鲤益生菌的筛选、生物学特性及作用机理的研究
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摘要
本研究从鲤(Cyprinus carpio)肠道及养殖水体分离筛选具有耐热和抑制病原菌功能的益生菌,并对其进行鉴定,在体外研究了部分益生菌对鲤前肠粘液的粘附特性及对病原菌粘附的抑制作用等生物学特性,并探讨了所筛选的枯草芽孢杆菌、肠球菌及光合细菌等3株益生菌对鲤肠道免疫机能、消化酶、肠道菌群结构与多样性及鲤肌肉品质、生长性能等的影响。各试验的研究结果如下:
试验一、试验二中,从养殖池、水族箱和健康鲤肠道等分离20株细菌,以及四川农业大学动物微生态研究中心提供的8株供试细菌共28株,通过耐热和体外拮抗试验筛选出5株细菌,经生化试验和细菌16S rDNA测序鉴定为2株芽孢杆菌(Bacillus)、2株肠球菌(Enterococci)和1株柠檬酸杆菌(Citrobacter)。进一步采用体外固定鲤前肠黏液蛋白,结合同位素~(32)P标记细菌并示踪的方法,研究来源于鲤肠道的肠球菌和柠檬酸杆菌以及鲤养殖水体中的芽孢杆菌对鲤前肠黏液的体外黏附活性,建立筛选鲤益生菌的方法。结果表明,5株细菌均能黏附到黏液体外模型,肠球菌的相对黏附率极显著高于芽孢杆菌与柠檬酸杆菌(P＜0.01),柠檬酸杆菌与芽孢杆菌的相对黏附率差异不显著(P＞0.05),而肠球菌L2的相对黏附率极显著高于肠球菌E2(P＜0.01)。结果证明,鱼源的肠球菌对鲤前肠黏液的黏附率高于异源的芽孢杆菌,而且不同种属的肠球菌在黏附能力上也存在差异。
试验三、试验四中,选择枯草芽孢杆菌制剂(B组,由Z2菌株制成),枯草芽孢杆菌和光合细菌(四川农业大学动物微生态研究中心提供)混合制剂(C组),枯草芽孢杆菌和肠球菌(由L2菌株制成)混合制剂(D组)3种益生菌组合,并将每一种制剂在同一浓度水平与对照组(A组)进行比较,分析试验组对鲤鱼免疫功能及鲤肠道消化酶的影响。结果表明:试验B组、C组中鲤的白细胞吞噬、杀菌活性、血清溶菌酶活性与对照组A差异显著(P＜0.05),而试验D组上述3项指标都高于对照组A,但效果没有试验B、C组好,而3个试验组与对照组中淋巴细胞数差异不显著(P＞0.05)。证明:单独使用枯草芽孢杆菌比枯草芽孢杆菌和光合细菌复合制剂,以及枯草芽孢杆菌和肠球菌复合制剂更能有效地提高鲤鱼的白细胞的吞噬活性、杀菌活性和血清溶菌酶活性。同时,结果还表明:试验后期B组淀粉酶比活显著提高(P＜0.05),脂肪酶、胰蛋白酶比活力极显著提高(P＜0.01)。C组、D组胰蛋白酶比活力极显著提高(P＜0.01),但对淀粉酶和脂肪酶比活力的影响差异不显著(P＞0.05),特别是D组与A组的淀粉酶比活力在各阶段差异均不显著(P＞0.05)。在整个试验中,B组对鲤肠道消化酶比活的提高效果最好。
试验五中,采用PCR-DGGE技术研究了不同组合的益生菌制剂对鲤肠道菌群结构与多样性的影响。结果表明:B组、C组和D组均能提高鲤肠道菌群的多样性。其中,试验B组鲤肠道菌群的多样性、优势菌群条带较对照组A明显提高,到42 d时条带增加量为9条;而试验C组鲤肠道菌群的多样性、优势菌群高于对照组A,到42 d时条带增加量为4条,但效果没有试验B组好,而D组的效果不明显。
试验六中,研究了不同益生菌组合对鲤肌肉氨基酸含量、肉质及生长性能的影响。结果表明,在生长方面,与对照组相比,试验B、C、D组试验组鲤的生长性能指标均高于对照组。其中B组的丰满度、绝对生长率、相对生长率、瞬时生长率、生长比速等均达到同阶段的峰值,与对照组相比,试验B、C组绝对生长率与瞬时生长率显著提高(P＜0.05);在肌肉品质方面,与对照组相比,添加益生菌的3组试验组(B、C、D组),鲤肌肉粗脂肪含量、失水率极显著降低(P＜0.01);肌肉粗蛋白含量显著增加(P＜0.05);其中,试验B组显著程度最高。上述结果表明,3种益生菌组合对鲤均有一定的促生长作用,并对肌肉品质有明显的改善作用,其中,又以枯草芽孢杆菌菌剂单独使用效果最明显。
综上所述,在鲤肠道、养殖水体广泛分布着益生菌。本研究通过普通培养和选择培养的方法,并通过耐热及体外抑制试验,筛选出枯草芽孢杆菌Z2、肠球菌L2等2株益生菌对致病性大肠杆菌、气单胞杆菌等有害菌有较强的抑制活性,并首次采用体外固定鲤前肠黏液,结合同位素~(32)p示踪的方法,对来源于鲤肠道的肠球菌和柠檬酸杆菌以及养殖水体的芽孢杆菌的细菌表面凝集素黏液蛋白受体的化学组成、其对病源菌附着鲤前肠黏液的影响等进行了研究,进一步探讨了益生菌的抑菌机理。同时本研究还首次探讨了不同益生菌组合对鲤免疫机理、肠道消化道酶活、肠道菌群结构与多样性、鲤肌肉品质及生产性能等的影响,结果表明,3种益生菌组合均可提高鲤免疫机能、消化道酶活性;增加鲤肠道菌群微生物多样性;改善鲤肌肉品质;提高生长性能。其中又以单株枯草芽孢杆菌制剂对以上指标的影响最大,作用效果最明显。该研究项目筛选的枯草芽孢杆(Z2)有望开发成鲤和其它鱼类养殖应用的益生菌。
In this study,probiotics with the ability of heat tolerant and to inhibit pathogenic bacterium were screened from intestinal tract of Cyprinus carpio and breeding water. Followed identification,some probiotics strains were tested for ability to adhere to the foregut mucus of Cyprinus carpio and against binding of pathogenic bacteria.Mechanism of three isolates,which are Bacillus subtilis,Enterococcus and Photosynthetic Bacteria effect on immune function,digestive enzymes,and flora diversity of Cyprinus carpio intestinal tract, muscle quality,and growth performance of Cyprinus carpio were discussed.The results of study are as follows:
In the first and second experiments,twenty strains isolated from breeding pools, aquariums and intestinal tracts of healthy Cyprinus carpios,and eight strains provided by Animal Micro-ecology Research Center of Sichuan Agricultural University were tested by heat-treatment and inhibitory experiment.There were five strains were screened according to the results,and then identified as two strains of Bacillus,two strains of Enterococci,and one strain of Citrobacter by 16S rDNA sequencing.Further,the method of immobilizing Cyprinus carpio foregut mucus albumen in vitro,combined with labeling bacteria with isotope ~(32)P was conducted to study the adhesion activity of Citrobacter and Enterococci strains from Cyprinus carpio intestinal tract,and Bacillus strains from breeding water.The method of screening probiotics from Cyprinus carpio was established in this study.The results indicated that all five strains can adhere to the mucus of in vitro model,and while the adhesion ratio of Enterococci strains was extremely higher than Citrobacter and Bacillus strains(P＜0.01),the difference between Citrobacter and Bacillus strains was not significant (P＞0.05),and the adhesion ratio of Enterococci strain L2 was extremely higher than Enterococci strain E2(P＜0.01).The results suggested that the Enterococci strains isolated from fish are easier to adhere to Cyprinus carpio foregut mucus than the heterogenous Bacillus strains,and the adhesion activity depended on different strain genus.
In the third and forth experiments,in order to investigate the effect of probiotics on immune function and intestinal digestive enzymes of Cyprinus carpio,four treatment groups fed with different probiotics preparations were conducted,including Group A(control group), Group B(fed with Bacillus subtilis strain Z2),Group C(fed with mixing preparation of Bacillus subtilis and Photosynthetic bacteria),Group D(fed with mixing preparation of Bacillus subtilis and Enterococci strain L2),and each preparation was compared with Group A within the same level.The results showed that phagocytosis and bactericidal activity of leukocyte and serum enzymatic activity of Group B and C was significantly different from Group A(P＜0.05),and the performance of Group D was not as well as Group B and C. There was no significant difference in amount of lymphocyte among Group A,B and C(P＞0.05).The results indicated that Bacillus subtilis rather than compound of Bacillus subtilis and Photosynthetic bacteria,Bacillus subtilis and Enterococci efficiently improved the phagocytosis and bactericida activity of leukocyte,and serum lysozyme activity.Further, amylase specific activity of Group B was enhanced significantly in the later stage of experiment(P＜0.05),while lipase and trypsin specific activity were extremely improved(P＜0.01).Trypsin specific activity of Group C and D was extremely enhanced,but the change of amylase and lipase were not as well as trypsin,especially the difference of amylase was not significant in every stage of Group D and A(P＞0.05).In this study,Group B showed the best performance to improve the activity of intestinal digestive enzymes.
In the fifth experiment,Polymerase Chain Reaction-Denaturing Gradient Gel Electrophoresis(PCR-DGGE) method was used to investigate the effect of different preparation of probiotics on intestinal tract flora construction and diversity of Cyprinus carpio. The results indicated that Group B,C and D can improve the intestinal tract flora diversity. Flora diversity and belts of dominance strains of Group B were significantly increased compared with Group A,and the belts added with nine at 42 days,meanwhile,Group C showed less efficiency than Group B,and the belts increased with four at 42 days.The effect of Group D was not significant.
In the sixth experiment,the effect of different probiotics preparations on amino acid content,muscle quality and growth performance was studied.The results showed that growth performance index of Group B,C and D were higher than control group.The fullness, absolute growth rate,relative growth rate,transient growth rate and growth specific speed of Group B reached the peak at every stage.Absolute growth rate and transient growth rate of Group B and C were significantly improved(P＜0.05).Compared with the control group, muscle crude fat content and dehydration rate of treatment groups fed with probiotics were decreased significantly(P＜0.01),and muscle crude protein was increased significantly(P＜0.05).The results indicated that three strains of probiotics in this study can promote the growth performance,and improve the muscle quality,and Bacillus subtilis has the best effect.
In conclusion,probiotics are distributed widely in intestinal tracts of Cyprinus carpio and breeding water.Bacillus subtilis strain Z2 and Enterococci strain L2 screened by common and selecting cultivation method,heat-treatment,and inhibitory experiment in vitro showed strong inhibition activity to enteropathogenic E.coli and Aeromonas.For the first time,the method of immobilizing Cyprinus carpio foregut mucus albumen in vitro,combined with labeling bacteria with isotope ~(32)P was used to study chemical composition of the lectin receptors on bacterial surface,and ability of pathogenic bacteria to adhere to mucus.Further, the mechanism of bacteriostasis of prebiotics was discussed.Three different prebiotics combination groups enhanced immune function and digestive enzymes activity,increased intestinal tract flora diversity,and improved muscle quality and growth performance of Cyprinus carpio.Bacillus subtilis strain Z2 has the potential to utilize as prebiotics in breeding Cyprinus carpio and other fish.
试验一、试验二中,从养殖池、水族箱和健康鲤肠道等分离20株细菌,以及四川农业大学动物微生态研究中心提供的8株供试细菌共28株,通过耐热和体外拮抗试验筛选出5株细菌,经生化试验和细菌16S rDNA测序鉴定为2株芽孢杆菌(Bacillus)、2株肠球菌(Enterococci)和1株柠檬酸杆菌(Citrobacter)。进一步采用体外固定鲤前肠黏液蛋白,结合同位素~(32)P标记细菌并示踪的方法,研究来源于鲤肠道的肠球菌和柠檬酸杆菌以及鲤养殖水体中的芽孢杆菌对鲤前肠黏液的体外黏附活性,建立筛选鲤益生菌的方法。结果表明,5株细菌均能黏附到黏液体外模型,肠球菌的相对黏附率极显著高于芽孢杆菌与柠檬酸杆菌(P＜0.01),柠檬酸杆菌与芽孢杆菌的相对黏附率差异不显著(P＞0.05),而肠球菌L2的相对黏附率极显著高于肠球菌E2(P＜0.01)。结果证明,鱼源的肠球菌对鲤前肠黏液的黏附率高于异源的芽孢杆菌,而且不同种属的肠球菌在黏附能力上也存在差异。
试验三、试验四中,选择枯草芽孢杆菌制剂(B组,由Z2菌株制成),枯草芽孢杆菌和光合细菌(四川农业大学动物微生态研究中心提供)混合制剂(C组),枯草芽孢杆菌和肠球菌(由L2菌株制成)混合制剂(D组)3种益生菌组合,并将每一种制剂在同一浓度水平与对照组(A组)进行比较,分析试验组对鲤鱼免疫功能及鲤肠道消化酶的影响。结果表明:试验B组、C组中鲤的白细胞吞噬、杀菌活性、血清溶菌酶活性与对照组A差异显著(P＜0.05),而试验D组上述3项指标都高于对照组A,但效果没有试验B、C组好,而3个试验组与对照组中淋巴细胞数差异不显著(P＞0.05)。证明:单独使用枯草芽孢杆菌比枯草芽孢杆菌和光合细菌复合制剂,以及枯草芽孢杆菌和肠球菌复合制剂更能有效地提高鲤鱼的白细胞的吞噬活性、杀菌活性和血清溶菌酶活性。同时,结果还表明:试验后期B组淀粉酶比活显著提高(P＜0.05),脂肪酶、胰蛋白酶比活力极显著提高(P＜0.01)。C组、D组胰蛋白酶比活力极显著提高(P＜0.01),但对淀粉酶和脂肪酶比活力的影响差异不显著(P＞0.05),特别是D组与A组的淀粉酶比活力在各阶段差异均不显著(P＞0.05)。在整个试验中,B组对鲤肠道消化酶比活的提高效果最好。
试验五中,采用PCR-DGGE技术研究了不同组合的益生菌制剂对鲤肠道菌群结构与多样性的影响。结果表明:B组、C组和D组均能提高鲤肠道菌群的多样性。其中,试验B组鲤肠道菌群的多样性、优势菌群条带较对照组A明显提高,到42 d时条带增加量为9条;而试验C组鲤肠道菌群的多样性、优势菌群高于对照组A,到42 d时条带增加量为4条,但效果没有试验B组好,而D组的效果不明显。
试验六中,研究了不同益生菌组合对鲤肌肉氨基酸含量、肉质及生长性能的影响。结果表明,在生长方面,与对照组相比,试验B、C、D组试验组鲤的生长性能指标均高于对照组。其中B组的丰满度、绝对生长率、相对生长率、瞬时生长率、生长比速等均达到同阶段的峰值,与对照组相比,试验B、C组绝对生长率与瞬时生长率显著提高(P＜0.05);在肌肉品质方面,与对照组相比,添加益生菌的3组试验组(B、C、D组),鲤肌肉粗脂肪含量、失水率极显著降低(P＜0.01);肌肉粗蛋白含量显著增加(P＜0.05);其中,试验B组显著程度最高。上述结果表明,3种益生菌组合对鲤均有一定的促生长作用,并对肌肉品质有明显的改善作用,其中,又以枯草芽孢杆菌菌剂单独使用效果最明显。
综上所述,在鲤肠道、养殖水体广泛分布着益生菌。本研究通过普通培养和选择培养的方法,并通过耐热及体外抑制试验,筛选出枯草芽孢杆菌Z2、肠球菌L2等2株益生菌对致病性大肠杆菌、气单胞杆菌等有害菌有较强的抑制活性,并首次采用体外固定鲤前肠黏液,结合同位素~(32)p示踪的方法,对来源于鲤肠道的肠球菌和柠檬酸杆菌以及养殖水体的芽孢杆菌的细菌表面凝集素黏液蛋白受体的化学组成、其对病源菌附着鲤前肠黏液的影响等进行了研究,进一步探讨了益生菌的抑菌机理。同时本研究还首次探讨了不同益生菌组合对鲤免疫机理、肠道消化道酶活、肠道菌群结构与多样性、鲤肌肉品质及生产性能等的影响,结果表明,3种益生菌组合均可提高鲤免疫机能、消化道酶活性;增加鲤肠道菌群微生物多样性;改善鲤肌肉品质;提高生长性能。其中又以单株枯草芽孢杆菌制剂对以上指标的影响最大,作用效果最明显。该研究项目筛选的枯草芽孢杆(Z2)有望开发成鲤和其它鱼类养殖应用的益生菌。
In this study,probiotics with the ability of heat tolerant and to inhibit pathogenic bacterium were screened from intestinal tract of Cyprinus carpio and breeding water. Followed identification,some probiotics strains were tested for ability to adhere to the foregut mucus of Cyprinus carpio and against binding of pathogenic bacteria.Mechanism of three isolates,which are Bacillus subtilis,Enterococcus and Photosynthetic Bacteria effect on immune function,digestive enzymes,and flora diversity of Cyprinus carpio intestinal tract, muscle quality,and growth performance of Cyprinus carpio were discussed.The results of study are as follows:
In the first and second experiments,twenty strains isolated from breeding pools, aquariums and intestinal tracts of healthy Cyprinus carpios,and eight strains provided by Animal Micro-ecology Research Center of Sichuan Agricultural University were tested by heat-treatment and inhibitory experiment.There were five strains were screened according to the results,and then identified as two strains of Bacillus,two strains of Enterococci,and one strain of Citrobacter by 16S rDNA sequencing.Further,the method of immobilizing Cyprinus carpio foregut mucus albumen in vitro,combined with labeling bacteria with isotope ~(32)P was conducted to study the adhesion activity of Citrobacter and Enterococci strains from Cyprinus carpio intestinal tract,and Bacillus strains from breeding water.The method of screening probiotics from Cyprinus carpio was established in this study.The results indicated that all five strains can adhere to the mucus of in vitro model,and while the adhesion ratio of Enterococci strains was extremely higher than Citrobacter and Bacillus strains(P＜0.01),the difference between Citrobacter and Bacillus strains was not significant (P＞0.05),and the adhesion ratio of Enterococci strain L2 was extremely higher than Enterococci strain E2(P＜0.01).The results suggested that the Enterococci strains isolated from fish are easier to adhere to Cyprinus carpio foregut mucus than the heterogenous Bacillus strains,and the adhesion activity depended on different strain genus.
In the third and forth experiments,in order to investigate the effect of probiotics on immune function and intestinal digestive enzymes of Cyprinus carpio,four treatment groups fed with different probiotics preparations were conducted,including Group A(control group), Group B(fed with Bacillus subtilis strain Z2),Group C(fed with mixing preparation of Bacillus subtilis and Photosynthetic bacteria),Group D(fed with mixing preparation of Bacillus subtilis and Enterococci strain L2),and each preparation was compared with Group A within the same level.The results showed that phagocytosis and bactericidal activity of leukocyte and serum enzymatic activity of Group B and C was significantly different from Group A(P＜0.05),and the performance of Group D was not as well as Group B and C. There was no significant difference in amount of lymphocyte among Group A,B and C(P＞0.05).The results indicated that Bacillus subtilis rather than compound of Bacillus subtilis and Photosynthetic bacteria,Bacillus subtilis and Enterococci efficiently improved the phagocytosis and bactericida activity of leukocyte,and serum lysozyme activity.Further, amylase specific activity of Group B was enhanced significantly in the later stage of experiment(P＜0.05),while lipase and trypsin specific activity were extremely improved(P＜0.01).Trypsin specific activity of Group C and D was extremely enhanced,but the change of amylase and lipase were not as well as trypsin,especially the difference of amylase was not significant in every stage of Group D and A(P＞0.05).In this study,Group B showed the best performance to improve the activity of intestinal digestive enzymes.
In the fifth experiment,Polymerase Chain Reaction-Denaturing Gradient Gel Electrophoresis(PCR-DGGE) method was used to investigate the effect of different preparation of probiotics on intestinal tract flora construction and diversity of Cyprinus carpio. The results indicated that Group B,C and D can improve the intestinal tract flora diversity. Flora diversity and belts of dominance strains of Group B were significantly increased compared with Group A,and the belts added with nine at 42 days,meanwhile,Group C showed less efficiency than Group B,and the belts increased with four at 42 days.The effect of Group D was not significant.
In the sixth experiment,the effect of different probiotics preparations on amino acid content,muscle quality and growth performance was studied.The results showed that growth performance index of Group B,C and D were higher than control group.The fullness, absolute growth rate,relative growth rate,transient growth rate and growth specific speed of Group B reached the peak at every stage.Absolute growth rate and transient growth rate of Group B and C were significantly improved(P＜0.05).Compared with the control group, muscle crude fat content and dehydration rate of treatment groups fed with probiotics were decreased significantly(P＜0.01),and muscle crude protein was increased significantly(P＜0.05).The results indicated that three strains of probiotics in this study can promote the growth performance,and improve the muscle quality,and Bacillus subtilis has the best effect.
In conclusion,probiotics are distributed widely in intestinal tracts of Cyprinus carpio and breeding water.Bacillus subtilis strain Z2 and Enterococci strain L2 screened by common and selecting cultivation method,heat-treatment,and inhibitory experiment in vitro showed strong inhibition activity to enteropathogenic E.coli and Aeromonas.For the first time,the method of immobilizing Cyprinus carpio foregut mucus albumen in vitro,combined with labeling bacteria with isotope ~(32)P was used to study chemical composition of the lectin receptors on bacterial surface,and ability of pathogenic bacteria to adhere to mucus.Further, the mechanism of bacteriostasis of prebiotics was discussed.Three different prebiotics combination groups enhanced immune function and digestive enzymes activity,increased intestinal tract flora diversity,and improved muscle quality and growth performance of Cyprinus carpio.Bacillus subtilis strain Z2 has the potential to utilize as prebiotics in breeding Cyprinus carpio and other fish.
引文
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