抗生素、益生菌及银耳孢子发酵物对猪肠道硬壁门菌和拟杆门菌、脂肪沉积和脂肪代谢相关基因表达的影响及其作用机理
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摘要
肠道菌群种类和数量可以影响宿主脂肪沉积,而硬壁菌门(Firmicutes)和拟杆菌门(Bacteroidetes)是哺乳动物肠道中的优势菌群。前人的研究主要集中在粪便或食糜中菌群,反映大肠肠腔内菌群,不能反映小肠内及肠粘膜菌群。小肠内菌群与大肠明显不同,肠粘膜菌群与肠腔菌群明显不同。本研究在考察不同沉脂能力猪脂肪沉积和回肠内容物、回肠粘膜菌群差异的基础上,研究回肠粘膜菌群与脂肪沉积的关系和G-细菌细胞壁成分LPS及其受体TLR4与肠道粘膜菌群的关系,在此基础上考察不同微生态调节剂对猪回肠粘膜菌群、脂肪沉积、血液生化指标、炎性因子基因和脂肪代谢相关基因表达的影响,初步探讨肠粘膜菌群影响脂肪沉积的作用途径。研究包括以下四个试验：
试验一猪回肠粘膜和食糜菌群与脂肪沉积的关系
为研究肠粘膜上菌群与脂肪沉积的关系,本试验考察不同沉脂能力猪回肠粘膜和回肠食糜菌群差异和脂肪沉积差异。选取日龄接近,体重20kg左右杜洛克x梅山猪(DM)和杜长大猪(DLY)各6头,单笼饲养,饲喂同一玉米-豆粕日粮,试猪达110kg时全部屠宰,测定背膘厚度、眼肌面积、板油重和肌内脂肪含量,并采集回肠粘膜样品,采用根据细菌16S rDNA基因序列的实时荧光定量PCR方法测定回肠粘膜中硬壁菌门、拟杆菌门和总细菌拷贝数。结果表明：(1)DLY猪试验期极显著短于DM猪(P<0.01),平均日增重极显著高于DM猪(P<0.01),而平均日采食量显著低于DM猪(P<0.05),料肉比极显著低于DM猪(P<0.01)。DM猪和DLY猪板油重无显著差异,而DM猪的屠宰率、平均背膘厚度、眼肌面积和肌内脂肪含量均极显著高于DLY猪(P<0.01)。(2)两种猪回肠食糜中总细菌、硬壁菌和拟杆菌数量均高于回肠粘膜(P<0.01)。不同种类猪之间对比,DM猪回肠食糜中硬壁菌数量显著高于DLY猪(P<0.05)。而回肠食糜总细菌、拟杆菌及粘膜中总细菌、硬壁菌和拟杆菌在两种猪间无显著差异。DM猪回肠食糜和粘膜中硬壁菌和拟杆菌的比值均高于DLY猪(P<0.01)。(3)猪食糜中占总细菌比例最大的是硬壁菌(P<0.01),但回肠粘膜中占总细菌比例最大的是拟杆菌(P<0.01)。(4)肠粘膜和食糜中硬壁菌相对丰度、硬壁菌/拟杆菌比值与平均背膘厚正相关(P<0.01,P<0.01),而拟杆菌相对丰度与平均背膘厚度负相关(P<0.01)。硬壁菌／拟杆菌比值对背膘厚的影响大于硬壁菌／总细菌比值。
试验二猪回肠粘膜菌群与血浆LPS水平、TLR4、TLR5基因表达量的关系
本试验考察猪回肠粘膜菌群与血浆LPS水平、TLR4、TLR5基因的关系,试验动物和试验设计同试验一。屠宰后取回肠粘膜和皮下脂肪样品,测定回肠粘膜中硬壁菌门、拟杆菌门和总细菌数量,测定回肠粘膜和皮下脂肪中TLR4、TLR5基因相对于β-actin基因的表达量以及宰前血浆中LPS水平。结果表明：DM猪回肠粘膜和皮下脂肪中TLR4mRNA表达水平显著高于DLY猪(P<0.05)。DM猪和DLY猪回肠粘膜中TLR5mRNA相对表达量无显著差异(P>0.05)。两种猪皮下脂肪TLR4mRNA表达量均极显著高于回肠粘膜(P<0.01)。皮下脂肪和回肠粘膜中TLR4mRNA表达水平与肠道硬壁菌/拟杆菌比值均呈正相关关系(P<0.01),其决定系数R2分别为0.7437和0.7739。血浆LPS水平与硬壁菌/拟杆菌比值呈正的线性关系(P<0.01),其中血浆LPS水平与回肠粘膜中硬壁菌/拟杆菌比值相关性强于血浆LPS水平与回肠食糜中硬壁菌/拟杆菌比值的相关性。
试验三微生态调节剂对猪生产性能、胴体品质和回肠粘膜菌群的影响
试验一证明肠粘膜硬壁菌/拟杆菌比值与背膘厚呈正相关,且其相关关系强于食糜中菌群这一比值与背膘厚的关系。试验二证明肠粘膜硬壁菌/拟杆菌比值与血浆LPS、LPS受体TLR4基因表达水平呈正相关。为了进一步验证试验一和实验二结果,本试验考察不同微生态调节剂：硫酸粘杆菌素、芽孢杆菌和银耳孢子发酵物对猪生产性能、胴体品质和回肠粘膜菌群的影响。选取日龄接近,体重20+0.63kg的DLY猪24头,分为4个处理,每个处理6个重复,每个重复1头猪。4个处理组分别为对照组(C组,不加抗生素),硫酸粘杆菌素组(CS组),芽孢杆组(BS组),银耳孢子发酵物组(TFS组)。试猪达110kg左右时屠宰,每个处理屠宰4头猪,测定生产性能、胴体指标和回肠粘膜硬壁菌、拟杆菌及总细菌数量。结果表明：(1)硫酸粘杆菌素、芽孢杆菌和银耳孢子发酵物均可提高试猪生产性能。(2)各种微生态调节剂对DLY猪屠宰率和板油重无显著影响。硫酸粘杆菌素极显著降低试猪背膘厚度(P<0.01),有降低硬壁菌/总细菌比例和硬壁菌/拟杆菌比值及升高拟杆菌/总细菌比例的趋势；而芽孢杆菌极显著提高试猪背膘厚度(P<0.01),显著提高硬壁菌/总细菌比例(P<0.05)、有降低拟杆菌/总细菌比例和提高硬壁菌/拟杆菌比值的趋势。银耳孢子发酵物极显著提高试猪背膘厚度(P<0.01),极显著降低拟杆菌/总细菌比例(P<0.01)。(3)不同肠道微生态调节剂作用下猪平均背膘厚与回肠粘膜硬壁菌/总细菌和硬壁菌/拟杆菌比值呈正的线性关系(P<0.05)。
试验四微生态调节剂对猪血清指标、炎性因子和脂肪代谢相关基因表达量的影响
试验三结果验证了试验一的结论,为探讨肠道菌群影响脂肪沉积的作用途径,本试验在试验三的基础上考察硫酸粘杆菌素、芽孢杆菌和银耳孢子发酵物对猪血液生化指标、脂肪代谢相关基因和炎症因子基因表达量的影响。试验动物和试验设计同试验三,在试验三试猪体重达20kg.50kg和110kg体重时禁食13h后,早上8点左右前腔静脉采血,用以测定血浆LPS、血清中葡萄糖、胰岛素、瘦素和脂质水平。并在试猪达110kg体重时采集皮下脂肪和回肠粘膜,用以测定炎性因子基因和脂肪代谢相关基因的表达量。试验结果表明：(1)芽孢杆菌极显著提高试猪血浆脂多糖水平(P<0.01),硫酸粘杆菌素和银耳孢子发酵物有提高血浆脂多糖水平的趋势(P>0.05)。(2)微生态调节剂对不同体重阶段猪血液指标影响不同,但表现出一定趋势：硫酸粘杆菌素提高血糖、胰岛素水平,降低血清瘦素水平；芽孢杆菌降低血糖水平,提高血清瘦素水平；银耳孢子发酵物降低血糖、胰岛素水平,提高血清瘦素水平。硫酸粘杆菌素有降低猪血清总甘油三酯、胆固醇、游离脂肪酸水平的趋势；芽孢杆菌有提高血清甘油三酯,总胆固醇水平的趋势；银耳孢子发酵物有降低110kg阶段猪血清甘油三酯和总胆固醇水平的趋势。(3)不同微生态调节剂对不同组织炎性因子基因表达量影响不同,但总体上讲,硫酸粘杆菌素降低TLR4.TNF-α、IL1β和IL6表达量,芽孢杆菌提高TLR4、 TNF-α、IL1β和IL6表达量,银耳孢子发酵物提高TLR4-TNF-α、IL1β表达量。(4)硫酸粘杆菌素降低猪脂肪合成相关基因的表达,提高脂肪分解转运相关基因的表达；芽孢杆菌提高脂肪合成相关基因的表达量,对脂肪分解基因表达量影响较小；银耳孢子发酵物提高猪脂肪分解相关基因表达量,降低脂肪合成相关基因表达量。
综上所述：
(1)猪回肠粘膜中总细菌、硬壁菌和拟杆菌数量均显著低于回肠食糜中各类细菌数量
(2)硬壁菌/拟杆菌比值对背膘厚的影响大于硬壁菌/总细菌比值；猪回肠粘膜硬壁菌/拟杆菌比值与脂肪沉积的相关关系强于回肠食糜硬壁菌/拟杆菌比值与脂肪沉积的相关关系
(3)猪血浆LPS水平与回肠硬壁菌/拟A杆菌比值正相关；猪脂肪组织和回肠粘膜TLR4基因表达水平与肠粘膜硬壁菌/拟杆菌比值正相关
(4)根据实验一得出的相关关系,CS降低试猪背膘厚与其减小硬/总和硬/拟比值结果一致；BS增加试猪背膘厚与其增加硬/总和硬/拟比值结果一致；而TSF增加试猪背膘厚与其增加硬/总结果一致,但减小硬/拟比值结果不一致
(5)不同微生态制剂作用下背膘厚和硬壁门菌/拟杆菌的变化规律与血浆LPS,TLR4基因的变化不完全一致
(6)硫酸粘杆菌素、芽孢杆菌可能通过影响猪肠道食糜和粘膜中硬壁菌、拟杆菌数量及丰度,从而影响试猪皮下脂肪和回肠粘膜炎性因子基因的表达,脂肪代谢关键酶基因的表达,而最终影响脂肪的沉积能力。而银耳孢子发酵物影响脂肪沉积的作用与回肠菌群变化无关。
Gut microbiota which is mainly consisted of Firmicutes and Bacteroidetes divisions has been shown to affect fat storage and energy harvesting of host.Previous studies mainly focused on the flora in the feces or digesta, reflecting flora of the intestine lumen not the intestinal mucosa flora. Mucosa flora composition was significantly different from intestine lumen.Duroc x Meishan and DLY pigs has different capable of fat deposition. The amount of fat deposition and the intestinal flora of the two kinds of rossbred pigs were investigated. Then the correlation between intestinal mucosa flora and the amount of fat deposition was studied. And the correlation between intestinal mucosa flora and the expression level of TLR4was studied. The effect of different microbial modulator on intestinal mucosa flora, the amount of fat deposition, blood biochemical parameters, the inflammatory factor gene and fat metabolism gene expression were investigated. The mechanism that the gut microbiota regulates fat storage in pigs was also investigated. Four experiments were conducted as follows:
EXP.1The correlation between intestinal mucosa flora and the amount of fat deposition of pigs
To study the relationship between intestinal mucosa flora and the amount of fat deposition, the amount of fat deposition and the intestinal flora of Duroc x Meishan(DM) and DLY pigs were investigated. Six DM pigs and Six DLY pigs(about20kg) were feed the same corn-soybean meal. Each pig was removed from the test on the day it most nearly approximated110kg. Every pig from each treatment were slaughtered, and the backfat depth, longissimus muscle area, leaf lard weight and intramuscular fat content of pigs were measured. Ileum mucosa were collected for detecting the16S rDNA gene sequence of total bacterial, Firmicutes and Bacteroidetes divisions using the RT-PCR method. Results of the experiment indicated that:(1) DLY pigs trial period was significantly shorter than the DM pigs(P<0.01), average daily gain was significantly higher than DM pigs (P<0.01), while the average daily feed intake was significantly lower than DM pigs (P <0.05), F/G was significantly lower than DM pigs (P<0.01). Leaf lard weight was no significant different between DM and DLY pigs. The yield of carcass, average backfat depth, longissimus muscle area, intramuscular fat content of DM pigs were significantly higher than DLY pigs (P<0.01).(2)The copy numbers of total bacterial,Firmicutes and Bacteroidetes divisions of ileum contents from both kind pigs were significantly higher than ileum mucosa(P<0.01). The copy numbers of Firmicutes divisions of ileum contents from DM pigs were significantly higher than DLY pigs(P<0.05). There were no significant different of total bacterial, Firmicutes and Bacteroidetes divisions of ileum mucosa between two kinds of rossbred pigs. The copy numbers of Firmicutes and Bacteroidetes divisions of ileum contents and ileum mucosa frome DM pigs were significantly higher than DLY pigs(P<0.01).(3)Both kind pigs having the sum of percents of Firmicutes and Bacteroidetes divisions of ileum contents and ileum mucosa which were about85%. Firmicutes of ileum contents from both kind pig has the largest proportion(P<0.01) while Bacteroidetes of ileum mucosa has the largest proportion(P<0.01).(4) The relative abundance of Firmicutes, Firmicutes/Bacteroides ratio of gut lumenal contents and intestinal mucosa were positively correlated to average backfat(.P<0.01,.P<0.01), while the relative abundance of Bacteroides and average backfat thickness negatively correlated (.P<0.01).
EXP.2The relationship between ileum mucosa flora and plasma LPS level, TLR4, TLR5gene expression level of pigs
The relationship between ileum mucosa flora and plasma LPS level, TLR4, TLR5gene expression level was studied,animal and the experimental design were the same to EXP.1. After slaughter, ileum mucosa and subcutaneous fat were collected for measuring the copy numbers of Firmicutes and Bacteroidetes divisions and expression level of TLR4and TLR5. Results of the experiment indicated that:TLR4mRNA expression levels of intestinal mucosa and subcutaneous fat from DM pigs were significantly higher than DLY pigs (P<0.05). TLR5mRNA expression levels of intestinal mucosa from DM pigs was no significant different from DLY pigs. TLR4mRNA expression levels of subcutaneous fat from both kind pigs was significantly higher than intestinal mucosa(P<0.01). TLR4mRNA expression levels of intestinal mucosa and subcutaneous fat were positive correlated to intestine Firmicutes/Bacteroidetes, The coefficient of determination R2were0.7437and0.7739respectively.
EXP.3The effect of the of microbial regulators on pig performance, carcass quality and mucosal flora
The EXP.1proved that the ratio of Firmicutes/Bacteroidetes was positively correlated to fat deposition amount, and the coefficient of determination was higher than the one of ileum contents. To further validate this relationship, the effect of colistin sulfate, Bacillus and white fungus spores fermentation products on pig production performance, carcass quality and mucosa flora were investigated.24pigs (20±0.63kg) were randomly allotted into4dietary treatments with6replicate pens per treatment and1pig per pen:(1) control diets containing no antibiotic (C group),(2) colistin sulfate added to the control diets (CS group),(3) Bacillus added to the control diets (BS group),(4) TFS added to the control diets (TFS group). Each pig was removed from the test on the day it most nearly approximated110kg.4pigs from each treatment were slaughtered, and the backfat depth, longissimus muscle area, leaf lard weight, intramuscular fat content and the copy number of total bacterial, Firmicutes and Bacteroidetes of ileum mucosa were measured. Results of the experiment indicated that:(1) The colistin sulfate, probiotics and white fungus spores fermentation products improved pig performance.(2) Variety of microbial modulator showed no significant effect on yield of carcass and leaf lard weight.The colistin sulfate significantly reduced the pig backfat thickness (P<0.01), and probiotics and white fungus spores fermentation products significantly improved pig backfat depth, longissimus muscle area and intramuscular fat content.(3)There were no significantly differences among the copy numbers of total bacterial,Firmicutes and Bacteroidetes. Colistin sulfate had no effect on the rate of Firmicutes/total bacterial and Firmicutes/Bacteroidetes, but had trends of reducing the rate of Firmicutes/total bacterial, Firmicutes/Bacteroidetes and increasing the rate of Bacteroidetes/total bacterial. Probiotic significantly increased the rate of Firmicutes/total bacterial(P<0.05) and had trends of reducing the rate of Bacteroidetes/total bacterial and increasing the rate of Firmicutes/Bacteroidetes. White fungus spores fermentation products significantly reduced the rate of Bacteroidetes/total bacterial. The BS and TSF goup had higher rate of Firmicutes/total bacterial than CS group(P<0.05). The TSF goup had the lowest rate of Bacteroidetes/total bacterial than CS group(P<0.01).
EXP.4Effect of microbial modulator on pig serum indicators inflammatory factors and fat metabolism gene expression levels
The EXP.3verified the conclusion of the EXP.1, to study the pathway of intestinal flora affecting fat deposition, the effect of microbial regulators on pig blood biochemical indicators, the expression levels of fat metabolism-related genes and the inflammatory cytokines were investigated. Animals and experimental design were same to EXP.3. Vena cava blood were sampled for determination of plasma LPS, serum glucose, insulin, leptin, and lipid levels when pigs have weights up to20kg,50kg and110kg. Subcutaneous fat and intestinal mucosa were sampled for determination of inflammatory factors and fat metabolism gene expression levels. Results of the experiment indicated that:(1)The probiotic significantly increased the level of plasma lipopolysaccharide, colistin sulfate and white fungus spores fermentation products trend to increase plasma lipopolysaccharide levels.(2)Microbial modulator has different effect on blood indicators of different weight stage pigs, but showed some trend:the colistin sulfate raise plasma glucose and insulin levels, and lower serum leptin levels; probiotics reduce plasma glucose levels, serum leptin levels; TSF lower plasma glucose and insulin levels, serum leptin levels. Colistin sulfate trend to reduce the pig serum total triglycerides, cholesterol, free fatty acids; probiotics trend to increase serum triglycerides, total cholesterol; TSF trend to reduce110kg stage pig serum triglycerides and total cholesterol.(3) Microbial modulator has different effect on inflammatory factor gene expression levels of different tissue, but generally speaking, colistin sulfate reduce TLR4, TNF-a, IL1β and IL6expression levels, probiotics increase TLR4, TNF-a, IL1β and IL6expression levels, and TSF increase TLR4, TNF-a, IL1β.(4)The colistin sulfate reduced the pig fat synthesis gene expression, increased lipolysis and fat transport related gene expression; probiotics increased fat synthesis genes expression, reduce lipolysis gene expression; TSF increase pig fat decomposition related gene expression levels, reduce fat synthesis gene expression.
In summary, the results of this study showed that:
(1) The total bacteria, relative abundance of Firmicutes and Bacteroides from ileal mucosa were significantly lower than the lumenal
(2) The effect of Firmicutes/Bacteroides ratio on backfat thickness is stronger than Firmicutes/total bacteria ratio; Lleum mucosa Firmicutes/Bacteroides ratios has stronger correlation to fat deposition than ileal digesta in pig
(3) Porcine plasma LPS levels and ileum Firmicutes/Bacteroides ratio were positively correlated; TLR4gene expression levels in porcine adipose tissue and the Ileum mucosa were positively correlated to intestinal mucosa Firmicutes/Bacteroides ratio
(4) According to the results of Exp.1, CS decrease backfat thickness is consistent with decreasing Firmicutes/Bacteroides ratio; BS increase backfat thickness is consistent with increasing Firmicutes/Bacteroides ratio; TSF increase backfat thickness is inconsistent with decreasing Firmicutes/Bacteroides ratio
(5) Under the action of different probiotics backfat thickness and Firmicutes/Bacteroides variation are not entirely consistent with plasma LPS, TLR4gene changes
(6) Colistin sulfate, Bacillus may influence the number and abundance of Firmicutes and Bacteroides in pig intestine, thus affecting the trial pig backfat and intestinal inflammatory cytokines gene expression, fat metabolism key enzyme gene expression, and ultimately affect the ability of fat deposition.The Tremella fuciformis ferment substance affect the role of fat deposition has nothing to do with ileal flora..
试验一猪回肠粘膜和食糜菌群与脂肪沉积的关系
为研究肠粘膜上菌群与脂肪沉积的关系,本试验考察不同沉脂能力猪回肠粘膜和回肠食糜菌群差异和脂肪沉积差异。选取日龄接近,体重20kg左右杜洛克x梅山猪(DM)和杜长大猪(DLY)各6头,单笼饲养,饲喂同一玉米-豆粕日粮,试猪达110kg时全部屠宰,测定背膘厚度、眼肌面积、板油重和肌内脂肪含量,并采集回肠粘膜样品,采用根据细菌16S rDNA基因序列的实时荧光定量PCR方法测定回肠粘膜中硬壁菌门、拟杆菌门和总细菌拷贝数。结果表明：(1)DLY猪试验期极显著短于DM猪(P<0.01),平均日增重极显著高于DM猪(P<0.01),而平均日采食量显著低于DM猪(P<0.05),料肉比极显著低于DM猪(P<0.01)。DM猪和DLY猪板油重无显著差异,而DM猪的屠宰率、平均背膘厚度、眼肌面积和肌内脂肪含量均极显著高于DLY猪(P<0.01)。(2)两种猪回肠食糜中总细菌、硬壁菌和拟杆菌数量均高于回肠粘膜(P<0.01)。不同种类猪之间对比,DM猪回肠食糜中硬壁菌数量显著高于DLY猪(P<0.05)。而回肠食糜总细菌、拟杆菌及粘膜中总细菌、硬壁菌和拟杆菌在两种猪间无显著差异。DM猪回肠食糜和粘膜中硬壁菌和拟杆菌的比值均高于DLY猪(P<0.01)。(3)猪食糜中占总细菌比例最大的是硬壁菌(P<0.01),但回肠粘膜中占总细菌比例最大的是拟杆菌(P<0.01)。(4)肠粘膜和食糜中硬壁菌相对丰度、硬壁菌/拟杆菌比值与平均背膘厚正相关(P<0.01,P<0.01),而拟杆菌相对丰度与平均背膘厚度负相关(P<0.01)。硬壁菌／拟杆菌比值对背膘厚的影响大于硬壁菌／总细菌比值。
试验二猪回肠粘膜菌群与血浆LPS水平、TLR4、TLR5基因表达量的关系
本试验考察猪回肠粘膜菌群与血浆LPS水平、TLR4、TLR5基因的关系,试验动物和试验设计同试验一。屠宰后取回肠粘膜和皮下脂肪样品,测定回肠粘膜中硬壁菌门、拟杆菌门和总细菌数量,测定回肠粘膜和皮下脂肪中TLR4、TLR5基因相对于β-actin基因的表达量以及宰前血浆中LPS水平。结果表明：DM猪回肠粘膜和皮下脂肪中TLR4mRNA表达水平显著高于DLY猪(P<0.05)。DM猪和DLY猪回肠粘膜中TLR5mRNA相对表达量无显著差异(P>0.05)。两种猪皮下脂肪TLR4mRNA表达量均极显著高于回肠粘膜(P<0.01)。皮下脂肪和回肠粘膜中TLR4mRNA表达水平与肠道硬壁菌/拟杆菌比值均呈正相关关系(P<0.01),其决定系数R2分别为0.7437和0.7739。血浆LPS水平与硬壁菌/拟杆菌比值呈正的线性关系(P<0.01),其中血浆LPS水平与回肠粘膜中硬壁菌/拟杆菌比值相关性强于血浆LPS水平与回肠食糜中硬壁菌/拟杆菌比值的相关性。
试验三微生态调节剂对猪生产性能、胴体品质和回肠粘膜菌群的影响
试验一证明肠粘膜硬壁菌/拟杆菌比值与背膘厚呈正相关,且其相关关系强于食糜中菌群这一比值与背膘厚的关系。试验二证明肠粘膜硬壁菌/拟杆菌比值与血浆LPS、LPS受体TLR4基因表达水平呈正相关。为了进一步验证试验一和实验二结果,本试验考察不同微生态调节剂：硫酸粘杆菌素、芽孢杆菌和银耳孢子发酵物对猪生产性能、胴体品质和回肠粘膜菌群的影响。选取日龄接近,体重20+0.63kg的DLY猪24头,分为4个处理,每个处理6个重复,每个重复1头猪。4个处理组分别为对照组(C组,不加抗生素),硫酸粘杆菌素组(CS组),芽孢杆组(BS组),银耳孢子发酵物组(TFS组)。试猪达110kg左右时屠宰,每个处理屠宰4头猪,测定生产性能、胴体指标和回肠粘膜硬壁菌、拟杆菌及总细菌数量。结果表明：(1)硫酸粘杆菌素、芽孢杆菌和银耳孢子发酵物均可提高试猪生产性能。(2)各种微生态调节剂对DLY猪屠宰率和板油重无显著影响。硫酸粘杆菌素极显著降低试猪背膘厚度(P<0.01),有降低硬壁菌/总细菌比例和硬壁菌/拟杆菌比值及升高拟杆菌/总细菌比例的趋势；而芽孢杆菌极显著提高试猪背膘厚度(P<0.01),显著提高硬壁菌/总细菌比例(P<0.05)、有降低拟杆菌/总细菌比例和提高硬壁菌/拟杆菌比值的趋势。银耳孢子发酵物极显著提高试猪背膘厚度(P<0.01),极显著降低拟杆菌/总细菌比例(P<0.01)。(3)不同肠道微生态调节剂作用下猪平均背膘厚与回肠粘膜硬壁菌/总细菌和硬壁菌/拟杆菌比值呈正的线性关系(P<0.05)。
试验四微生态调节剂对猪血清指标、炎性因子和脂肪代谢相关基因表达量的影响
试验三结果验证了试验一的结论,为探讨肠道菌群影响脂肪沉积的作用途径,本试验在试验三的基础上考察硫酸粘杆菌素、芽孢杆菌和银耳孢子发酵物对猪血液生化指标、脂肪代谢相关基因和炎症因子基因表达量的影响。试验动物和试验设计同试验三,在试验三试猪体重达20kg.50kg和110kg体重时禁食13h后,早上8点左右前腔静脉采血,用以测定血浆LPS、血清中葡萄糖、胰岛素、瘦素和脂质水平。并在试猪达110kg体重时采集皮下脂肪和回肠粘膜,用以测定炎性因子基因和脂肪代谢相关基因的表达量。试验结果表明：(1)芽孢杆菌极显著提高试猪血浆脂多糖水平(P<0.01),硫酸粘杆菌素和银耳孢子发酵物有提高血浆脂多糖水平的趋势(P>0.05)。(2)微生态调节剂对不同体重阶段猪血液指标影响不同,但表现出一定趋势：硫酸粘杆菌素提高血糖、胰岛素水平,降低血清瘦素水平；芽孢杆菌降低血糖水平,提高血清瘦素水平；银耳孢子发酵物降低血糖、胰岛素水平,提高血清瘦素水平。硫酸粘杆菌素有降低猪血清总甘油三酯、胆固醇、游离脂肪酸水平的趋势；芽孢杆菌有提高血清甘油三酯,总胆固醇水平的趋势；银耳孢子发酵物有降低110kg阶段猪血清甘油三酯和总胆固醇水平的趋势。(3)不同微生态调节剂对不同组织炎性因子基因表达量影响不同,但总体上讲,硫酸粘杆菌素降低TLR4.TNF-α、IL1β和IL6表达量,芽孢杆菌提高TLR4、 TNF-α、IL1β和IL6表达量,银耳孢子发酵物提高TLR4-TNF-α、IL1β表达量。(4)硫酸粘杆菌素降低猪脂肪合成相关基因的表达,提高脂肪分解转运相关基因的表达；芽孢杆菌提高脂肪合成相关基因的表达量,对脂肪分解基因表达量影响较小；银耳孢子发酵物提高猪脂肪分解相关基因表达量,降低脂肪合成相关基因表达量。
综上所述：
(1)猪回肠粘膜中总细菌、硬壁菌和拟杆菌数量均显著低于回肠食糜中各类细菌数量
(2)硬壁菌/拟杆菌比值对背膘厚的影响大于硬壁菌/总细菌比值；猪回肠粘膜硬壁菌/拟杆菌比值与脂肪沉积的相关关系强于回肠食糜硬壁菌/拟杆菌比值与脂肪沉积的相关关系
(3)猪血浆LPS水平与回肠硬壁菌/拟A杆菌比值正相关；猪脂肪组织和回肠粘膜TLR4基因表达水平与肠粘膜硬壁菌/拟杆菌比值正相关
(4)根据实验一得出的相关关系,CS降低试猪背膘厚与其减小硬/总和硬/拟比值结果一致；BS增加试猪背膘厚与其增加硬/总和硬/拟比值结果一致；而TSF增加试猪背膘厚与其增加硬/总结果一致,但减小硬/拟比值结果不一致
(5)不同微生态制剂作用下背膘厚和硬壁门菌/拟杆菌的变化规律与血浆LPS,TLR4基因的变化不完全一致
(6)硫酸粘杆菌素、芽孢杆菌可能通过影响猪肠道食糜和粘膜中硬壁菌、拟杆菌数量及丰度,从而影响试猪皮下脂肪和回肠粘膜炎性因子基因的表达,脂肪代谢关键酶基因的表达,而最终影响脂肪的沉积能力。而银耳孢子发酵物影响脂肪沉积的作用与回肠菌群变化无关。
Gut microbiota which is mainly consisted of Firmicutes and Bacteroidetes divisions has been shown to affect fat storage and energy harvesting of host.Previous studies mainly focused on the flora in the feces or digesta, reflecting flora of the intestine lumen not the intestinal mucosa flora. Mucosa flora composition was significantly different from intestine lumen.Duroc x Meishan and DLY pigs has different capable of fat deposition. The amount of fat deposition and the intestinal flora of the two kinds of rossbred pigs were investigated. Then the correlation between intestinal mucosa flora and the amount of fat deposition was studied. And the correlation between intestinal mucosa flora and the expression level of TLR4was studied. The effect of different microbial modulator on intestinal mucosa flora, the amount of fat deposition, blood biochemical parameters, the inflammatory factor gene and fat metabolism gene expression were investigated. The mechanism that the gut microbiota regulates fat storage in pigs was also investigated. Four experiments were conducted as follows:
EXP.1The correlation between intestinal mucosa flora and the amount of fat deposition of pigs
To study the relationship between intestinal mucosa flora and the amount of fat deposition, the amount of fat deposition and the intestinal flora of Duroc x Meishan(DM) and DLY pigs were investigated. Six DM pigs and Six DLY pigs(about20kg) were feed the same corn-soybean meal. Each pig was removed from the test on the day it most nearly approximated110kg. Every pig from each treatment were slaughtered, and the backfat depth, longissimus muscle area, leaf lard weight and intramuscular fat content of pigs were measured. Ileum mucosa were collected for detecting the16S rDNA gene sequence of total bacterial, Firmicutes and Bacteroidetes divisions using the RT-PCR method. Results of the experiment indicated that:(1) DLY pigs trial period was significantly shorter than the DM pigs(P<0.01), average daily gain was significantly higher than DM pigs (P<0.01), while the average daily feed intake was significantly lower than DM pigs (P <0.05), F/G was significantly lower than DM pigs (P<0.01). Leaf lard weight was no significant different between DM and DLY pigs. The yield of carcass, average backfat depth, longissimus muscle area, intramuscular fat content of DM pigs were significantly higher than DLY pigs (P<0.01).(2)The copy numbers of total bacterial,Firmicutes and Bacteroidetes divisions of ileum contents from both kind pigs were significantly higher than ileum mucosa(P<0.01). The copy numbers of Firmicutes divisions of ileum contents from DM pigs were significantly higher than DLY pigs(P<0.05). There were no significant different of total bacterial, Firmicutes and Bacteroidetes divisions of ileum mucosa between two kinds of rossbred pigs. The copy numbers of Firmicutes and Bacteroidetes divisions of ileum contents and ileum mucosa frome DM pigs were significantly higher than DLY pigs(P<0.01).(3)Both kind pigs having the sum of percents of Firmicutes and Bacteroidetes divisions of ileum contents and ileum mucosa which were about85%. Firmicutes of ileum contents from both kind pig has the largest proportion(P<0.01) while Bacteroidetes of ileum mucosa has the largest proportion(P<0.01).(4) The relative abundance of Firmicutes, Firmicutes/Bacteroides ratio of gut lumenal contents and intestinal mucosa were positively correlated to average backfat(.P<0.01,.P<0.01), while the relative abundance of Bacteroides and average backfat thickness negatively correlated (.P<0.01).
EXP.2The relationship between ileum mucosa flora and plasma LPS level, TLR4, TLR5gene expression level of pigs
The relationship between ileum mucosa flora and plasma LPS level, TLR4, TLR5gene expression level was studied,animal and the experimental design were the same to EXP.1. After slaughter, ileum mucosa and subcutaneous fat were collected for measuring the copy numbers of Firmicutes and Bacteroidetes divisions and expression level of TLR4and TLR5. Results of the experiment indicated that:TLR4mRNA expression levels of intestinal mucosa and subcutaneous fat from DM pigs were significantly higher than DLY pigs (P<0.05). TLR5mRNA expression levels of intestinal mucosa from DM pigs was no significant different from DLY pigs. TLR4mRNA expression levels of subcutaneous fat from both kind pigs was significantly higher than intestinal mucosa(P<0.01). TLR4mRNA expression levels of intestinal mucosa and subcutaneous fat were positive correlated to intestine Firmicutes/Bacteroidetes, The coefficient of determination R2were0.7437and0.7739respectively.
EXP.3The effect of the of microbial regulators on pig performance, carcass quality and mucosal flora
The EXP.1proved that the ratio of Firmicutes/Bacteroidetes was positively correlated to fat deposition amount, and the coefficient of determination was higher than the one of ileum contents. To further validate this relationship, the effect of colistin sulfate, Bacillus and white fungus spores fermentation products on pig production performance, carcass quality and mucosa flora were investigated.24pigs (20±0.63kg) were randomly allotted into4dietary treatments with6replicate pens per treatment and1pig per pen:(1) control diets containing no antibiotic (C group),(2) colistin sulfate added to the control diets (CS group),(3) Bacillus added to the control diets (BS group),(4) TFS added to the control diets (TFS group). Each pig was removed from the test on the day it most nearly approximated110kg.4pigs from each treatment were slaughtered, and the backfat depth, longissimus muscle area, leaf lard weight, intramuscular fat content and the copy number of total bacterial, Firmicutes and Bacteroidetes of ileum mucosa were measured. Results of the experiment indicated that:(1) The colistin sulfate, probiotics and white fungus spores fermentation products improved pig performance.(2) Variety of microbial modulator showed no significant effect on yield of carcass and leaf lard weight.The colistin sulfate significantly reduced the pig backfat thickness (P<0.01), and probiotics and white fungus spores fermentation products significantly improved pig backfat depth, longissimus muscle area and intramuscular fat content.(3)There were no significantly differences among the copy numbers of total bacterial,Firmicutes and Bacteroidetes. Colistin sulfate had no effect on the rate of Firmicutes/total bacterial and Firmicutes/Bacteroidetes, but had trends of reducing the rate of Firmicutes/total bacterial, Firmicutes/Bacteroidetes and increasing the rate of Bacteroidetes/total bacterial. Probiotic significantly increased the rate of Firmicutes/total bacterial(P<0.05) and had trends of reducing the rate of Bacteroidetes/total bacterial and increasing the rate of Firmicutes/Bacteroidetes. White fungus spores fermentation products significantly reduced the rate of Bacteroidetes/total bacterial. The BS and TSF goup had higher rate of Firmicutes/total bacterial than CS group(P<0.05). The TSF goup had the lowest rate of Bacteroidetes/total bacterial than CS group(P<0.01).
EXP.4Effect of microbial modulator on pig serum indicators inflammatory factors and fat metabolism gene expression levels
The EXP.3verified the conclusion of the EXP.1, to study the pathway of intestinal flora affecting fat deposition, the effect of microbial regulators on pig blood biochemical indicators, the expression levels of fat metabolism-related genes and the inflammatory cytokines were investigated. Animals and experimental design were same to EXP.3. Vena cava blood were sampled for determination of plasma LPS, serum glucose, insulin, leptin, and lipid levels when pigs have weights up to20kg,50kg and110kg. Subcutaneous fat and intestinal mucosa were sampled for determination of inflammatory factors and fat metabolism gene expression levels. Results of the experiment indicated that:(1)The probiotic significantly increased the level of plasma lipopolysaccharide, colistin sulfate and white fungus spores fermentation products trend to increase plasma lipopolysaccharide levels.(2)Microbial modulator has different effect on blood indicators of different weight stage pigs, but showed some trend:the colistin sulfate raise plasma glucose and insulin levels, and lower serum leptin levels; probiotics reduce plasma glucose levels, serum leptin levels; TSF lower plasma glucose and insulin levels, serum leptin levels. Colistin sulfate trend to reduce the pig serum total triglycerides, cholesterol, free fatty acids; probiotics trend to increase serum triglycerides, total cholesterol; TSF trend to reduce110kg stage pig serum triglycerides and total cholesterol.(3) Microbial modulator has different effect on inflammatory factor gene expression levels of different tissue, but generally speaking, colistin sulfate reduce TLR4, TNF-a, IL1β and IL6expression levels, probiotics increase TLR4, TNF-a, IL1β and IL6expression levels, and TSF increase TLR4, TNF-a, IL1β.(4)The colistin sulfate reduced the pig fat synthesis gene expression, increased lipolysis and fat transport related gene expression; probiotics increased fat synthesis genes expression, reduce lipolysis gene expression; TSF increase pig fat decomposition related gene expression levels, reduce fat synthesis gene expression.
In summary, the results of this study showed that:
(1) The total bacteria, relative abundance of Firmicutes and Bacteroides from ileal mucosa were significantly lower than the lumenal
(2) The effect of Firmicutes/Bacteroides ratio on backfat thickness is stronger than Firmicutes/total bacteria ratio; Lleum mucosa Firmicutes/Bacteroides ratios has stronger correlation to fat deposition than ileal digesta in pig
(3) Porcine plasma LPS levels and ileum Firmicutes/Bacteroides ratio were positively correlated; TLR4gene expression levels in porcine adipose tissue and the Ileum mucosa were positively correlated to intestinal mucosa Firmicutes/Bacteroides ratio
(4) According to the results of Exp.1, CS decrease backfat thickness is consistent with decreasing Firmicutes/Bacteroides ratio; BS increase backfat thickness is consistent with increasing Firmicutes/Bacteroides ratio; TSF increase backfat thickness is inconsistent with decreasing Firmicutes/Bacteroides ratio
(5) Under the action of different probiotics backfat thickness and Firmicutes/Bacteroides variation are not entirely consistent with plasma LPS, TLR4gene changes
(6) Colistin sulfate, Bacillus may influence the number and abundance of Firmicutes and Bacteroides in pig intestine, thus affecting the trial pig backfat and intestinal inflammatory cytokines gene expression, fat metabolism key enzyme gene expression, and ultimately affect the ability of fat deposition.The Tremella fuciformis ferment substance affect the role of fat deposition has nothing to do with ileal flora..
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