益生菌调节蛋鸡胆固醇代谢的研究进展发表时间:2020-10-25 15:57 胆固醇是动物细胞膜的重要组成成分,是胆汁酸、维生素D3和类固醇激素的合成前体物质,具有重要的生理功能[1]。 体内胆固醇含量过低(di)(di)或(huo)过高对健康都是不(bu)利的(de),胆固醇过低(di)(di)会导致类固醇激素(su)等(deng)物质合成不(bu)足,影响机(ji)体发育[2]; 胆固醇过高(gao)则会(hui)提高(gao)心血(xue)管疾病的(de)发病率[3],鸡蛋(dan)含(han)有(you)人体(ti)所(suo)必需(xu)的(de)蛋(dan)白质(zhi)、脂(zhi)肪、微量元(yuan)素、维生(sheng)素等(deng),所(suo)含(han)营养元(yuan)素丰富而均衡,且易(yi)于被人体(ti)消(xiao)化吸收(shou),所(suo)以(yi)(yi)自古以(yi)(yi)来(lai)一(yi)直是人们(men)所(suo)青睐的(de)营养食品之(zhi)一(yi)[4]。 然而,关于(yu)(yu)食(shi)用鸡(ji)蛋是否导致心血(xue)管(guan)疾(ji)病(bing)发病(bing)率(lv)提升仍存(cun)在争议(yi)。一些研究者认为食(shi)用鸡(ji)蛋会(hui)提高心血(xue)管(guan)疾(ji)病(bing)(冠心病(bing)、动脉壁粥(zhou)样硬化等(deng))的(de)(de)发生率(lv)[3,5-7],其危害几(ji)乎等(deng)于(yu)(yu)吸(xi)烟坏处的(de)(de)2/3[3]; 但另一种观(guan)点认为食用鸡蛋(dan)(dan)与心血管疾病的(de)发生不(bu)存(cun)在相(xiang)关关系,摄(she)入鸡蛋(dan)(dan)对血清(qing)胆固(gu)醇(chun)的(de)影(ying)响是微(wei)弱(ruo)的(de)[8-10],分歧(qi)存(cun)在的(de)主要(yao)原因是蛋(dan)(dan)黄在人体内的(de)具体代谢机(ji)制尚不(bu)清(qing)楚。 鸡蛋中胆固(gu)醇的含量约(yue)为213mg/枚,受(shou)到品(pin)种[11]、鸡龄[12-13]、产蛋率[14]以(yi)及饲粮成分[15-16]等多种因素(su)的影响。 由于胆(dan)固醇是家禽胚(pei)胎(tai)发育的(de)必需(xu)物质,降低鸡蛋中胆(dan)固醇含量(liang)将影响(xiang)胚(pei)胎(tai)的(de)发育,因(yin)此品种选育对调节蛋黄胆(dan)固醇的(de)作用很小(<8%)[17],因(yin)此,动物营(ying)养研(yan)究人员尝试(shi)通(tong)过改变饲(si)粮成分,如(ru)脂肪[16]、ω3脂肪酸[18]、微量(liang)元(yuan)素[19] 含(han)量和添加大蒜素[20-21]等调节(jie)蛋(dan)鸡胆固醇(chun)代(dai)谢(xie)。此外,益生菌作为一(yi)类(lei)活体微(wei)生物,具有多种生物活性,已(yi)有试验表明,调节(jie)机体胆固醇(chun)代(dai)谢(xie)是其活性之一(yi)。 1蛋鸡胆固醇代谢蛋鸡饲粮通常以植物性(xing)原料为主,很(hen)少含有动(dong)物性(xing)原料,因此,蛋鸡体内(nei)的胆固醇主要依靠自(zi)身(shen)合成。 蛋鸡体内的(de)胆(dan)固(gu)醇(chun)主要在肝脏和卵巢中合(he)成,其中肝脏不(bu)仅合(he)成速(su)度快,而且合(he)成量多,是蛋鸡最活跃的(de)胆(dan)固(gu)醇(chun)合(he)成部位和血液胆(dan)固(gu)醇(chun)来源(yuan)[22]。 产蛋鸡肝脏胆(dan)固(gu)醇(chun)合成代谢(xie)易(yi)受到饲(si)粮成分(fen),如脂肪、胆(dan)固(gu)醇(chun)、纤维含量的影响(xiang),而(er)卵巢受饲(si)粮影响(xiang)相对(dui)较小(xiao)[22-23]。Naber[22]推测1只体(ti)重约(yue)为1.7kg的产蛋鸡在饲(si)喂(wei)无胆(dan)固(gu)醇(chun)饲(si)粮的情况下,每(mei)天体(ti)内(nei)产生300mg的胆(dan)固(gu)醇(chun),其中大约(yue)2/3沉(chen)积在鸡蛋中[24],其次为粪便和胆(dan)酸代谢(xie)[25],部(bu)分(fen)胆(dan)固(gu)醇(chun)用于(yu)合成类固(gu)醇(chun)激素、维生素D3以及细胞膜。 在公鸡(ji)和未性成(cheng)(cheng)(cheng)熟母鸡(ji)体内,肝脏(zang)合成(cheng)(cheng)(cheng)的胆(dan)固(gu)醇(chun)(chun)载脂蛋(dan)白(bai)(bai)主要为极低(di)密(mi)度(du)(du)脂蛋(dan)白(bai)(bai)(VLDL)和高(gao)密(mi)度(du)(du)脂蛋(dan)白(bai)(bai)(HDL),VLDL到达血(xue)液(ye)后(hou)代谢成(cheng)(cheng)(cheng)为低(di)密(mi)度(du)(du)脂蛋(dan)白(bai)(bai)(LDL),因此血(xue)液(ye)中胆(dan)固(gu)醇(chun)(chun)主要以高(gao)密(mi)度(du)(du)脂蛋(dan)白(bai)(bai)胆(dan)固(gu)醇(chun)(chun)(HDLC)和低(di)密(mi)度(du)(du)脂蛋(dan)白(bai)(bai)胆(dan)固(gu)醇(chun)(chun)(LDLC)的形式(shi)存在[26]。 母(mu)鸡性成(cheng)熟以(yi)后,卵巢分(fen)泌(mi)的大量(liang)雌(ci)激素刺激肝细胞(bao)分(fen)泌(mi)VLDL小颗粒(现一般称作VLDLy,其直径(jing)小,甘(gan)油三(san)酯和胆固醇含(han)量(liang)高)[27]、卵黄蛋白原(VTG)[28]以(yi)及HDLC。 此时(shi),血液中胆固醇主要(yao)通过载(zai)(zai)(zai)体蛋(dan)白VLDLy运输到(dao)卵巢(chao),通过胞吞作(zuo)用进入卵黄,随蛋(dan)黄排(pai)出体外。公鸡和(he)未性(xing)成熟母鸡的VLDL的主要(yao)载(zai)(zai)(zai)体蛋(dan)白为(wei)载(zai)(zai)(zai)脂(zhi)(zhi)蛋(dan)白B(ApoB)和(he)载(zai)(zai)(zai)脂(zhi)(zhi)蛋(dan)白(ApoC),能(neng)够激活(huo)脂(zhi)(zhi)蛋(dan)白酯(zhi)酶(mei)(LPL)。 产蛋鸡VLDLy的主(zhu)要(yao)组(zu)成载体蛋白(bai)(bai)是ApoBApoVLDLⅡ[29]。ApoVLDLⅡ的相对分子(zi)(zi)质量为18,是一个小分子(zi)(zi)载脂蛋白(bai)(bai),能够(gou)抑制(zhi)LPL的活性,防止VLDLy被LPL降解[29]。 同时,ApoVLDLⅡ可(ke)以(yi)控制(zhi)VLDLy在形(xing)成过程中的大小(xiao),保(bao)证形(xing)成的VLDLy直径小(xiao)于30nm,从(cong)而顺利通过卵细胞外的基底层[29-30]。 此外,Hummel等[31]研究发(fa)现ApoVLDLⅡ可能具有(you)阻止(zhi)VLDLy与(yu)LDL受体(LDLR)结(jie)合的(de)(de)作用(yong)。VLDLy和VTG经过血液运输到达卵巢后,与(yu)卵巢膜上的(de)(de)受体LR8结(jie)合,经胞吞(tun)作用(yong)进入卵细胞[32]。 另有研(yan)究表(biao)明,LDLR相(xiang)关(guan)蛋白LRP380在胞吞(tun)过程中发挥了重(zhong)要作(zuo)用(yong),其可能与LR8在卵母(mu)细胞的最后(hou)生长(zhang)阶段具有协同作(zuo)用(yong)[33]。 研(yan)究表明,蛋黄中95%的胆(dan)固(gu)醇(chun)来(lai)自VLDLy,5%来(lai)自VTG[34],且蛋黄中90%以上的胆(dan)固(gu)醇(chun)以游离形式存在[35]。 2益生菌可(ke)调节蛋鸡胆固醇代谢益生菌对机(ji)(ji)体能(neng)够产生有益的作用[36]。以往研究(jiu)表(biao)明,微生物饲料添加剂具(ju)有维护动物肠(chang)道(dao)健康[37-39]、缓解不(bu)良应激[39]、改善畜舍环境(jing)[40-41]、调节机(ji)(ji)体脂肪(fang)代谢[42-44]、提(ti)高生产性能(neng) 和(he)改善(shan)畜产品品质[45]等(deng)功(gong)能。 研究表明,益生(sheng)菌在调(diao)节(jie)蛋鸡胆固醇代谢(xie)方面具有显著作用。 这些菌(jun)(jun)株主要有嗜酸乳杆菌(jun)(jun)、植物乳杆菌(jun)(jun)、瑞士乳杆菌(jun)(jun)、芽(ya)孢杆菌(jun)(jun)、屎肠球菌(jun)(jun)等[46]。 Mohan等[47]在饲(si)粮中(zhong)添(tian)加(jia)益生(sheng)菌产品(Probiolac)饲(si)喂28周龄白莱航鸡(ji),试验10周,发现其血清(qing)和蛋黄(huang)胆固醇含(han)量(liang)分别降低了27%和23%; Abdulrahim等[48]在饲(si)粮(liang)中添(tian)加嗜酸乳杆菌(jun)饲(si)喂20周(zhou)龄白壳蛋(dan)鸡,试验8周(zhou),发(fa)现(xian)其(qi)血浆和蛋(dan)黄胆(dan)固醇(chun)含量分别下降了56%和17%; Chen等(deng)[49]在饲(si)(si)粮中添加益生(sheng)菌产品(Ecozyme)饲(si)(si)喂27周(zhou)龄白(bai)莱航(hang)蛋鸡(ji),试验4周(zhou),发(fa)现益生(sheng)菌组蛋鸡(ji)的血清和蛋黄胆(dan)固(gu)醇(chun)含量(liang)分别下降了13.67%和11.31%; Xu等[50]在饲粮(liang)中添加(jia)枯草(cao)芽孢杆菌饲喂25周(zhou)龄(ling)罗曼(man)褐壳(qiao)蛋鸡,试验31周(zhou),发现蛋黄胆固醇含(han)量下降了12.21%; Panda等(deng)[51]在饲(si)粮中(zhong)添加(jia)益(yi)生(sheng)菌(jun)产品(乳杆(gan)菌(jun)、双歧(qi)杆(gan)菌(jun)等(deng))饲(si)喂(wei)24周(zhou)龄白莱航蛋(dan)鸡,试验48周(zhou),发现益(yi)生(sheng)菌(jun)组(zu)蛋(dan)鸡血(xue)清和(he)蛋(dan)黄胆(dan)固(gu)醇含量分别下降了14%和(he)19%; 随后(hou),Panda等[52]在饲(si)粮(liang)中添加芽孢乳杆(gan)菌饲(si)喂25周龄白莱航鸡,试验16周,发(fa)现其血浆和蛋(dan)黄胆固(gu)醇(chun)含量(liang)分别下降了20.2%和27.6%; Kurtoglu等[53]在饲粮中添(tian)加益生菌(jun)(jun)产品(芽孢杆菌(jun)(jun))饲喂(wei)27周龄褐壳蛋(dan)鸡(ji),试(shi)验90d,发现试(shi)验30d时,其(qi)(qi)血清和(he)(he)蛋(dan)黄(huang)胆固醇含(han)量无显著性差异,至90d时,其(qi)(qi)血清和(he)(he)蛋(dan)黄(huang)胆固醇含(han)量分别下降了40.8%和(he)(he)37.2%; Haddadin等[54]在饲粮中(zhong)添加嗜(shi)酸乳杆(gan)菌饲喂25周(zhou)龄的白莱航蛋(dan)鸡,试(shi)验(yan)48周(zhou),前40周(zhou)试(shi)验(yan)组(zu)饲喂试(shi)验(yan)饲粮,最后(hou)8周(zhou)试(shi)验(yan)组(zu)饲喂基础(chu)饲粮,发(fa)现(xian)试(shi)验(yan)进行(xing)到(dao)40周(zhou)时蛋(dan)鸡血浆(jiang)和(he)蛋(dan)黄胆(dan)固(gu)醇(chun)含量(liang)(liang)分别下(xia)(xia)降(jiang)了(le)55.0%和(he)18.8%,但(dan)当(dang)试(shi)验(yan)进行(xing)到(dao)44周(zhou)时血浆(jiang)胆(dan)固(gu)醇(chun)含量(liang)(liang)下(xia)(xia)降(jiang)了(le)24.0%,蛋(dan)黄胆(dan)固(gu)醇(chun)含量(liang)(liang)与对照组(zu)差(cha)异(yi)不显著,至48周(zhou)时,血浆(jiang)和(he)蛋(dan)黄胆(dan)固(gu)醇(chun)含量(liang)(liang)均(jun)无显著性变化(hua); Mahdavi等[55]在饲粮中添加(jia)益生菌产品(枯草芽(ya)孢杆菌和地(di)衣芽(ya)孢杆菌)饲喂28周(zhou)龄(ling)白莱航鸡,试验12周(zhou),发现其血浆胆(dan)固醇含(han)量(liang)下降了15.4%,但(dan)蛋黄(huang)胆(dan)固醇含(han)量(liang)无(wu)显著性差异; Kalavathy等[56]在饲粮中添加乳酸(suan)菌饲喂20周龄罗曼褐壳蛋鸡,试验48周,发(fa)现24和(he)28周龄时蛋黄(huang)(huang)胆固醇(chun)含(han)量分别下降了15.3%和(he)10.8%,但(dan)是(shi)32周龄时蛋黄(huang)(huang)胆固醇(chun)含(han)量无显著性变化; Choe等[57]在饲粮中添(tian)加(jia)植物乳杆菌代(dai)(dai)谢物饲喂19周(zhou)龄褐壳(qiao)蛋(dan)鸡(ji),试(shi)验12周(zhou),发(fa)现(xian)菌代(dai)(dai)谢物添(tian)加(jia)量(liang)(liang)为1.2%的(de)组血(xue)浆和蛋(dan)黄胆(dan)固(gu)醇含量(liang)(liang)分别下降了32.7%和4.9%,而(er)菌代(dai)(dai)谢物添(tian)加(jia)量(liang)(liang)为0.6%的(de)组血(xue)浆和蛋(dan)黄胆(dan)固(gu)醇含量(liang)(liang)分别下降了18.1%和10.6%。 上述研究表明,蛋(dan)鸡饲粮中添(tian)加益生(sheng)菌可以降低血(xue)清和蛋(dan)黄(huang)中胆(dan)固醇(chun)(chun)的含量,但降低程度(du)有所差(cha)别。这可能(neng)与(yu)试验所用(yong)益生(sheng)菌菌株、添(tian)加浓度(du)、蛋(dan)鸡品种以及试验周期等因素(su)有关。菌株本身存(cun)在极大(da)的差(cha)异,尤其(qi)是分泌胆(dan)盐水解酶的能(neng)力存(cun)在差(cha)异导致对胆(dan)固醇(chun)(chun)的作用(yong)不同; 益生菌调(diao)节蛋鸡胆(dan)(dan)固醇(chun)代谢具有最佳剂量(liang)范围,剂量(liang)过低或过高(gao)对(dui)机体(ti)胆(dan)(dan)固醇(chun)代谢无显(xian)著影响;试(shi)验周期过长,蛋鸡可能(neng)利用(yong)自身(shen)对(dui)胆(dan)(dan)固醇(chun)的调(diao)节能(neng)力而使(shi)机体(ti)胆(dan)(dan)固醇(chun)恢复到正(zheng)常(chang)水平。 另(ling)外(wai),需要注意(yi)的(de)(de)是结(jie)果(guo)中胆固醇(chun)的(de)(de)表(biao)(biao)示方式(shi),以上研究除了Mohan等(deng)[47]、Kurtoglu等(deng)[53]、Choe等(deng)[57]的(de)(de)研究外(wai),其他研究结(jie)果(guo)均没有涉及蛋黄重量这(zhei)一(yi)指标(biao),蛋黄胆固醇(chun)是以浓度(du)(mg/g蛋黄)的(de)(de)形(xing)式(shi)表(biao)(biao)示的(de)(de),在这(zhei)种情况(kuang)下,试验结(jie)果(guo)难以准确评价(jia)益生菌(jun)对鸡蛋胆固醇(chun)的(de)(de)降低(di)能(neng)力。 3益生(sheng)菌(jun)调节胆固醇的(de)可能机(ji)理关于(yu)益(yi)生菌降低胆固(gu)醇(chun)的机(ji)制,国内外(wai)学者进行了(le)大量的研究,研究结果分述如下(xia)。 3.1益(yi)生菌可水解肠道内胆盐胆(dan)盐(yan)(yan)是以(yi)胆(dan)固(gu)醇为前体在肝(gan)脏中合(he)成的(de)水(shui)溶性物(wu)质,经过胆(dan)囊储存和浓(nong)缩(suo),最(zui)终排入十二指肠内促进(jin)脂类物(wu)质的(de)消化和吸收。胆(dan)汁一般由(you)胆(dan)固(gu)醇、结合(he)胆(dan)盐(yan)(yan)、磷脂、色素和电(dian)解质组(zu)成。胆(dan)盐(yan)(yan)水(shui)解酶(bilesalthydrolase,BSH)为N末(mo)端亲核(he)水(shui)解酶,能(neng)特异性地水(shui)解结合(he)胆(dan)盐(yan)(yan)的(de)酰胺键,释放出游离(li)胆(dan)盐(yan)(yan)和甘氨酸(suan)或牛磺酸(suan)的(de)氨基(ji)酸(suan)残基(ji)[58]。 胆(dan)盐水解(jie)酶的(de)主(zhu)要作用部(bu)位(wei)(wei)在哺乳动(dong)物的(de)小(xiao)肠(chang)(chang)(chang)和大肠(chang)(chang)(chang),更为确切的(de)部(bu)位(wei)(wei)因动(dong)物的(de)种(zhong)类而(er)(er)异,如小(xiao)鼠(shu)的(de)主(zhu)要代(dai)谢(xie)部(bu)位(wei)(wei)在小(xiao)肠(chang)(chang)(chang),而(er)(er)人(ren)的(de)主(zhu)要代(dai)谢(xie)部(bu)位(wei)(wei)在回肠(chang)(chang)(chang),胆(dan)盐水解(jie)最终在大肠(chang)(chang)(chang)完成[59]。 研(yan)究(jiu)表明(ming),乳(ru)酸菌属、双(shuang)歧(qi)杆菌属、肠(chang)球(qiu)菌属、梭菌属和拟(ni)杆菌属等细(xi)菌均可分泌(mi)胆盐水解酶,具有水解结合胆盐的能力[58]。 每次(ci)肝肠(chang)循环中,大约有95%的结合胆盐(yan)在(zai)回肠(chang)被(bei)重新吸收。 游离胆盐与结合胆盐相(xiang)比(bi),其溶解度较低(di),不容(rong)易被肠道重新吸收(shou)从而随粪便排出体(ti)外[60],如图2[61]所(suo)示。 研(yan)究者(zhe)推测,机体为了(le)维持正常的(de)(de)肝肠(chang)循环,弥补胆盐损失,肝脏会利用血(xue)液中的(de)(de)胆固(gu)醇(chun)合成新的(de)(de)胆盐来补充排泄掉的(de)(de)那部(bu)分,从而引起血(xue)清(qing)胆固(gu)醇(chun)含量的(de)(de)降低[62-63]。 此外,体外试验表明,在(zai)pH低(di)(di)于(yu)5.5时,游(you)离胆(dan)盐(yan)(yan)与(yu)胆(dan)固(gu)醇发生(sheng)(sheng)共沉(chen)淀(dian)现象,降低(di)(di)肠(chang)(chang)道内(nei)胆(dan)固(gu)醇的吸收几率(lv)[46]。但事实上,蛋(dan)鸡(ji)(ji)肠(chang)(chang)道内(nei)pH通(tong)常介于(yu)7.39~7.53[4],因(yin)此共沉(chen)淀(dian)在(zai)蛋(dan)鸡(ji)(ji)肠(chang)(chang)道内(nei)很(hen)难发生(sheng)(sheng)。但是,Ramasamy等[65]从(cong)鸡(ji)(ji)肠(chang)(chang)道内(nei)分离得到12株乳酸菌,经体外研究(jiu)发现,12株乳酸菌都可(ke)以(yi)不同程度地分解甘氨酸胆(dan)盐(yan)(yan)(16.87%~100.00%)和(he)牛磺(huang)酸胆(dan)盐(yan)(yan)(1.69%~57.43%),具有清除培养(yang)基中胆(dan)固(gu)醇的能(neng)力。 3.2益生菌可吸收(shou)或吸附肠道(dao)内的胆固醇益生菌除了通(tong)过胆(dan)盐(yan)水解(jie)酶的解(jie)离作用降低机体胆(dan)固醇外,还(hai)可以通(tong)过另(ling)外一些(xie)途径发挥相(xiang)同(tong)的作用,这些(xie)途径包括: 1)益生菌细胞质同化胞外胆固醇; 2)益(yi)生(sheng)菌细(xi)胞(bao)膜(mo)整(zheng)合胞(bao)外胆固醇; 3)益生菌(jun)细胞(bao)壁(bi)吸附(fu)胞(bao)外胆固醇(chun); 4)以上3种途径的结合方(fang)式[46]。 一般认为,益(yi)生菌(jun)(jun)细胞质(zhi)同化和(he)细胞膜整(zheng)合胞外胆固醇(chun)的前(qian)提是益(yi)生菌(jun)(jun)必须为活菌(jun)(jun)。 Gilliland等[67]发现从猪粪便分离得到的嗜酸性乳杆菌RP32具有降低培养基中胆固醇的能力,以及抑制猪血清胆固醇含量的升高,进一步分离菌体细胞发现细胞内的胆固醇含量与对照组相比显著升高。 Noh等[68]发现生长中(zhong)的嗜酸性乳(ru)杆菌ATCC43121吸收胆(dan)固(gu)醇后并没有将胆(dan)固(gu)醇降解,而是(shi)将完整(zheng)的胆(dan)固(gu)醇整(zheng)合到了(le)细胞膜中(zhong)。 Liong等(deng)[69]报道当培养基中添加胆固(gu)醇(chun)后,细胞膜的(de)脂肪(fang)酸组成(cheng)会发生变化(hua),不饱(bao)和(he)脂肪(fang)酸的(de)含量(liang)(liang)上升(sheng),饱(bao)和(he)脂肪(fang)酸的(de)含量(liang)(liang)下降。 Lye等[70]利用(yong)荧光探针标(biao)记(ji)菌(jun)体(ti)磷脂双(shuang)分子层,发现胆(dan)固醇(chun)在细胞膜有富集现象,进一(yi)步证(zheng)明了益生菌(jun)可以(yi)通过细胞膜吸收胆(dan)固醇(chun)。 研究表明,益生菌能将肠道内(nei)胆固醇转化为肾固醇,而后者可随粪便排出(chu)体外(wai)[72]。 Chiang等[72]发现硝化细菌(jun)(Sterolibacteriumdenitrificans)可通过分泌(mi)胆(dan)固(gu)醇(chun)脱氢酶(mei)或异构酶(mei),将胆(dan)固(gu)醇(chun)转(zhuan)化为肾(shen)固(gu)醇(chun)的前体物质胆(dan)固(gu)-4-烯-3-酮。 Lye等(deng)[73]发(fa)现嗜酸乳杆菌、保(bao)加利(li)亚乳杆菌和干酪乳杆菌可以通过分泌胆固(gu)醇还(hai)原酶(mei)降低培(pei)养(yang)基中胆固(gu)醇的含量,提(ti)高培(pei)养(yang)基中肾固(gu)醇的浓度。 3.4益(yi)生菌(jun)可提高(gao)肠(chang)道内丙酸和丁酸含量益(yi)生菌(jun)代谢产(chan)物在(zai)肠(chang)(chang)道内通过发酵(jiao)肠(chang)(chang)道内容物产(chan)生短(duan)链脂(zhi)肪酸,如(ru)丙酸、丁酸等(deng)。丙酸可(ke)以抑(yi)制肝脏脂(zhi)肪酸和胆固(gu)醇合成,降低甘油三酯(zhi)和胆固(gu)醇的分泌速率,丁酸同样可(ke)以抑(yi)制肝脏胆固(gu)醇合成[74]。 4小结(jie) 综上所述,益(yi)(yi)生(sheng)(sheng)(sheng)菌(jun)可以(yi)降(jiang)低蛋(dan)鸡血清(qing)(血浆)和(he)蛋(dan)黄中胆(dan)固醇的(de)(de)含量。研(yan)究人(ren)员提出了很多益(yi)(yi)生(sheng)(sheng)(sheng)菌(jun)降(jiang)胆(dan)固醇的(de)(de)可能机(ji)理(li),但是这些(xie)机(ji)理(li)的(de)(de)主要依据来源于体(ti)(ti)外试(shi)验(yan)(yan)结果,而(er)且缺乏体(ti)(ti)内的(de)(de)验(yan)(yan)证试(shi)验(yan)(yan)结果。由于益(yi)(yi)生(sheng)(sheng)(sheng)菌(jun)在体(ti)(ti)内和(he)体(ti)(ti)外的(de)(de)生(sheng)(sheng)(sheng)长模(mo)式差异较大,所以(yi)只(zhi)有将体(ti)(ti)内和(he)体(ti)(ti)外试(shi)验(yan)(yan)相结合,才能清(qing)楚地(di)了解益(yi)(yi)生(sheng)(sheng)(sheng)菌(jun)调节胆(dan)固醇的(de)(de)真正机(ji)制,并以(yi)此指导益(yi)(yi)生(sheng)(sheng)(sheng)菌(jun)在动物和(he)人(ren)营养方面(mian)的(de)(de)进(jin)一步应用。 参考文献:[1] 汪仕奎,佟建明.蛋鸡的胆(dan)固醇代谢调(diao)控(kong)研究(jiu)进展 [J].动(dong)物营养学报,2002,14(3):7-11. 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