在高植物蛋白饲料中添加胆固醇、牛磺酸和大豆皂甙对大菱鲆生长性能和胆固醇代谢的影响
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摘要
以我国重要海水肉食性养殖鱼类大菱鲆（Scophthalmus maximus L.）为实验对象，在室内流水养殖系统中进行摄食生长实验，研究高植物蛋白饲料中添加胆固醇、牛磺酸和大豆皂甙以及限饲水平对大菱鲆生长性能和胆固醇代谢的影响。在此基础上探讨植物蛋白降低鱼类血浆胆固醇的机制。本文的研究内容和主要研究结果如下：
1、高植物蛋白饲料中添加胆固醇对大菱鲆幼鱼摄食、生长和胆固醇代谢的影响
用添加胆固醇的高植物蛋白饲料饲喂大菱鲆（Scophthalmus maximus L.）幼鱼9周，研究其对大菱鲆摄食、生长和胆固醇代谢的影响。实验设计了5组等氮等脂饲料，以含鱼粉58%的饲料作为基础饲料（FM），而其它4组饲料均含14.5%的鱼粉、42.0%豆粕和18.5%谷朊粉，在这4组饲料中分别补充0.0%（C-0.0%）、0.5%（C-0.5%）、1.0%（C-1.0%）和1.5%（C-1.5%）的胆固醇，其实测值分别为0.30%、0.77%、1.25%和1.78%，而基础饲料中胆固醇检测值为0.63%。结果表明，摄食饲料FM的大菱鲆增重率和饲料效率均显著高于其它饲料组（P<0.05）。与C-0.0%组相比，摄食饲料C-1.0%组显著提高了大菱鲆幼鱼的摄食率、增重率和血浆及肝脏胆固醇水平（P<0.05）。与C-1.0%组相比，摄食饲料C-1.5%组显著抑制了实验鱼的增重率和摄食率（P<0.05）。当饲料中胆固醇含量为1.25%（C-1.0%）时，其全鱼脂肪含量最高，且显著高于其它处理组（P<0.05）。饲料中胆固醇含量与血浆中总胆固醇、游离胆固醇、胆固醇酯、高密度脂蛋白胆固醇和低密度脂蛋白胆固醇的含量、肝脏中总胆固醇、胆固醇酯的含量及粪便中总胆固醇的含量呈显著相关关系，相关系数均大于0.74。饲料中含有0.30%的胆固醇（C-0.0%），其粪便总胆汁酸含量和肝脏胆固醇7A羟化酶的活性在所有处理组中最低。该研究结果表明，高植物蛋白饲料中含有1.25%的胆固醇有利于大菱鲆幼鱼的生长，且对其生理状态无不良影响。
2、大菱鲆HMG-CoAr基因的克隆、序列分析以及胆固醇添加到高植物蛋白饲料中对肝脏HMG-CoAr基因mRNA表达量的影响
以大菱鲆（Scophthalmus maximus L.）为实验材料，利用同源克隆技术和RACE技术从大菱鲆肝脏中成功扩增出3-羟基-3-甲基戊二酸单酰辅酶A还原酶（3-hydroxy-3-methylglutaryl-Coenzyme A reductase，HMG-CoAr）cDNA的全序列。HMG-CoAr基因cDNA全长序列为3048bp（Genbank No: JN542428），其中包括一个有2664bp的开放阅读框且编码887个氨基酸，预测分子量约为96.52kDa，理论等电点为6.44。BLAST分析显示大菱鲆HMG-CoAr基因与其它已知脊椎动物具有高度同源性，且N-端和C-端高度保守，仅连接N-端和C-端的中间疏水部分变异较大。序列分析指出HMG-CoAr基因在N-端氨基酸部分包括5个主要的跨膜域。实时定量PCR技术检测了全鱼粉组（FM）、高植物蛋白组（C-0.0%）和高植物蛋白组基础饲料中添加1.0%的胆固醇（C-1.0%）对大菱鲆肝脏HMG-CoAr基因mRNA表达量的影响。结果表明，与摄食全鱼粉组相比（DietFM），摄食高植物蛋白饲料（C-0.0%）能显著上调大菱鲆肝脏HMG-CoAr基因mRNA的表达（P<0.01）。然而，当高植物蛋白饲料中补充1.0%的胆固醇则显著抑制肝脏HMG-CoAr基因mRNA的表达量（P<0.01）。该研究结果表明，充足的饲料胆固醇能够显著抑制HMG-CoAr基因mRNA的表达，胆固醇的合成受反馈调节机制控制。
3、高植物蛋白饲料中添加胆固醇和牛磺酸对大菱鲆幼鱼摄食、生长、胆固醇代谢以及肝脏HMG-CoAr基因mRNA表达量的影响
在高植物蛋白饲料中添加胆固醇、牛磺酸或同时添加胆固醇和牛磺酸研究其对大菱鲆（Scophthalmus maximus L.）幼鱼生长性能和胆固醇代谢的影响。实验基础饲料（对照组）包括14.5%的鱼粉和50%的粗蛋白，在基础饲料中分别添加1.0%牛磺酸、1.0%胆固醇及1.0%胆固醇和1.0%牛磺酸的混合物（各组饲料分别命名为C-1.0%、T-1.0%和TC），配制成等氮等脂的实验饲料。实验结果表明摄食饲料C-1.0%、T-1.0%和TC的大菱鲆增重率显著高于对照组。其中摄食饲料TC的大菱鲆增重率最高，但与C-1.0%相比无显著差异（P>0.05）。与摄食饲料C-1.0%相比，摄食饲料TC显著降低了大菱鲆血浆总胆固醇、游离胆固醇和低密度脂蛋白胆固醇含量（P<0.05）。与对照组相比，摄食饲料C-1.0%能显著提高实验鱼胆固醇7A羟化酶（CYP7A1）的活性（P<0.05），且摄食饲料T-1.0%组CYP7A1活性最高。与摄食饲料C-1.0%相比，摄食饲料TC能显著下调HMG-CoAr基因mRNA的表达量（P<0.05）。该研究结果表明，高植物蛋白饲料中添加1.0%胆固醇和1.0%的牛磺酸有利于大菱鲆的生长。
4、饲料中添加大豆皂甙对大菱鲆幼鱼摄食、生长和胆固醇代谢的影响
本实验旨在研究大豆皂甙对大菱鲆幼鱼（Scophthalmus maximus L.）摄食、生长和胆固醇代谢的影响。以鱼粉饲料为基础饲料（对照组，不添加大豆皂甙），分别在对照饲料基础上添加0.3%（实际含量为0.25%）和0.6%（实际含量为0.5%）的大豆皂甙产品，配制出另外的2组实验饲料。3组饲料大豆皂甙的含量分别为0.0%（Diet1）、0.25%（Diet2）和0.5%（Diet3）。结果表明，与对照组相比，饲料中添加0.25%的大豆皂甙（Diet2）对实验鱼的摄食率、饲料利用率和生长无显著影响。但与对照组相比，饲料中添加0.5%的大豆皂甙显著抑制了实验鱼的生长（P<0.05）。血浆、肝脏和粪便胆固醇代谢相关指标在各处理组间均无显著差异。以上结果表明在鱼粉为基础的饲料中添加0.25%大豆皂甙对大菱鲆幼鱼的生长没有负面效应，而添加0.5%大豆皂甙则显著抑制大菱鲆幼鱼的生长。各组间实验鱼血浆、肝脏和粪便胆固醇含量无显著差异，这表明在鱼粉为基础的饲料中添加大豆皂甙对实验鱼胆固醇代谢无显著影响。
5、限饲水平对大菱鲆幼鱼生长性能、胆固醇代谢以及肝脏HMG-CoAr基因表达的影响
以初始体重为6.4±0.1g的大菱鲆幼鱼为实验研究对象，探讨限饲水平对大菱鲆幼鱼生长性能和胆固醇代谢的影响。实验分为4组，对照组以表观饱食投喂，其它3组摄食率相应调整为表观饱食的75%、50%和25%，并分别命名为100%组、75%组、50%组和25%组。实验结果表明，摄食率与大菱鲆增重率、血浆总胆固醇、游离胆固醇、高密度脂蛋白胆固醇和低密度脂蛋白胆固醇呈显著的正相关关系（P<0.05）。而摄食率与肝脏总胆固醇、游离胆固醇及胆固醇酯呈显著负相关关系（P<0.05）。与对照组相比，25%组的酰基辅酶A：胆固醇酰基转移酶（ACAT）活性显著降低（P<0.05），且其活性与摄食率呈显著正相关关系（P<0.05）。与对照组相比，摄食表观饱食50%或25%的实验组显著上调了3-羟基-3-甲基戊二酰-辅酶A还原酶（HMG-CoAr）基因的mRNA表达。以上结果表明，植物蛋白所致的鱼类血浆低胆固醇效应部分归因于其摄食植物蛋白后导致的摄食率下降。在50%或25%组中、显著升高的实验鱼肝脏胆固醇含量可能与其胆固醇合成能力升高有关。
Turbot (Scophthalmus maximus L.), an important commercial carnivorous fishin East Asia, is widely farmed. Four growth trials were conducted to investigate theeffects of dietary cholesterol, taurine, soya saponins supplementation and feedrestriction on growth performance, feed intake and cholesterol metabolism of juvenileturbot. The aim of these experiments was to study the mechanism ofhypocholesterolemic effect caused by plant protein and to clarify the mechanism ofthe fish growth promoting effects of cholesterol.The results are summarized asfollows:
1. Effects of dietary cholesterol on growth performance, feed intake andcholesterol metabolism in juvenile turbot (Scophthalmus maximus L.) fedhigh plant protein diets
A9-week growth trial was conducted to investigate the effects of dietarycholesterol supplementation on growth performance, feed intake and cholesterolmetabolism of juvenile turbot (Scophthalmus maximus L.) fed high plant proteindiets. A fish meal diet (diet FM) with58%FM was formulated, and this diet was usedas control. The other four isonitrogenous and isolipidic diets were formulated with0.0%,0.5%,1.0%and1.5%cholesterol supplementation, respectively, to the diet FM,whichcontained14.5%FM,42.0%soybean meal (SBM), and18.5%wheat glutenmeal. They were named as diet C-0.0%, C-0.5%, C-1.0%and C-1.5%, respectively.The final dietary cholesterol concentrations were0.30%,0.77%,1.25%, and1.78%,respectively, and that in control diet was0.63%. The results showed that weight gainrate (WGR) and feed efficiency rate in fish fed the diet FM were significantly higherthan those in fish fed other diets (P<0.05). Furthermore, compared with fish fed diet C-0.0%, fish fed diet C-1.0%significantly enhanced WGR, feed intake (FI) andcholesterol levels in plasma and liver. However, WGR and FI in fish fed diet C-1.5%were significantly lower than those in fish fed diet C-1.0%(P<0.05). Fish fed dietC-1.0%showed significantly higher whole-body lipid content than that of fish fedother diets (P<0.05). The total cholesterol (TC), free cholesterol (FC), cholesterolesters, high-density lipoprotein cholesterol (HDL-C) and low-density lipoproteincholesterol (LDL-C) contents in fish plasma, TC and cholesterol esters in fish liver,and TC in fish faeces were significantly correlated with dietary cholesterol contents,and correlated coefficients were above0.74.Fish fed diet C-0.0%showed the lowestfecal total bile acid and activity of cholesterol7α hydroxylase among dietarytreatments. These results suggested that1.25%of dietary cholesterol is helpful forjuvenile turbot fed high plant protein diets to get significantly better growth ratewithout negative effects.
2. Molecular cloning, characterization and mRNA expression of3-hydroxy-3-methylglutaryl-Coenzyme A reductase from turbotScophthalmus maximus L in response to dietary cholesterol
In the present study, the cDNA of3-hydroxy-3-methylglutaryl-Coenzyme Areductase (HMG-CoAr) from turbot Scophthalmus maximus L was cloned byhomology cloning with degenerate primer and RACE techniques. Thefull-length cDNA of HMG-CoAr was of3048bp, including an open reading frame(ORF) of2664bp encoding a polypeptide887amino acids with predicted molecularweight of96.52kDa and theoretical isoelectric point of6.44. BLAST analysisrevealed that HMG-CoAr shared high similarity with other known HMG-CoAr, andthe N-and C-terminus amino acids portions are the most conserved and are linked byhydrophilic region that appears to be quite variable. The expression levels ofHMG-CoAr in hepatic tissue was measured by real-time PCR after turbot were feddiets FM, C-0.0%and C-1.0%containing graded levels of cholesterol (0.63,0.30and1.25%) for9weeks. The results showed that the expression levels of HMG-CoAr transcript were significantly up-regulated in hepatic tissue of turbot fed diet C-0.0%(0.30%cholesterol) compared with that of fish fed diet FM (0.63%cholesterol)(p<0.01). However, these levels significantly decreased in hepatic tissue when fishwas fed diet C-1.0%(1.25%cholesterol)(p<0.01). These results indicated thatadequate dietary cholesterol could inhibit the mRNA expression of HMG-CoAr,suggesting that biosynthesis of cholesterol is controlled by a feedback mechanism inturbot.
3. Synergistic effects of dietary cholesterol and taurine on growth performanceand cholesterol metabolism in juvenile turbot (Scophthalmus maximus L.)fed high plant protein diets
The study was conducted to investigate the effects of cholesterol, taurine andcombination of dietary cholesterol and taurine on growth performance and cholesterolmetabolism in juvenile turbot (Scophthalmus maximus L.) fed high plant proteindiets. A basal diet (control) with14.5%fish meal and50%crude protein wasformulated. The other three isonitrogenous and isolipidic experimental diets wereprepared with the supplementation of1.0%cholesterol,1.0%taurine and thecombination of1.0%cholesterol and1.0%taurine to the basal diet, which werenamed as C-1.0%, T-1.0%and TC, respectively. The results showed that the weightgain rate in fish fed C-1.0%, T-1.0%and TC diets was significantly higher than that infish fed the control diet. Especially, fish fed TC diet showed the highest growthperformance in data among dietary treatments. The plasma total cholesterol, freecholesterol and low-density lipoprotein cholesterol levels were significantly lower infish fed TC diet compared to those in fish fed C-1.0%diet (P<0.05). Fish fed C-1.0%diet showed significantly higher activity of cholesterol7α hydroxylase (CYP7A1)than the control diet (P<0.05), and activity of CYP7A1in fish fed T-1.0%diet was thehighest among dietary treatments. The HMG-CoAr mRNA levels were significantlylower in fish fed TC diet compared to that in fish fed C-1.0%diet (P<0.05). Theseresults suggested that combination of1.0%cholesterol and1.0%taurine is helpful for juvenile turbot fed high plant protein diets to obtain significantly better growthwithout negative effects.
4. Effects of soya saponins on feed intake, growth performance and cholesterolmetabolism in juvenile turbot (Scophthalmus maximus L)
An8-week growth trial was conducted to investigate the effects of soya saponinson feed intake, growth performance and cholesterol metabolism in juvenile turbot(Scophthalmus maximus L). A control diet was formulated without soya saponinssupplementation and other two experimental diets were prepared with thesupplementation of3.0g kg-1and6.0g kg-1soya saponins product to the control diet,respectively. The planned soya saponin concentrations in diets were0.0g kg-1(Diet1),2.5g kg-1(Diet2) and5.0g kg-1(Diet3), respectively. The results showed that2.5g kg-1of dietary soya saponins exhibited comparable feed intake, feed utilization andgrowth performance with the control diet (P<0.05). However, fish fed the diet with5.0g kg-1soya saponins significantly depressed growth. All the selected parameters inplasma, liver and feces of each group were not significantly different. These resultssuggested that2.5g kg-1of dietary soya saponins did not show negative effects, but5.0g kg-1of dietary soya saponins significantly depressed growth of fish. Dietarysoya saponins supplementation to FM-based diet did not show significant effect oncholesterol metabolism.
5. Effects of feed restriction on growth performance and cholesterol metabolismin juvenile turbot (Scophthalmus maximus L)
The study was conducted to investigate the effects of feed restriction on growthperformance and cholesterol metabolism in juvenile turbot Scophthalmus maximusL. Fish from the control group was fed to apparent satiation following adjustedfeeding rates. In the experimental treatments, the control group feeding rates wasreduced to75%,50%and25%, and the four diets were named as100%,75%,50%and25%, respectively. The results showed that weight gain rate (WGR) was linearly related to feed intake (FI)(P<0.05). Significant positive correlations were observedbetween the FI and some the selected plasma parameters, total cholesterol (TC), freecholesterol (FC), high-density lipoprotein cholesterol (HDL-C) and low-densitylipoprotein cholesterol (LDL-C)(P<0.05). The FI was significant negativecorrelations with the TC, FC and cholesterol esters in fish liver (P<0.05). Fish fed to25%of satiation showed significant low activity of acyl-coenzyme A: cholesterolacyltransferase (ACAT) compared with fish fed to the control group(P<0.05), and asignificant positive correlations was observed between FI and activity of ACAT(P<0.05). The expression levels of3-hydroxy-3-methylglutaryl-Coenzyme Areductase (HMG-CoAr) transcript were significantly up-regulated in liver of turbotfed fed to50%or25%of satiation compared with that in fish fed to the control group.These results suggested thatthe hypocholesterolemic effect was observed in plasma offish fed plant protein diets, which were related to, in part, significantly decreasedFI.Moreover, the increased hepatic TC level in fish fed to25%or50%of satiation may beattributed to the increased ability of cholesterol synthesis.
1、高植物蛋白饲料中添加胆固醇对大菱鲆幼鱼摄食、生长和胆固醇代谢的影响
用添加胆固醇的高植物蛋白饲料饲喂大菱鲆（Scophthalmus maximus L.）幼鱼9周，研究其对大菱鲆摄食、生长和胆固醇代谢的影响。实验设计了5组等氮等脂饲料，以含鱼粉58%的饲料作为基础饲料（FM），而其它4组饲料均含14.5%的鱼粉、42.0%豆粕和18.5%谷朊粉，在这4组饲料中分别补充0.0%（C-0.0%）、0.5%（C-0.5%）、1.0%（C-1.0%）和1.5%（C-1.5%）的胆固醇，其实测值分别为0.30%、0.77%、1.25%和1.78%，而基础饲料中胆固醇检测值为0.63%。结果表明，摄食饲料FM的大菱鲆增重率和饲料效率均显著高于其它饲料组（P<0.05）。与C-0.0%组相比，摄食饲料C-1.0%组显著提高了大菱鲆幼鱼的摄食率、增重率和血浆及肝脏胆固醇水平（P<0.05）。与C-1.0%组相比，摄食饲料C-1.5%组显著抑制了实验鱼的增重率和摄食率（P<0.05）。当饲料中胆固醇含量为1.25%（C-1.0%）时，其全鱼脂肪含量最高，且显著高于其它处理组（P<0.05）。饲料中胆固醇含量与血浆中总胆固醇、游离胆固醇、胆固醇酯、高密度脂蛋白胆固醇和低密度脂蛋白胆固醇的含量、肝脏中总胆固醇、胆固醇酯的含量及粪便中总胆固醇的含量呈显著相关关系，相关系数均大于0.74。饲料中含有0.30%的胆固醇（C-0.0%），其粪便总胆汁酸含量和肝脏胆固醇7A羟化酶的活性在所有处理组中最低。该研究结果表明，高植物蛋白饲料中含有1.25%的胆固醇有利于大菱鲆幼鱼的生长，且对其生理状态无不良影响。
2、大菱鲆HMG-CoAr基因的克隆、序列分析以及胆固醇添加到高植物蛋白饲料中对肝脏HMG-CoAr基因mRNA表达量的影响
以大菱鲆（Scophthalmus maximus L.）为实验材料，利用同源克隆技术和RACE技术从大菱鲆肝脏中成功扩增出3-羟基-3-甲基戊二酸单酰辅酶A还原酶（3-hydroxy-3-methylglutaryl-Coenzyme A reductase，HMG-CoAr）cDNA的全序列。HMG-CoAr基因cDNA全长序列为3048bp（Genbank No: JN542428），其中包括一个有2664bp的开放阅读框且编码887个氨基酸，预测分子量约为96.52kDa，理论等电点为6.44。BLAST分析显示大菱鲆HMG-CoAr基因与其它已知脊椎动物具有高度同源性，且N-端和C-端高度保守，仅连接N-端和C-端的中间疏水部分变异较大。序列分析指出HMG-CoAr基因在N-端氨基酸部分包括5个主要的跨膜域。实时定量PCR技术检测了全鱼粉组（FM）、高植物蛋白组（C-0.0%）和高植物蛋白组基础饲料中添加1.0%的胆固醇（C-1.0%）对大菱鲆肝脏HMG-CoAr基因mRNA表达量的影响。结果表明，与摄食全鱼粉组相比（DietFM），摄食高植物蛋白饲料（C-0.0%）能显著上调大菱鲆肝脏HMG-CoAr基因mRNA的表达（P<0.01）。然而，当高植物蛋白饲料中补充1.0%的胆固醇则显著抑制肝脏HMG-CoAr基因mRNA的表达量（P<0.01）。该研究结果表明，充足的饲料胆固醇能够显著抑制HMG-CoAr基因mRNA的表达，胆固醇的合成受反馈调节机制控制。
3、高植物蛋白饲料中添加胆固醇和牛磺酸对大菱鲆幼鱼摄食、生长、胆固醇代谢以及肝脏HMG-CoAr基因mRNA表达量的影响
在高植物蛋白饲料中添加胆固醇、牛磺酸或同时添加胆固醇和牛磺酸研究其对大菱鲆（Scophthalmus maximus L.）幼鱼生长性能和胆固醇代谢的影响。实验基础饲料（对照组）包括14.5%的鱼粉和50%的粗蛋白，在基础饲料中分别添加1.0%牛磺酸、1.0%胆固醇及1.0%胆固醇和1.0%牛磺酸的混合物（各组饲料分别命名为C-1.0%、T-1.0%和TC），配制成等氮等脂的实验饲料。实验结果表明摄食饲料C-1.0%、T-1.0%和TC的大菱鲆增重率显著高于对照组。其中摄食饲料TC的大菱鲆增重率最高，但与C-1.0%相比无显著差异（P>0.05）。与摄食饲料C-1.0%相比，摄食饲料TC显著降低了大菱鲆血浆总胆固醇、游离胆固醇和低密度脂蛋白胆固醇含量（P<0.05）。与对照组相比，摄食饲料C-1.0%能显著提高实验鱼胆固醇7A羟化酶（CYP7A1）的活性（P<0.05），且摄食饲料T-1.0%组CYP7A1活性最高。与摄食饲料C-1.0%相比，摄食饲料TC能显著下调HMG-CoAr基因mRNA的表达量（P<0.05）。该研究结果表明，高植物蛋白饲料中添加1.0%胆固醇和1.0%的牛磺酸有利于大菱鲆的生长。
4、饲料中添加大豆皂甙对大菱鲆幼鱼摄食、生长和胆固醇代谢的影响
本实验旨在研究大豆皂甙对大菱鲆幼鱼（Scophthalmus maximus L.）摄食、生长和胆固醇代谢的影响。以鱼粉饲料为基础饲料（对照组，不添加大豆皂甙），分别在对照饲料基础上添加0.3%（实际含量为0.25%）和0.6%（实际含量为0.5%）的大豆皂甙产品，配制出另外的2组实验饲料。3组饲料大豆皂甙的含量分别为0.0%（Diet1）、0.25%（Diet2）和0.5%（Diet3）。结果表明，与对照组相比，饲料中添加0.25%的大豆皂甙（Diet2）对实验鱼的摄食率、饲料利用率和生长无显著影响。但与对照组相比，饲料中添加0.5%的大豆皂甙显著抑制了实验鱼的生长（P<0.05）。血浆、肝脏和粪便胆固醇代谢相关指标在各处理组间均无显著差异。以上结果表明在鱼粉为基础的饲料中添加0.25%大豆皂甙对大菱鲆幼鱼的生长没有负面效应，而添加0.5%大豆皂甙则显著抑制大菱鲆幼鱼的生长。各组间实验鱼血浆、肝脏和粪便胆固醇含量无显著差异，这表明在鱼粉为基础的饲料中添加大豆皂甙对实验鱼胆固醇代谢无显著影响。
5、限饲水平对大菱鲆幼鱼生长性能、胆固醇代谢以及肝脏HMG-CoAr基因表达的影响
以初始体重为6.4±0.1g的大菱鲆幼鱼为实验研究对象，探讨限饲水平对大菱鲆幼鱼生长性能和胆固醇代谢的影响。实验分为4组，对照组以表观饱食投喂，其它3组摄食率相应调整为表观饱食的75%、50%和25%，并分别命名为100%组、75%组、50%组和25%组。实验结果表明，摄食率与大菱鲆增重率、血浆总胆固醇、游离胆固醇、高密度脂蛋白胆固醇和低密度脂蛋白胆固醇呈显著的正相关关系（P<0.05）。而摄食率与肝脏总胆固醇、游离胆固醇及胆固醇酯呈显著负相关关系（P<0.05）。与对照组相比，25%组的酰基辅酶A：胆固醇酰基转移酶（ACAT）活性显著降低（P<0.05），且其活性与摄食率呈显著正相关关系（P<0.05）。与对照组相比，摄食表观饱食50%或25%的实验组显著上调了3-羟基-3-甲基戊二酰-辅酶A还原酶（HMG-CoAr）基因的mRNA表达。以上结果表明，植物蛋白所致的鱼类血浆低胆固醇效应部分归因于其摄食植物蛋白后导致的摄食率下降。在50%或25%组中、显著升高的实验鱼肝脏胆固醇含量可能与其胆固醇合成能力升高有关。
Turbot (Scophthalmus maximus L.), an important commercial carnivorous fishin East Asia, is widely farmed. Four growth trials were conducted to investigate theeffects of dietary cholesterol, taurine, soya saponins supplementation and feedrestriction on growth performance, feed intake and cholesterol metabolism of juvenileturbot. The aim of these experiments was to study the mechanism ofhypocholesterolemic effect caused by plant protein and to clarify the mechanism ofthe fish growth promoting effects of cholesterol.The results are summarized asfollows:
1. Effects of dietary cholesterol on growth performance, feed intake andcholesterol metabolism in juvenile turbot (Scophthalmus maximus L.) fedhigh plant protein diets
A9-week growth trial was conducted to investigate the effects of dietarycholesterol supplementation on growth performance, feed intake and cholesterolmetabolism of juvenile turbot (Scophthalmus maximus L.) fed high plant proteindiets. A fish meal diet (diet FM) with58%FM was formulated, and this diet was usedas control. The other four isonitrogenous and isolipidic diets were formulated with0.0%,0.5%,1.0%and1.5%cholesterol supplementation, respectively, to the diet FM,whichcontained14.5%FM,42.0%soybean meal (SBM), and18.5%wheat glutenmeal. They were named as diet C-0.0%, C-0.5%, C-1.0%and C-1.5%, respectively.The final dietary cholesterol concentrations were0.30%,0.77%,1.25%, and1.78%,respectively, and that in control diet was0.63%. The results showed that weight gainrate (WGR) and feed efficiency rate in fish fed the diet FM were significantly higherthan those in fish fed other diets (P<0.05). Furthermore, compared with fish fed diet C-0.0%, fish fed diet C-1.0%significantly enhanced WGR, feed intake (FI) andcholesterol levels in plasma and liver. However, WGR and FI in fish fed diet C-1.5%were significantly lower than those in fish fed diet C-1.0%(P<0.05). Fish fed dietC-1.0%showed significantly higher whole-body lipid content than that of fish fedother diets (P<0.05). The total cholesterol (TC), free cholesterol (FC), cholesterolesters, high-density lipoprotein cholesterol (HDL-C) and low-density lipoproteincholesterol (LDL-C) contents in fish plasma, TC and cholesterol esters in fish liver,and TC in fish faeces were significantly correlated with dietary cholesterol contents,and correlated coefficients were above0.74.Fish fed diet C-0.0%showed the lowestfecal total bile acid and activity of cholesterol7α hydroxylase among dietarytreatments. These results suggested that1.25%of dietary cholesterol is helpful forjuvenile turbot fed high plant protein diets to get significantly better growth ratewithout negative effects.
2. Molecular cloning, characterization and mRNA expression of3-hydroxy-3-methylglutaryl-Coenzyme A reductase from turbotScophthalmus maximus L in response to dietary cholesterol
In the present study, the cDNA of3-hydroxy-3-methylglutaryl-Coenzyme Areductase (HMG-CoAr) from turbot Scophthalmus maximus L was cloned byhomology cloning with degenerate primer and RACE techniques. Thefull-length cDNA of HMG-CoAr was of3048bp, including an open reading frame(ORF) of2664bp encoding a polypeptide887amino acids with predicted molecularweight of96.52kDa and theoretical isoelectric point of6.44. BLAST analysisrevealed that HMG-CoAr shared high similarity with other known HMG-CoAr, andthe N-and C-terminus amino acids portions are the most conserved and are linked byhydrophilic region that appears to be quite variable. The expression levels ofHMG-CoAr in hepatic tissue was measured by real-time PCR after turbot were feddiets FM, C-0.0%and C-1.0%containing graded levels of cholesterol (0.63,0.30and1.25%) for9weeks. The results showed that the expression levels of HMG-CoAr transcript were significantly up-regulated in hepatic tissue of turbot fed diet C-0.0%(0.30%cholesterol) compared with that of fish fed diet FM (0.63%cholesterol)(p<0.01). However, these levels significantly decreased in hepatic tissue when fishwas fed diet C-1.0%(1.25%cholesterol)(p<0.01). These results indicated thatadequate dietary cholesterol could inhibit the mRNA expression of HMG-CoAr,suggesting that biosynthesis of cholesterol is controlled by a feedback mechanism inturbot.
3. Synergistic effects of dietary cholesterol and taurine on growth performanceand cholesterol metabolism in juvenile turbot (Scophthalmus maximus L.)fed high plant protein diets
The study was conducted to investigate the effects of cholesterol, taurine andcombination of dietary cholesterol and taurine on growth performance and cholesterolmetabolism in juvenile turbot (Scophthalmus maximus L.) fed high plant proteindiets. A basal diet (control) with14.5%fish meal and50%crude protein wasformulated. The other three isonitrogenous and isolipidic experimental diets wereprepared with the supplementation of1.0%cholesterol,1.0%taurine and thecombination of1.0%cholesterol and1.0%taurine to the basal diet, which werenamed as C-1.0%, T-1.0%and TC, respectively. The results showed that the weightgain rate in fish fed C-1.0%, T-1.0%and TC diets was significantly higher than that infish fed the control diet. Especially, fish fed TC diet showed the highest growthperformance in data among dietary treatments. The plasma total cholesterol, freecholesterol and low-density lipoprotein cholesterol levels were significantly lower infish fed TC diet compared to those in fish fed C-1.0%diet (P<0.05). Fish fed C-1.0%diet showed significantly higher activity of cholesterol7α hydroxylase (CYP7A1)than the control diet (P<0.05), and activity of CYP7A1in fish fed T-1.0%diet was thehighest among dietary treatments. The HMG-CoAr mRNA levels were significantlylower in fish fed TC diet compared to that in fish fed C-1.0%diet (P<0.05). Theseresults suggested that combination of1.0%cholesterol and1.0%taurine is helpful for juvenile turbot fed high plant protein diets to obtain significantly better growthwithout negative effects.
4. Effects of soya saponins on feed intake, growth performance and cholesterolmetabolism in juvenile turbot (Scophthalmus maximus L)
An8-week growth trial was conducted to investigate the effects of soya saponinson feed intake, growth performance and cholesterol metabolism in juvenile turbot(Scophthalmus maximus L). A control diet was formulated without soya saponinssupplementation and other two experimental diets were prepared with thesupplementation of3.0g kg-1and6.0g kg-1soya saponins product to the control diet,respectively. The planned soya saponin concentrations in diets were0.0g kg-1(Diet1),2.5g kg-1(Diet2) and5.0g kg-1(Diet3), respectively. The results showed that2.5g kg-1of dietary soya saponins exhibited comparable feed intake, feed utilization andgrowth performance with the control diet (P<0.05). However, fish fed the diet with5.0g kg-1soya saponins significantly depressed growth. All the selected parameters inplasma, liver and feces of each group were not significantly different. These resultssuggested that2.5g kg-1of dietary soya saponins did not show negative effects, but5.0g kg-1of dietary soya saponins significantly depressed growth of fish. Dietarysoya saponins supplementation to FM-based diet did not show significant effect oncholesterol metabolism.
5. Effects of feed restriction on growth performance and cholesterol metabolismin juvenile turbot (Scophthalmus maximus L)
The study was conducted to investigate the effects of feed restriction on growthperformance and cholesterol metabolism in juvenile turbot Scophthalmus maximusL. Fish from the control group was fed to apparent satiation following adjustedfeeding rates. In the experimental treatments, the control group feeding rates wasreduced to75%,50%and25%, and the four diets were named as100%,75%,50%and25%, respectively. The results showed that weight gain rate (WGR) was linearly related to feed intake (FI)(P<0.05). Significant positive correlations were observedbetween the FI and some the selected plasma parameters, total cholesterol (TC), freecholesterol (FC), high-density lipoprotein cholesterol (HDL-C) and low-densitylipoprotein cholesterol (LDL-C)(P<0.05). The FI was significant negativecorrelations with the TC, FC and cholesterol esters in fish liver (P<0.05). Fish fed to25%of satiation showed significant low activity of acyl-coenzyme A: cholesterolacyltransferase (ACAT) compared with fish fed to the control group(P<0.05), and asignificant positive correlations was observed between FI and activity of ACAT(P<0.05). The expression levels of3-hydroxy-3-methylglutaryl-Coenzyme Areductase (HMG-CoAr) transcript were significantly up-regulated in liver of turbotfed fed to50%or25%of satiation compared with that in fish fed to the control group.These results suggested thatthe hypocholesterolemic effect was observed in plasma offish fed plant protein diets, which were related to, in part, significantly decreasedFI.Moreover, the increased hepatic TC level in fish fed to25%or50%of satiation may beattributed to the increased ability of cholesterol synthesis.
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