鼠李糖乳杆菌影响仔猪肠道屏障功能的研究

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英文题名：Effects of Lactobacillus Rhamnosus on Intestinal Barrier in Piglets 作者：崔志文 论文级别：博士 学科专业名称：动物营养与饲料科学 中文关键词：鼠李糖乳杆菌LR6001 ; 肠道屏障 ; 细胞因子 ; 免疫调节 ; TLR ; 仔猪 ; IPEC-J2 英文关键词：Lactobacillus rhamnosus LR6001 ; intestinal barrier ; piglet ; cytokine ; microbial flora ; immunoregulation ; TLR ; IPEC-J2 学位年度：2013 导师：李卫芬 学科代码：090502 学位授予单位：浙江大学 论文提交日期：2013-06-01 答辩委员会主席：刘建新

摘要

鼠李糖乳杆菌(Lactobacillus rhamnosus)是一种被广泛应用的益生菌,因其具有改善宿主肠道健康、促进生长、降低腹泻及免疫调节等作用而备受关注。但有关鼠李糖乳杆菌对仔猪肠道屏障功能的影响及其机理研究较少。本论文通过体内和体外试验研究鼠李糖乳杆菌LR6001对仔猪肠道屏障功能的影响及可能的作用机理。主要的研究内容和结果如下：
     试验1：鼠李糖乳杆菌LR6001对哺乳仔猪肠黏膜屏障的影响及机理研究。随机选取6窝杜长大外三元初生仔猪24头,分为2个组,每组3窝。在仔猪出生后开始吮吸母乳前,经口灌服10%的灭菌脱脂乳溶液(对照组)或含鼠李糖乳杆菌LR6001(5-6×108CFU/mL)的10%灭菌脱脂乳溶液(鼠李糖乳杆菌组),灌服量为2mL/(头·天),之后每2天灌服1次,共3次。由母猪按常规哺乳至25日龄,母猪和乳猪饲料均不添加抗生素,并在断奶前屠宰取样。结果显示,与对照组相比,鼠李糖乳杆菌组哺乳仔猪的日增重提高14.68%(P<0.05),腹泻率下降30.73%(P<0.05)；肠道菌群多样性增加,厚壁菌的比例显著增加,拟杆菌、变形菌和梭杆菌的比例都明显减少,显著降低了盲肠内容物中肠杆菌的数量(P<0.01)；空肠绒毛高度：隐窝深度降低了10.01%(P<0.05),但绒毛长度差异不显著(P>0.05),隐窝深度的比值增加了20.13%(P<0.05)；空肠黏膜OCLN基因表达下调了48.12%(P<0.05),但ZO-1基因表达无显著差异(P>0.05)；空肠黏膜的pBD-1和PMAP-37二种抗菌肽基因表达显著上调(P<0.05),LYS基因表达极显著下调(P<0.01),MUC2基因表达和胃肠道pH值差异不显著(P>0.05)；回肠黏膜DAO活性和ITF含量分别增加了375.15%(P<0.01)和100.44%(P<0.01),空肠黏膜DAO活性和ITF含量分别降低了60.51%(P<0.01)和33.98%(P<0.01),表明小肠完整性得到改善；空肠黏膜麦芽糖酶活性提高41.29%(P<0.05),回肠黏膜SIgA和C3的含量分别增加了25.70%(P<0.05)和37.54%(P<0.05),空肠黏膜IgG、C3和SIgA的含量和回肠黏膜IgG含量略有提高(P>0.05)；空肠黏膜的抗炎因子TGF-β1、IL-10、促炎因子IFN-Y和回肠黏膜的抗炎因子TGF-β1分泌极显著增加(P<0.01),空肠和回肠黏膜的促炎因子IL-1、IL-6、IL-12和TNF-α的分泌都极显著降低(P<0.01),趋化因子IL-8也显著降低(P<0.05),空肠黏膜的促炎因子TNF-α、IFN-γ、趋化因子MCP-1和抗炎因子TGF-β1的基因表达都显著上调(P<0.01),说明空肠黏膜免疫功能得到提高；空肠黏膜的信号传导途径相关分子MD-2、TLR2、 TLR9和TRAF6基因表达分别上调了21.29%(P<0.05)、230.77%(P<0.01)、107.34%(P<0.01)和81.92%(P<0.05)。
     试验2：鼠李糖乳杆菌LR6001对断奶仔猪小肠黏膜屏障的影响。试验1的仔猪于25日龄断奶,之后的一周内(至32日龄)继续饲喂教槽料,屠宰取样。结果表明,与对照组相比,鼠李糖乳杆菌组断奶仔猪腹泻率降低15.32%(P>0.05),日增重略有提高但差异不显著(P>0.05)；肠道菌群多样性增加,厚壁菌数量显著增加,拟杆菌和变形菌的数量明显减少,肠道菌群结构得到改善；空肠黏膜结构形态比较完整,绒毛高度与隐窝深度的比值、绒毛长度分别极显著增加了34.93%(P<0.01)和13.92%(P<0.01),隐窝深度降低了14.53%(P<0.05)；胃的pH值显著下降；空肠黏膜的DAO含量下降21.77%(P<0.01),而ITF和血清内毒素的含量没有显著变化(P>0.05)；空肠黏膜的乳糖酶活性下降13.60%(P<0.05),而蔗糖酶和麦芽糖酶的活性无显著差异(P>0.05)；空肠黏膜sIgA的含量增加50%(P<0.01),IgG和C3含量则差异不显著(P>0.05)；空肠黏膜抗炎因子TGF-β1和促炎因子TNF-α含量都极显著提高(P<0.01),促炎因子IL-1β、IL-6、IL-12和IFN-y的水平则极显著降低(P<0.01),趋化因子IL-8的含量显著降低(P<0.05),抗炎因子IL-10的分泌量略有提高但差异不显著(P>0.05)。
     试验3：鼠李糖乳杆菌LR6001对猪肠上皮细胞IEPC-J2TLR基因表达影响的研究。将培养的IEPC-J2细胞分为4组,对照组(CK)、处理组I(T6)、处理组II(T7)和处理组III(T8)分别添加鼠李糖乳杆菌(约0、106、107、108CFU/mL)。IEPC-J2细胞培养3h时收集细胞,采用实时荧光定量PCR方法分别测定TLR2、TLR4、TLR6、TLR9、TLR10的基因表达量。结果发现,与对照组相比,T6组TLR基因表达无显著变化(P>0.05),T7组TLR6及T8组TLR2(P<0.01)和TLR9(P<0.01)基因表达量显著增加,TLR4显著降低(P<0.01)；T6和T7组TLR2和TLR9基因表达上显著低于T8(P<0.01),但TLR4基因表达显著高于T8(P<0.01),T7组TLR10基因表达显著少于T8(P<0.05)。结果提示,不同浓度的鼠李糖乳杆菌LR6001对IPEC-J2细胞TLR基因表达调控不同。
     综上可见,口服鼠李糖乳杆菌LR6001能够通过促进肠道微生态平衡,增强肠黏膜物理屏障、化学屏障以及免疫屏障功能来改善肠道健康促进仔猪断奶前后的生长并减少腹泻,其对哺乳仔猪肠黏膜免疫功能的调控可能是通过TLR信号途径介导而调控的。
Lactobacillus rhamnosus has been widely used due to its function of improving intestinal health, enhancing growth and reducing the rate of diarrhea in the host. Pre-sently, much attention is being focused on the immunomodulatory properties of Lac-tobacillus rhamnosus. However, little is known about the immunomodulatory proper-ties of Lactobacillus rhamnosus and its mechanisms of effects as an intestinal barrier in the piglets. Therefore, the present study was designed to investigate the effects of Lactobacillus rhamnosus6001(LR6001) as an immunomodulatory properties and intestinal barrier in vivo and in vitro. The main results are listed as follows:
     Trial1:The effects of orally administered Lactobacillus rhamnosus6001(LR6001) on the intestinal barrier of sucking piglets. Six litters (n=24) newborn piglets (Large White×Landrace×Duroc) were randomly divided into two groups,3litters per group. After birth and before the first sucking, piglets of control group orally received10%sterilized skim milk2mL piglet-1day-1. The LR6001group re-ceived10%sterilized skim milk2mL piglet-1day-1in addition to viable LR6001(5-6×108CFU·mL-1) for the first time, and another two of oral gavages were admi-nisted on the alternative odd days (3rd and5th day) of post partum. The feeding trial was conducted for25days and during experimental no antibiotic was added in the diet of sows and piglets throughout the trial. Results showed that oral administration of LR6001increased daily body weight gain by14.68%, while decreased diarrhea rate30.73%(P<0.05). Significantly Modulatory changes were appeared in the diversity of intestinal flora and the balance of microflora significantly, which led the beneficial effects by exerting prominent decrease in the growth of gram-negative bacteria (e.g., Bacteroidetes spp., Proteobacteria spp., Fusobacteria spp., and Enterobacteria), and an increased number of gram-positive bacteria noted (e.g., Firmicutes spp.). The ratio of villus height and crypt depth increased by20.13%(P<0.05) in jejunum of treat-ment group piglets, while the crypt depth decreased by10.01%(P<0.05). The gene expression of OCLN decreased by48.12%(P<0.05) in jejunum, but the expression of ZO-1remained unchanged. The gene expression of pBD-1and PMAP-37increased in jejunum of LR6001treated group, but LYS mRNA level noted lower significanlty. It was observed that the gene expression of MUC2and pH values numerically altered in the jejunum. The activities of DAO and ITF in ileul mucosa increased by375.15%(P<0.01) and100.44%(P<0.01) in the treatment group, respectively, while the activities of DAO and ITF in jejunal mucosa decreased by60.51%(P<0.01) and33.98%(P<0.01). The activity of maltase increased by41.29%(P<0.05) in jejunal mucosa of LR6001group, but the activities of sucrose and lactase were unaltered. Moreover, the concentrations of sIgA and C3in ileal mucosa increased by25.70%(P<0.05) and37.54%(P<0.05) in treatment group respectively, while the con-centrations of sIgA, C3and IgG in jejunal mucosa and IgG in ileal mucosa numari-cally improved. Additionally, LR6001treatment up-regulated the anti-inflammatory cytokines of TGF-β1and1L-10in jejuna mucosa, while the production of pro-inflammatory cytokines of IL-1, IL-6, IL-12and TNF-a, and chemokine of IL-8down-regulated (P<0.01) in both jejunal and ileal mucosa. It was also observed that pro-inflammatory cytokines of TNF-α, IFN-γ and chemokine of MCP-1up-regulated (P<0.01). Therefore, it provides the information that LR6001adminis-tration modulates the immune response by regulating the secretion of an-ti-inflammatory cytokines, pro-inflammatory cytokines, and chemokine in both jejun-al and ileal mucosa. Moreover, the gene expressions of MD-2, TLR2, TLR9, and TRAF6up-regulated21.29%(P<0.05),230.77%(P<0.01),107.34%(P<0.01), and81.92%(P<0.05), respectively, which suggests that MD-2, TLR2, TLR9, and TRAF6activated by immunomodulatory function of LR6001. All together, oral administra-tion of LR6001after birth promotes growth performance and improves the intestine health, regulates microbial ecological balance, functions of chemical barrier and physical barrier beside the TLR-meditated signaling pathway, as well as innate im-mune responses to reduce diarrhea in sucking piglets.
     Trial2:The effect of LR6001on intestinal mucosa barrier of weanling piglets was evaluated in the2nd trail. The same experimental protocol was used as described in Trial1. The results as compared to control group showed that average daily weight gain slightly increased in LR6001-treated group, and diarrhea incidence decreased by 15.32%(P>0.05). Oral administration of LR6001increased the diversity of intestinal flora, and also improved the composition of microflora. The amount of Firmicutes bacteria increased remarkably, while the amount of Bacteroid and Proteus reduced markedly. The structure of jejunal mucosa was more integrated in LR6001group than that in control group. The ratio of villus length and villus height to crypt depth in-creased by13.92%(P<0.01) and34.93%(P<0.01) in LR6001group respectively, and the crypt depth reduced by14.53%(P<0.05). Additionally, pH values in stomach re-duced in LR6001-treated piglets group. There was a decrease in the jejunal mucosal DAO activity by21.77%(P<0.01), whereas the serum concentrations of ITF and en-dotoxin were remained unchanged. Lactase activity in jejunal mucosa reduced13.60%(P<0.05) in LR6001group, but no significant difference was observed for the activities of sucrase and maltase. The mucosal concentration of sIgA increased by50%(P<0.01) in jejunum of LR6001-treated piglets, but the concentrations of IgG and C3slightly changed (P>0.05). Furthermore, LR6001treatment increased (P<0.01) anti-inflammatory cytokine TGF-β1and pro-inflammatory cytokine TNF-α, but re-duced (P<0.05) the level of inflammatory cytokine of IL-1β,IL-6,IL-12, IFN-γ and chemokine IL-8. Conclusively, we found that LR6001administered orally after birth improves the intestine health, prevents harmful effects of weaning stress in weaning piglets. It induces the increased capacity of absorption and balance the ecological en-vironment of intestine. Further, enhances the functions of chemical and physical bar-riers as well as innate immune responses in the piglets during the first week of wean-ing.
     Trial3:The effects of LR6001on expression level of TLR in piglets epithelial cells (IPEC-J2). The cultured IPEC-J2cells were randomly assigned into four treat-ment groups. Control (CK), treatmentⅠ (T6), treatmentⅡ (T7) and treatmentⅢ(T8), and were stimulated with0,106,107,108CFU·mL-1of Lactobacillus rhamnosus6001(LR6001). After stimulation period of3h, the IPEC-2cells were collected to investi-gate the mRNA expression of TLR2, TLR4, TLR6, TLR9and TLR10using real-time PCR. The results showed that the T8group increased the mRNA levels of TLR2(P<0.01) and TLR9(P<0.01), however TLR4mRNA levels reduced significantly as compared to CK, while T7showed increased level of TLR6mRNA expression. Among the experimental groups, the expression levels of TLR2and TLR9were lower in T6and T7as compared to T8(P<0.01), however, the TLR4gene expression level was higher in T6and T7in comparision of T8(P<0.01), and mRNA expression of TLR10in T7group was significanlty lower than T8group. The results indicated that the different concentrations of LR6001promote the proinfalammatory and antiinfa-lamatory response through activation of TLR.
     In summary, the mechanism of LR6001for the promotion of the intestinal health in piglets is due to the enhancement functions of biological, physical, chemical and immune barriers. Moreover, LR6001can also regulate the non-specificity immune response via TLR-meditated signaling pathways.

引文

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