 肠绒毛高度显著增加(P<0.05), 血浆内毒素含量显著降低。结果表明, 在大鲵配合饲料中添加MLE和DNJ均能不同程度改善大鲵胃和肠的消化吸收功能, 提高生长性能、肠道抗氧化能力和免疫能力, 提高肠道微生物多样性, 促进肠道发育; 综合胃和肠结构和功能, 复合MLE较单体DNJ效果更优。Abstract: This study explored the effects of feeding basal diet, mulberry leaf extract diet (containing 0.75% MLE) and 1-deoxynojirimycin diet (containing 0.04‰ DNJ) on growth, digestion and immunity capacity, and intestinal microorganism in giant salamanders for a 90-day trail. Giant salamanders with an initial body weight of (47.81±0.23) g were randomly divided into 3 groups for these 3 diets. Compared with the control diet, the weight gain rate and feed efficiency in MLE and DNJ groups were significantly improved (P<0.05); lipase and trypsin activities were also significantly increased (P<0.05). The contents of endotoxin in plasma acid and malondialdehyde in intestine in MLE and DNJ groups were significantly lower than those in control group (P<0.05). Compared with control group, the number of intestinal villi in MLE and DNJ groups were increased, and the tight junction between intestinal mucosa cells in MLE and DNJ groups were closer. Compared with the control group, the mRNA expression levels of target of rapamycin (TOR), interleukin-10 (IL-10), polymeric immunoglobulin receptor (pIgR), toll-like receptor-2 (TLR2) and myeloid differentiation primary response gene88 (MyD88) in MLE group were significantly increased (P<0.05), and the mRNA expression level of pIgR in the DNJ group was significantly increased (P<0.05). The diversity of intestinal bacteria in MLE and DNJ group was higher than that in control group. There was no significant difference in weight gain rate and feed efficiency among 3 groups (P>0.05); MLE group significantly increased the activities of pepsin, H+-K+-ATPase, Na+-K+-ATPase and the total intestinal antioxidant capacity compared with the control group (P<0.05). In addition, MLE group had closer tight junction between intestinal mucosa cells, higher intestinal villi height (P<0.05), and lower plasma endotoxin content (P<0.05), compared with DNJ group. In summary, MLE and DNJ can improve growth performance, digestion and absorption functions, intestinal antioxidant and immune capabilities, promote the growth of beneficial bacteria in the intestine, increase the diversity of intestinal microorganisms, and promote the intestinal development of the giant salamander. Combined the structure and function of the stomach and intestines, MLE is more effective than DNJ.
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图 1 MLE与DNJ对大鲵前肠形态结构的影响(HE染色，40×)
a. 对照组；b. MLE组；c. DNJ组
Figure 1. Effects of MLE and DNJ on intestinal morphology and structure of Andrias davidianus
a. control group; b. MLE group; c. DNJ group
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图 2 MLE与DNJ对大鲵肠黏膜细胞紧密连接结构的影响
A1. 对照组(4000×)；B1. MLE组(4000×)；C1. DNJ组(4000×)；A2. 对照组(8000×)；B2. MLE组(8000×)；C2. DNJ组(8000×)
Figure 2. Effects of MLE and DNJ on tight junctions in intestinal mucosa of Andrias davidianus
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图 3 MLE和DNJ对大鲵肠道免疫相关基因相对表达量的影响
Figure 3. Effects of MLE and DNJ on intestinal immune related gene expression of Andrias davidianus
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图 4 MLE和DNJ对大鲵肠道菌群的结构分析
a. 不同组OUT分析；b. 样品UPGMA聚类树；c. 门水平样品序列分类；d. 属水平样品序列分类
Figure 4. Effects of MLE and DNJ on intestinal flora structure of Andrias davidianus
表 1 实验饲料配方及主要营养成分(风干基础)
Table 1 Formulation and nutrient composition of diets (air-dry basis)
项目Item 组别Group 对照组Control MLE组 DNJ组 鱼粉Fish meal (%) 15 15 15 肉粉Meat meal (%) 3 3 3 酪蛋白Casein (%) 6 6 6 明胶Gelatin (%) 6 6 6 复合蛋白粉1)Compound protein powder (%) 17 17 17 面粉Wheat flour (%) 28 28 28 α-预糊化淀粉α-pregelatinized starch (%) 11 11 11 大豆油Soybean oil (%) 4.8 4.8 4.8 预混料Premix2) (%) 7.7 7.7 7.7 桑叶提取物MLE (%) 0 0.75 0 1-脱氧野尻霉素DNJ(‰) 0 0 0.04 膨润土Bentonite (%) 1.5 0.75 1.5 合计Total (%) 100 100 100 营养组成(干基实测)Nutrient composition (measured dry basis, %) 粗蛋白Crude protein 55.36 55.60 55.41 粗脂肪Crude lipid 8.70 8.81 8.73 粗灰分Crude ash 10.30 10.15 10.12 粗纤维Crude fiber 0.91 0.95 0.92 注：1)复合蛋白粉成分保密；2)预混料含多维、多矿、功能性添加剂和载体，多维、多矿的组成参照肉食性鱼类Note: 1)The ingredients of compound protein powder are confidential; 2)Premix contains multi-vitamins, complex mineral, functional additives and carrier, multi-vitamin and complex mineral refer to carnivorous fishes 
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表 2 免疫相关基因及内参基因引物序列
Table 2 Primers used for quantitative real-time PCR
基因Gene 引物序列Sequences (5′—3′) 长度Length (bp) EF-1α GGACAGACCCGTGAACATGC
CTTCCTTAGTGATCTCCTCGTAGC130 IL-8 TGCAGACAGACGCACTTCAA
CACAGCCATCAGTCACCCTT112 IL-10 GCCGCGTGCATCTATTTTGG
CAGCAACCTGGAGAACCTCA104 TOR CGCTTCACCTCGAACTCCAC
ATCTCCTGCCCTCCAGTGAC109 pIgR CGCTTGGAGTCACTGAACCT
AGCCTCCTCCAAAGCATCAC102 TLR2 TGGACAATGTCACCCCTGTG
TCATCCGCTGAAGAAACCCC150 MyD88 TCCCTGCCATTTCTTGGGAC
CTCACCGACCTGAAACCGAA130 IRAK4 TTCTGGGTCACTGTCTTGCG
ATCAAAGGAGGCATCTGGGC112 注: EF-1α. 延伸因子-1α elongation factor-1α; IL-8. 白细胞介素-8 interleukin-8; IL-10. 白细胞介素-10 interleukin-10; TOR. 雷帕霉素靶蛋白target of rapamycin; TLR2.类Toll受体2 toll-like receptor 2; IRAK4.白细胞介素-1受体相关激酶4 interleukin-1 receptor-associated kinase 4; MyD88. 髓样分化初级反应基因88 myeloid differentiation primary response gene 88; pIgR. 多聚免疫球蛋白受体polymeric immunoglobulin receptor
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表 3 MLE和DNJ对大鲵生长性能及饲料效率的影响
Table 3 Effects of MLE and DNJ on growth performance and feed utilization of Andrias davidianus
项目Item 组别Group 对照组Control MLE组 DNJ组 终末鲵尾均重FBW (g) 76.92±4.31a 88.13±1.82b 90.15±3.11b 增重率WGR (%) 60.8±9.3a 84.5±2.9b 88.5±6.1b 平均日增重ADG (g/d) 0.32±0.05a 0.45±0.02b 0.47±0.04b 摄食量FI (g) 119.73±11.36a 138.79±3.57b 137.29±10.36ab 饲料效率FE (%) 145.3±8.3a 174.5±7.2b 185.1±6.1b
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表 4 MLE和DNJ对大鲵胃功能的影响
Table 4 Effects of MLE and DNJ on gastric function of Andrias davidianus (U/mg)
项目Item 组别Group 对照组Control MLE组 DNJ组 胃蛋白酶Pepsin 14.6±0.6a 23.0±1.9b 15.9±0.4a H+-K+-ATPase 4.14±0.14a 4.65±0.12b 4.25±0.15a 碳酸酐酶CA 191±9a 212±12b 208±8b
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表 5 MLE和DNJ对大鲵肠道消化吸收功能的影响
Table 5 Effects of MLE and DNJ on intestinal digestion and absorption function of Andrias davidianus (U/mg)
项目Item 组别Group 对照组Control MLE组 DNJ组 胰蛋白酶Trypsin 155±5a 263±6c 247±9b 脂肪酶Lipase 742±48a 858±35b 835±48b 淀粉酶Amylase 0.212±0.008b 0.168±0.007a 0.159±0.007a Na+-K+-ATPase 3.16±0.13a 3.59±0.09b 3.14±0.12a
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表 6 MLE和DNJ对大鲵肠黏膜通透性影响的比较
Table 6 Effects of MLE and DNJ on intestinal permeability of Andrias davidianus
项目Item 组别Group 对照组Control MLE组 DNJ组 内毒素ET (EU/mL) 64.3±1.6c 51.7±2.6a 61.9±1.9b D-乳酸D-LA (μmol/L) 241±14b 183±16a 209±23ab
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表 7 MLE与DNJ对大鲵肠道形态结构的影响
Table 7 Effects of MLE and DNJ on intestinal morphology and structure of Andrias davidianus
项目Item 组别Group 对照组Control MLE组 DNJ组 绒毛数量
Villus number12.1±0.9a 14.0±1.3b 13.5±0.5b 绒毛高度
Villus height (μm)350.3±70.2a 470.3±52.4b 319.7±27.4a 空腔率*
Cavity rate (%)22.4±3.2c 16.3±2.8a 19.8±3.6b 注: *空腔率(%)=肠道横切空腔面积/肠道横切面积×100Note: *Cavity rate(%)=Scavity of intestine/Sintestine×100
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表 8 MLE与DNJ对大鲵肠道抗氧化能力的影响
Table 8 Effects of MLE and DNJ on intestinal antioxidant of Andrias davidianus
项目Item 组别Group 对照组Control MLE组 DNJ组 总抗氧化能力T-AOC (U/mg) 4.28±0.09a 4.72±0.09b 4.45±0.11a 总超氧化物歧化酶T-SOD (U/mg) 48.6±1.2 52.9±1.5 50.6±2.8 丙二醛MDA (nmol/mg) 2.76±0.09b 2.15±0.10a 2.38±0.20a
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表 9 肠道菌群Alpha多样性指数
Table 9 Alpha diversity indices of samples
样品Simple Alpha多样性指数 ACE Chao1 香农指数Shannon 辛普森指数Simpson 覆盖率Coverage (%) 对照组Control 189.8 191.4 2.63 0.201 99.98 MLE组 204.2 205.3 2.82 0.152 99.98 DNJ组 228.1 231.2 2.48 0.244 99.95 P-valve 0.014 0.017 0.254 0.377 99.90 注: 每组数值分别为每组三个样品的均值Note: The values are the mean values of three samples in each group
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