瓦氏黄颡鱼（Pelteobagrus vachelli）脂肪代谢相关基因cDNA的克隆及表达分析
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摘要
随着水生动物营养学研究的不断深入和完善,近年来,鱼类的脂类营养已逐渐成为营养学的一个研究热点。迄今不同学者分别从脂类营养与动物的生长性能、生理学参数,以及生化酶和代谢之间的关系等方面做了大量的工作,但从分子水平上,尤其是有关外源营养素与鱼类代谢相关酶基因之间关系的研究仍颇为缺乏。本论文以瓦氏黄颡鱼(darkbarbel catfish, Pelteobagrus vachelli)为研究对象,主要从分子角度探讨了脂肪种类、水平等对瓦氏黄颡鱼幼鱼生长和脂肪代谢的影响,研究结果可以为完善瓦氏黄颡鱼人工饲料配方和全价配合饲料的开发提供参考。
本论文利用分子生物学技术和生物信息学方法,首先克隆了瓦氏黄颡鱼的主要脂肪代谢相关基因肝脂酶(hepatic lipase, HL),脂肪酸结合蛋白(fatty acid binding protein, FABP),脂肪酸合成酶(fatty acid synthase, FAS)和肉碱酰基转运酶(carnitine acetyltransferase, CAT)的cDNA片段,分析了这些基因的cDNA序列和分子特征,以及在不同组织中的表达模式。在此基础上,进行了室内饲养试验,研究了饲料脂肪种类和水平等对瓦氏黄颡鱼生长参数、脂肪代谢相关基因mRNA表达的影响,探讨了它们在机体脂肪的分解、转运,以及体内合成等过程发挥的作用。
论文的主要结果和结论如下：
1瓦氏黄颡鱼脂肪代谢相关基因的研究
本研究根据GenBnak中提供的生物学信息设计简并引物,采用RT-PCR和克隆测序及生物信息学方法、从瓦氏黄颡鱼肝脏中克隆得到肝脂酶,肝型脂肪酸结合蛋白,心型脂肪酸结合蛋白,脂肪酸合成酶和肉碱酰基转运酶基因的cDNA片段。
肝脂酶是血液循环中内源性甘油三酯代谢的关键酶之一。本试验克隆了瓦氏黄颡鱼肝脂酶的部分cDNA序列。该片段长为1065bp序列,软件分析获得了343个氨基酸,其序列与斑马鱼(Danio rerio)、日本鳗鲡(Anguilla japonica)、鳜鱼(Siniperca chuatsi)、斜带石斑鱼(Epinephelus coioides)、真鲷(Pagrus major)的肝脂酶具有很高的同源性,分别为73%、68%、67%、66%和65%。经分析,所得瓦氏黄颡鱼肝脂酶氨基酸残基中含有1个糖基化位点,催化中心三联体位点1个和多肽“盖”等主要的结构功能位点,因而该基因属于脂酶家族。定量PCR分析表明肝脂酶只在肝脏组织中表达,表明瓦氏黄颡鱼肝脂酶的表达具有组织特异性,肝脏是分泌肝脂酶及其行使功能的主要场所。
脂肪酸结合蛋白是一类分子量较小,而对脂肪酸有高亲和力的可溶性蛋白质。本试验采用简并引物扩增获得了瓦氏黄颡鱼脂肪酸结合蛋白部分cDNA序列。肝型脂肪酸结合蛋白cDNA片段长为335bp,软件分析获得111个氨基酸残基,与鲤鱼(Cyprinus carpio)的肝型脂肪酸结合蛋白的氨基酸序列同源性最高,可达到90%。瓦氏黄颡鱼心型脂肪酸结合蛋白cDNA序列片段长为425bp序列,包括26bp 3’端非编码区。软件分析后获得114个氨基酸残基,与斜齿鳊(Rutilus rutilus)的心型脂肪酸结合蛋白氨基酸序列同源性高达85%。结果表明克隆所得基因序列均属于脂肪酸结合蛋白家族。定量PCR分析表明,瓦氏黄颡鱼肝型脂肪酸结合蛋白在肝脏、腹腔脂肪、肌肉和心脏组织有较高的表达量。心型脂肪酸结合蛋白主要在肝脏、脂肪组织和心脏中表达,表明肝脏和腹腔脂肪组织均是肝型和心型脂肪酸结合蛋白表达的主要场所。
脂肪酸合成酶能催化机体内脂肪酸的从头合成,生成内源性脂肪酸。本试验采用RT-PCR技术成功分离了瓦氏黄颡鱼脂肪酸合成酶cDNA序列片段,序列长为674bp,软件分析后获得216个氨基酸残基,与斑马鱼(D. rerio)脂肪酸合成酶氨基酸序列同源性最高(87%),其次是红原鸡(Gallus gallus,79%)、南方短尾负鼠(Monodelphis domestica,70%)、褐家鼠(Rattus norvegicus,68%)等。定量PCR分析表明,瓦氏黄颡鱼脂肪酸合成酶mRNA在肝脏、肠道、腹腔脂肪、心脏组织的表达水平显著高于肌肉等组织(P<0.05),脂肪酸合成酶mRNA的表达量上升,有利于促进脂肪酸的生物合成,以满足不同组织细胞分化增殖对脂质营养和能量的需要。
肉碱酰基转运酶属于转运酶家族,能转运从乙酰辅酶A到L-肉碱等酰基基团。本试验获得了瓦氏黄颡鱼肉碱酰基转运酶部分cDNA序列,片段长为494bp,序列软件分析后获得156个氨基酸残基,与斑马鱼(D. rerio)和红原鸡(G. gallus)的肉碱酰基转运酶氨基酸序列同源性分别为81%和70%,分析表明所得序列属于酰基转运酶家族。PCR检测发现瓦氏黄颡鱼肉碱酰基转运酶主要在肝脏、腹腔脂肪组织和肌肉等组织中表达,提示这些组织均为能量代谢的重要场所。
瓦氏黄颡鱼肝脂酶、心型脂肪酸结合蛋白、脂肪酸合成酶和肉碱酰基转运酶cDNA片段的成功克隆,为研究瓦氏黄颡鱼脂肪代谢提供了基础资料。根据肝脂酶、心型脂肪酸结合蛋白、脂肪酸合成酶和肉碱酰基转运酶的组织表达特点,后续的生长试验中,选择了鱼的肝脏和腹腔脂肪组织为目标靶器官,研究了饲料脂肪种类和水平对瓦氏黄颡鱼的生长和脂肪代谢的影响。
2饲料脂肪种类对瓦氏黄颡鱼的生长和脂肪代谢相关基因表达的影响
研究了饲料中不同脂肪种类对瓦氏黄颡鱼幼鱼生长、主要脂肪代谢相关基因表达的影响。试验以鱼粉和豆粕为蛋白源,分别添加8%的全鱼油、豆油、猪油、混合油1(豆油和鱼油混合=3：1)、混合油2(豆油和鱼油混合=1：3),配制了等氮等能的5种饲料。每组设三重复,投喂初始平均体重约为1.0 g的幼鱼80天。结果表明,鱼油、豆油、混合油1和混合油2饲料4个处理组均表现出较好的生长趋势,终末体重和特定生长率分别达到7.67g、7.47g、8.21g、8.31g和2.52%、2.50%、2.65%、2.63%,均显著高于猪油组6.13g和2.32%(P<0.05)。饲料脂肪源对瓦氏黄颡鱼的腹腔脂肪率和肝体指数无显著影响,腹腔脂肪率分别为3.52%,4.49%,3.70%,3.92%,4.42%；肝体指数分别为2.66%,2.88%,3.06%,3.08%,2.54%。根据所得的结果,可以认为短期内以豆油作为脂肪源不会显著影响瓦氏黄颡鱼的生长。
定量PCR分析主要脂肪代谢相关基因mRNA表达显示,在肝脏组织中,混合油2组的脂蛋白脂酶,肝脂酶,肝型脂肪酸结合蛋白和肉碱酰基转运酶mRNA表达量均显著高于其它组(P<0.05)。而猪油组能促进脂肪酸合成酶mRNA表达上调,且显著高于其它组(P<0.05)。同时,在腹腔脂肪组织中,脂蛋白脂酶和肝型脂肪酸结合蛋白mRNA表达具有组织特异性,各组间均无显著差异。混合油1组脂肪酸合成酶表达显著高于鱼油组(P<0.05),而其它组间无显著差异。肉碱酰基转运酶mRNA的表达在混合油1和混合油2组中表现为上调,且显著高于猪油组(P<0.05)。混合油2组饲料中多不饱和脂肪酸百分含量(46%)最高,能促进肝脏脂蛋白脂酶、肝脂酶、肝型脂肪酸结合蛋白和肉碱酰基转运酶mRNA的表达显著上调(P<0.05),提高饲料多不饱和脂肪酸的利用率,从而促进了瓦氏黄颡鱼幼鱼的生长。上述结果提示,不同的饲料脂肪种类主要通过调节瓦氏黄颡鱼肝脏的脂蛋白脂酶和肝脂酶分解脂蛋白的能力,使得肝型脂肪酸结合蛋白转运脂肪酸、脂肪酸的合成活动发生相应的变化,最终使机体生长和代谢所需的脂肪酸得到满足。
3脂肪水平对瓦氏黄颡鱼幼鱼的生长和脂肪代谢相关基因表达的影响
探讨了饲料中不同脂肪水平对瓦氏黄颡鱼幼鱼生长和主要脂肪代谢相关基因表达的影响。以鱼粉和脱脂豆粕为饲料蛋白源,鱼油和大豆油等比例混合为脂肪源,分别配制含有5%,11%和20%三个脂肪水平的等氮饲料。试验组每组各设三个重复,投喂初始平均体重1.0g左右的幼鱼,70天后5%、11%和20%处理组平均体重分别为8.45g、7.92g、6.94g。经统计分析,5%和11%处理组的终末体重、特定生长率显著高于20%处理组。从生长试验来看,饲料脂肪水平对瓦氏黄颡鱼的生长有显著的影响,随着饲料脂肪水平增加,腹腔脂肪率显著升高(P<0.05),分别是2.56%,4.27%和6.06%；但随饲料脂肪水平提高,鱼的肝体指数没有显著变化,分别为2.59%；2.47%和2.58%。
对主要脂肪代谢相关基因mRNA的定量PCR分析显示,随着饲料脂肪水平的升高,肝脏脂蛋白脂酶,肝脂酶和肝型脂肪酸结合蛋白的mRNA表达水平显著提高(P<0.05)；而脂肪酸合成酶和肉碱酰基转运酶的mRNA表达水平却显著下降(P<0.05),肉碱酰基转运酶mRNA表达在11%处理组和20%处理组之间无显著差异(P>0.05)。同时,在腹腔脂肪组织中,11%处理组脂蛋白脂酶基因mRNA的表达量显著低于20%处理组(P<0.05),略低于5%处理组。肝型脂肪酸结合蛋白mRNA表达在11%处理组中最低,并显著低于5%处理组(P<0.05)。肉碱酰基准转运酶mRNA表达在20%处理组最高,显著高于11%处理组和5%处理组(P<0.05)。此外,5%处理组脂肪酸合成酶表达水平显著高于11%处理组和20%处理组(P<0.05)。上述研究结果表明,饲料脂肪水平会影响肝脏和腹腔脂肪组织脂肪代谢相关基因mRNA的表达；过高的饲料脂肪水平,能通过增强肝脏和腹腔脂肪组织中的脂肪分解,促进脂肪酸转运,抑制脂肪酸生成等环节来调节脂肪的代谢,以减少脂肪在机体内组织器官的沉积。
4瘦素对瓦氏黄颡鱼脂肪代谢相关基因表达的影响
本试验分析了腹腔注射小鼠瘦素重组蛋白对瓦氏黄颡鱼肝脏脂肪代谢的影响。试验设置4个处理组,分别注射20μg/100μl,5μg/100μl,磷酸缓冲液(PBS,100μl)和未注射空白组(对照组),每个组含三个重复,每个重复放养9尾试验鱼(体重30±0.5g)。对试验鱼隔天注射一次,连续饲养15天,按常规饲养投喂和管理。试验结果表明,与未注射组的鱼相比,注射不同含量的瘦素均能显著地提高鱼肝脏中肉碱酰基转运酶mRNA的表达量(P<0.05),但不同注射剂量的各瘦素处理组间无显著性差异。低瘦素浓度组能显著地促进肝型脂肪酸结合蛋白mRNA的表达(P<0.05),但注射瘦素会显著抑制脂肪酸合成酶mRNA的表达,且随注射浓度的增加,其mRNA的表达随之明显下降(P<0.05)。此外,瘦素注射能增加脂蛋白脂酶和肝脂酶mRNA的表达,但均与对照组无显著差异(P>0.05)。结果提示外源瘦素可能是通过促进肝型脂肪酸结合蛋白mRNA的表达,增加了体内脂肪酸的转运能力；同时,由于脂肪酸合成酶mRNA表达水平的降低,抑制了体内脂肪酸的合成,从而促进机体脂肪代谢活动,维持鱼体的脂肪贮存量和能量的平衡。
Due to the importance of dietary lipid utilization in fish rearing, increasing attention has been paid to different aspects of fish lipid nutrition. However, different explanations have been proposed but the exact mechanisms behind these observations have not been completely clarified. So this study was conducted to investigate that the lipid level and source effect the growth and lipid metabolism related gene expression of freshwater omnivorous darkbarbel catfish, Pelteobagrus vachelli.
1 Cloning and tissues expression of the lipid metabolism related gene in darkbarbel catfish, Pelteobagrus vachelli
To investigated the lipid nutritional regulation of lipid molecular metabolism of fish, hepatic lipase (HL), fatty acid binding protein (FABP), fatty acid synthase (FAS) and carnitine acetyltransferase (CAT) gene of darkbarbel catfish P. vachelli were isolated from the liver. (1):The partial cDNA of HL (1065bp) was cloned, and was capable of encoding 343 amino acids residues, and showed significant identity to the HL gene of Danio rerio (73%). The partial deduced amino acids residues carried conserved features of this family, and found the"lid"sites. HL gene mRNA expression was detected in liver, but not in gill, muscle, intestine, spleen, heart and brain. (2)The length of cloned liver and heart FABP (L-and H-FABP) cDNA sequences were 335 and 425bp, and encoded 111 and 114 amino acids residues, respectively. Homologous alignment analysis showed L-and H-FABP gene had 82% and 85% identity to that of Cyprinus carpio and Rutilus rutilus, respectively. FABP gene mRNA was mainly distributed in liver, visceral adipose tissue, muscle and heart for liver FABP, and in liver, visceral adipose tissue and heart for heart FABP. (3):The partial cDNA of FAS gene consisted of 674bp, finding 216 amino acids residues. Sequence comparison showed that the partial FAS amino acids residues had the highest identity with D. rerio (90%). FAS transcripts were detected in all sampled tissues, and significantly distributed in liver, visceral adipose tissue, muscle, intestine, heart. (4):cDNA encoding CAT mRNA was obtained from the liver, partial cDNA with 494bp encoded 156 amino acids residues. Comparison the sequence of CAT amino acids residues showed the highest identity with D. rerio (81%).
2 Effect of lipid sources on the growth and expression of related metabolism gene of lipid of darkbarbel catfish, Pelteobagrus vachelli
The effects of dietary lipid sources on fatty acid absorption and expression regulation of genes related to lipid metabolism in darkbarkbel catfish, Pelteobagrus vachelli, were evaluated. Fingerlings of P. vachelli (1.0±0.2 g) were fed five experimental diets with fish oil (FO), soybean oil (SO), pig oil (PO),75% fish oil and 25% soybean oil (Mix group 1), 25% fish oil and 75% soybean oil (Mix group 2) as lipid sources respectively for 80 days. Fish weight gains of SO, M1 and M2 groups were similar to those of FO group, but total replacement of fish oil by PO significantly reduced fish weight gain (P<0.05). Though there were increasing trends in hepatosomatic index (HSI) and intraperitoneal fat ratio (IPF) of fish feeding SO, M1, M2 and PO deits, no significant differences were found (P<0.05). Fish of PO group had higher SFA fraction and lower PUFA fraction in feces relative to those feeding FO diets. Fish of SO, M1 and M2 groups had significantly higher PUFA fraction and lower SFA fraction (p<0.05) in feces, and the PUFA fraction in feces increased with the replacement content of fish oil by soybean oil. Furthermore, replacement of fish oil with SO, PO and 75% SO could significantly increased the fatty acid binding protein (FABP) and fatty acid synthase (FAS) gene expressions in liver and adipose tissue (P<0.05), and significantly reduced the carnitine acyltransferase gene expression in liver (P<0.05). No significant changes were found in lipoprotein lipase (LPL) gene expression in liver and adipose tissue, and hepatic lipase (HL) gene expression in liver (P<0.05). All these findings could suggest that soybean oil could increase lipid biosynthesis and slow down the lipid catabolism of P. vachelli. Therefore, partly replacement of fish oil by soybean oil did not reduce the weight gain of darkbarkbel catfish.
3 Effect of lipid level on the growth and expression of related lipid metabolism gene of lipid of darkbarbel catfish, Pelteobagrus vachelli
An experiment was conducted to determine the effect of dietary lipid levels on growth performance and expression regulation of genes of darkbarbel catfish, P. vachelli juveniles cultured in plastic boxes. Three isonitrogenous diets (42% dietary protein) with increasing dietary lipid concentration (5%,11%,20% of dry material, DM) were fed to satiation to triplicate groups of 30 fish (mean weight:1.0±0.1 g) for 70 days. Fish weight gains of 5% lipid and 11% lipid groups were similar, but weight gains of 20%lipid group significantly reduced (P<0.05). Furthermore, intraperitoneal fat ratio (IPF) significantly increased with lipid level (P<0.05), but no significant differences were found for hepatosomatic index (HSI) between different lipid level diets (P<0.05). Furthermore, in liver, the mRNA expression of LPL, HL and L-FABP significantly increased, FAS and CAT significantly reduced with the augment of diet lipid level (P<0.05). In visceral adipose tissue, gene expression characterization of lipid metabolism was multiplex. Both LPL and L-FABP mRNA expression were the highest in 5% lipid group. The 20% lipid group significantly increased the expression of CAT mRNA, but there was no significant difference between 5% lipid and 11% lipid groups. CAT mRNA was the highest in 20% lipid group. However, the FAS transcripts was decreased with increasing of dietary lipid level (P<0.05), but there was no significant difference between 11% and 20% lipid groups. These results suggested that increasing dietary lipid level could induce the metabolism of lipid in liver and visceral adipose tissue, and reduce the lipid biosynthesis.
4 Effect of murine leptin injections on expression of the related metabolism gene of lipid of darkbarbel catfish, Pelteobagrus vachelli
An experiment was conducted to determine the effect of leptin on regulation of genes related to lipid metabolism in darkbarkbel catfish, Pelteobagrus vachelli, were evaluated. The darkbarbel catfish, P. vachelli, (30±0.5g) were injected daily with higher murine leptin (20μg/100μl), lower murine leptin (5μg/100μl), phosphate-buffered saline (PBS), or simply handled without injection for 15 days. At the end of the experiment, fish were assayed for expression regulation of genes related to lipid metabolism in liver. Injecting leptin could remarkably increase the mRNA expression of CAT and L-FABP, but no significant difference was found between high and low concentration. Though there were increasing trends with leptin injection for HL and LPL mRNA expression, but no significant differences were found in all groups (P<0.05). However, the FAS mRNA expression in leptin treated groups was significantly decreased. These results suggest that fish respond to murine leptin injections by increasing fat transport, and depressing the lipid biosynthesis.
本论文利用分子生物学技术和生物信息学方法,首先克隆了瓦氏黄颡鱼的主要脂肪代谢相关基因肝脂酶(hepatic lipase, HL),脂肪酸结合蛋白(fatty acid binding protein, FABP),脂肪酸合成酶(fatty acid synthase, FAS)和肉碱酰基转运酶(carnitine acetyltransferase, CAT)的cDNA片段,分析了这些基因的cDNA序列和分子特征,以及在不同组织中的表达模式。在此基础上,进行了室内饲养试验,研究了饲料脂肪种类和水平等对瓦氏黄颡鱼生长参数、脂肪代谢相关基因mRNA表达的影响,探讨了它们在机体脂肪的分解、转运,以及体内合成等过程发挥的作用。
论文的主要结果和结论如下：
1瓦氏黄颡鱼脂肪代谢相关基因的研究
本研究根据GenBnak中提供的生物学信息设计简并引物,采用RT-PCR和克隆测序及生物信息学方法、从瓦氏黄颡鱼肝脏中克隆得到肝脂酶,肝型脂肪酸结合蛋白,心型脂肪酸结合蛋白,脂肪酸合成酶和肉碱酰基转运酶基因的cDNA片段。
肝脂酶是血液循环中内源性甘油三酯代谢的关键酶之一。本试验克隆了瓦氏黄颡鱼肝脂酶的部分cDNA序列。该片段长为1065bp序列,软件分析获得了343个氨基酸,其序列与斑马鱼(Danio rerio)、日本鳗鲡(Anguilla japonica)、鳜鱼(Siniperca chuatsi)、斜带石斑鱼(Epinephelus coioides)、真鲷(Pagrus major)的肝脂酶具有很高的同源性,分别为73%、68%、67%、66%和65%。经分析,所得瓦氏黄颡鱼肝脂酶氨基酸残基中含有1个糖基化位点,催化中心三联体位点1个和多肽“盖”等主要的结构功能位点,因而该基因属于脂酶家族。定量PCR分析表明肝脂酶只在肝脏组织中表达,表明瓦氏黄颡鱼肝脂酶的表达具有组织特异性,肝脏是分泌肝脂酶及其行使功能的主要场所。
脂肪酸结合蛋白是一类分子量较小,而对脂肪酸有高亲和力的可溶性蛋白质。本试验采用简并引物扩增获得了瓦氏黄颡鱼脂肪酸结合蛋白部分cDNA序列。肝型脂肪酸结合蛋白cDNA片段长为335bp,软件分析获得111个氨基酸残基,与鲤鱼(Cyprinus carpio)的肝型脂肪酸结合蛋白的氨基酸序列同源性最高,可达到90%。瓦氏黄颡鱼心型脂肪酸结合蛋白cDNA序列片段长为425bp序列,包括26bp 3’端非编码区。软件分析后获得114个氨基酸残基,与斜齿鳊(Rutilus rutilus)的心型脂肪酸结合蛋白氨基酸序列同源性高达85%。结果表明克隆所得基因序列均属于脂肪酸结合蛋白家族。定量PCR分析表明,瓦氏黄颡鱼肝型脂肪酸结合蛋白在肝脏、腹腔脂肪、肌肉和心脏组织有较高的表达量。心型脂肪酸结合蛋白主要在肝脏、脂肪组织和心脏中表达,表明肝脏和腹腔脂肪组织均是肝型和心型脂肪酸结合蛋白表达的主要场所。
脂肪酸合成酶能催化机体内脂肪酸的从头合成,生成内源性脂肪酸。本试验采用RT-PCR技术成功分离了瓦氏黄颡鱼脂肪酸合成酶cDNA序列片段,序列长为674bp,软件分析后获得216个氨基酸残基,与斑马鱼(D. rerio)脂肪酸合成酶氨基酸序列同源性最高(87%),其次是红原鸡(Gallus gallus,79%)、南方短尾负鼠(Monodelphis domestica,70%)、褐家鼠(Rattus norvegicus,68%)等。定量PCR分析表明,瓦氏黄颡鱼脂肪酸合成酶mRNA在肝脏、肠道、腹腔脂肪、心脏组织的表达水平显著高于肌肉等组织(P<0.05),脂肪酸合成酶mRNA的表达量上升,有利于促进脂肪酸的生物合成,以满足不同组织细胞分化增殖对脂质营养和能量的需要。
肉碱酰基转运酶属于转运酶家族,能转运从乙酰辅酶A到L-肉碱等酰基基团。本试验获得了瓦氏黄颡鱼肉碱酰基转运酶部分cDNA序列,片段长为494bp,序列软件分析后获得156个氨基酸残基,与斑马鱼(D. rerio)和红原鸡(G. gallus)的肉碱酰基转运酶氨基酸序列同源性分别为81%和70%,分析表明所得序列属于酰基转运酶家族。PCR检测发现瓦氏黄颡鱼肉碱酰基转运酶主要在肝脏、腹腔脂肪组织和肌肉等组织中表达,提示这些组织均为能量代谢的重要场所。
瓦氏黄颡鱼肝脂酶、心型脂肪酸结合蛋白、脂肪酸合成酶和肉碱酰基转运酶cDNA片段的成功克隆,为研究瓦氏黄颡鱼脂肪代谢提供了基础资料。根据肝脂酶、心型脂肪酸结合蛋白、脂肪酸合成酶和肉碱酰基转运酶的组织表达特点,后续的生长试验中,选择了鱼的肝脏和腹腔脂肪组织为目标靶器官,研究了饲料脂肪种类和水平对瓦氏黄颡鱼的生长和脂肪代谢的影响。
2饲料脂肪种类对瓦氏黄颡鱼的生长和脂肪代谢相关基因表达的影响
研究了饲料中不同脂肪种类对瓦氏黄颡鱼幼鱼生长、主要脂肪代谢相关基因表达的影响。试验以鱼粉和豆粕为蛋白源,分别添加8%的全鱼油、豆油、猪油、混合油1(豆油和鱼油混合=3：1)、混合油2(豆油和鱼油混合=1：3),配制了等氮等能的5种饲料。每组设三重复,投喂初始平均体重约为1.0 g的幼鱼80天。结果表明,鱼油、豆油、混合油1和混合油2饲料4个处理组均表现出较好的生长趋势,终末体重和特定生长率分别达到7.67g、7.47g、8.21g、8.31g和2.52%、2.50%、2.65%、2.63%,均显著高于猪油组6.13g和2.32%(P<0.05)。饲料脂肪源对瓦氏黄颡鱼的腹腔脂肪率和肝体指数无显著影响,腹腔脂肪率分别为3.52%,4.49%,3.70%,3.92%,4.42%；肝体指数分别为2.66%,2.88%,3.06%,3.08%,2.54%。根据所得的结果,可以认为短期内以豆油作为脂肪源不会显著影响瓦氏黄颡鱼的生长。
定量PCR分析主要脂肪代谢相关基因mRNA表达显示,在肝脏组织中,混合油2组的脂蛋白脂酶,肝脂酶,肝型脂肪酸结合蛋白和肉碱酰基转运酶mRNA表达量均显著高于其它组(P<0.05)。而猪油组能促进脂肪酸合成酶mRNA表达上调,且显著高于其它组(P<0.05)。同时,在腹腔脂肪组织中,脂蛋白脂酶和肝型脂肪酸结合蛋白mRNA表达具有组织特异性,各组间均无显著差异。混合油1组脂肪酸合成酶表达显著高于鱼油组(P<0.05),而其它组间无显著差异。肉碱酰基转运酶mRNA的表达在混合油1和混合油2组中表现为上调,且显著高于猪油组(P<0.05)。混合油2组饲料中多不饱和脂肪酸百分含量(46%)最高,能促进肝脏脂蛋白脂酶、肝脂酶、肝型脂肪酸结合蛋白和肉碱酰基转运酶mRNA的表达显著上调(P<0.05),提高饲料多不饱和脂肪酸的利用率,从而促进了瓦氏黄颡鱼幼鱼的生长。上述结果提示,不同的饲料脂肪种类主要通过调节瓦氏黄颡鱼肝脏的脂蛋白脂酶和肝脂酶分解脂蛋白的能力,使得肝型脂肪酸结合蛋白转运脂肪酸、脂肪酸的合成活动发生相应的变化,最终使机体生长和代谢所需的脂肪酸得到满足。
3脂肪水平对瓦氏黄颡鱼幼鱼的生长和脂肪代谢相关基因表达的影响
探讨了饲料中不同脂肪水平对瓦氏黄颡鱼幼鱼生长和主要脂肪代谢相关基因表达的影响。以鱼粉和脱脂豆粕为饲料蛋白源,鱼油和大豆油等比例混合为脂肪源,分别配制含有5%,11%和20%三个脂肪水平的等氮饲料。试验组每组各设三个重复,投喂初始平均体重1.0g左右的幼鱼,70天后5%、11%和20%处理组平均体重分别为8.45g、7.92g、6.94g。经统计分析,5%和11%处理组的终末体重、特定生长率显著高于20%处理组。从生长试验来看,饲料脂肪水平对瓦氏黄颡鱼的生长有显著的影响,随着饲料脂肪水平增加,腹腔脂肪率显著升高(P<0.05),分别是2.56%,4.27%和6.06%；但随饲料脂肪水平提高,鱼的肝体指数没有显著变化,分别为2.59%；2.47%和2.58%。
对主要脂肪代谢相关基因mRNA的定量PCR分析显示,随着饲料脂肪水平的升高,肝脏脂蛋白脂酶,肝脂酶和肝型脂肪酸结合蛋白的mRNA表达水平显著提高(P<0.05)；而脂肪酸合成酶和肉碱酰基转运酶的mRNA表达水平却显著下降(P<0.05),肉碱酰基转运酶mRNA表达在11%处理组和20%处理组之间无显著差异(P>0.05)。同时,在腹腔脂肪组织中,11%处理组脂蛋白脂酶基因mRNA的表达量显著低于20%处理组(P<0.05),略低于5%处理组。肝型脂肪酸结合蛋白mRNA表达在11%处理组中最低,并显著低于5%处理组(P<0.05)。肉碱酰基准转运酶mRNA表达在20%处理组最高,显著高于11%处理组和5%处理组(P<0.05)。此外,5%处理组脂肪酸合成酶表达水平显著高于11%处理组和20%处理组(P<0.05)。上述研究结果表明,饲料脂肪水平会影响肝脏和腹腔脂肪组织脂肪代谢相关基因mRNA的表达；过高的饲料脂肪水平,能通过增强肝脏和腹腔脂肪组织中的脂肪分解,促进脂肪酸转运,抑制脂肪酸生成等环节来调节脂肪的代谢,以减少脂肪在机体内组织器官的沉积。
4瘦素对瓦氏黄颡鱼脂肪代谢相关基因表达的影响
本试验分析了腹腔注射小鼠瘦素重组蛋白对瓦氏黄颡鱼肝脏脂肪代谢的影响。试验设置4个处理组,分别注射20μg/100μl,5μg/100μl,磷酸缓冲液(PBS,100μl)和未注射空白组(对照组),每个组含三个重复,每个重复放养9尾试验鱼(体重30±0.5g)。对试验鱼隔天注射一次,连续饲养15天,按常规饲养投喂和管理。试验结果表明,与未注射组的鱼相比,注射不同含量的瘦素均能显著地提高鱼肝脏中肉碱酰基转运酶mRNA的表达量(P<0.05),但不同注射剂量的各瘦素处理组间无显著性差异。低瘦素浓度组能显著地促进肝型脂肪酸结合蛋白mRNA的表达(P<0.05),但注射瘦素会显著抑制脂肪酸合成酶mRNA的表达,且随注射浓度的增加,其mRNA的表达随之明显下降(P<0.05)。此外,瘦素注射能增加脂蛋白脂酶和肝脂酶mRNA的表达,但均与对照组无显著差异(P>0.05)。结果提示外源瘦素可能是通过促进肝型脂肪酸结合蛋白mRNA的表达,增加了体内脂肪酸的转运能力；同时,由于脂肪酸合成酶mRNA表达水平的降低,抑制了体内脂肪酸的合成,从而促进机体脂肪代谢活动,维持鱼体的脂肪贮存量和能量的平衡。
Due to the importance of dietary lipid utilization in fish rearing, increasing attention has been paid to different aspects of fish lipid nutrition. However, different explanations have been proposed but the exact mechanisms behind these observations have not been completely clarified. So this study was conducted to investigate that the lipid level and source effect the growth and lipid metabolism related gene expression of freshwater omnivorous darkbarbel catfish, Pelteobagrus vachelli.
1 Cloning and tissues expression of the lipid metabolism related gene in darkbarbel catfish, Pelteobagrus vachelli
To investigated the lipid nutritional regulation of lipid molecular metabolism of fish, hepatic lipase (HL), fatty acid binding protein (FABP), fatty acid synthase (FAS) and carnitine acetyltransferase (CAT) gene of darkbarbel catfish P. vachelli were isolated from the liver. (1):The partial cDNA of HL (1065bp) was cloned, and was capable of encoding 343 amino acids residues, and showed significant identity to the HL gene of Danio rerio (73%). The partial deduced amino acids residues carried conserved features of this family, and found the"lid"sites. HL gene mRNA expression was detected in liver, but not in gill, muscle, intestine, spleen, heart and brain. (2)The length of cloned liver and heart FABP (L-and H-FABP) cDNA sequences were 335 and 425bp, and encoded 111 and 114 amino acids residues, respectively. Homologous alignment analysis showed L-and H-FABP gene had 82% and 85% identity to that of Cyprinus carpio and Rutilus rutilus, respectively. FABP gene mRNA was mainly distributed in liver, visceral adipose tissue, muscle and heart for liver FABP, and in liver, visceral adipose tissue and heart for heart FABP. (3):The partial cDNA of FAS gene consisted of 674bp, finding 216 amino acids residues. Sequence comparison showed that the partial FAS amino acids residues had the highest identity with D. rerio (90%). FAS transcripts were detected in all sampled tissues, and significantly distributed in liver, visceral adipose tissue, muscle, intestine, heart. (4):cDNA encoding CAT mRNA was obtained from the liver, partial cDNA with 494bp encoded 156 amino acids residues. Comparison the sequence of CAT amino acids residues showed the highest identity with D. rerio (81%).
2 Effect of lipid sources on the growth and expression of related metabolism gene of lipid of darkbarbel catfish, Pelteobagrus vachelli
The effects of dietary lipid sources on fatty acid absorption and expression regulation of genes related to lipid metabolism in darkbarkbel catfish, Pelteobagrus vachelli, were evaluated. Fingerlings of P. vachelli (1.0±0.2 g) were fed five experimental diets with fish oil (FO), soybean oil (SO), pig oil (PO),75% fish oil and 25% soybean oil (Mix group 1), 25% fish oil and 75% soybean oil (Mix group 2) as lipid sources respectively for 80 days. Fish weight gains of SO, M1 and M2 groups were similar to those of FO group, but total replacement of fish oil by PO significantly reduced fish weight gain (P<0.05). Though there were increasing trends in hepatosomatic index (HSI) and intraperitoneal fat ratio (IPF) of fish feeding SO, M1, M2 and PO deits, no significant differences were found (P<0.05). Fish of PO group had higher SFA fraction and lower PUFA fraction in feces relative to those feeding FO diets. Fish of SO, M1 and M2 groups had significantly higher PUFA fraction and lower SFA fraction (p<0.05) in feces, and the PUFA fraction in feces increased with the replacement content of fish oil by soybean oil. Furthermore, replacement of fish oil with SO, PO and 75% SO could significantly increased the fatty acid binding protein (FABP) and fatty acid synthase (FAS) gene expressions in liver and adipose tissue (P<0.05), and significantly reduced the carnitine acyltransferase gene expression in liver (P<0.05). No significant changes were found in lipoprotein lipase (LPL) gene expression in liver and adipose tissue, and hepatic lipase (HL) gene expression in liver (P<0.05). All these findings could suggest that soybean oil could increase lipid biosynthesis and slow down the lipid catabolism of P. vachelli. Therefore, partly replacement of fish oil by soybean oil did not reduce the weight gain of darkbarkbel catfish.
3 Effect of lipid level on the growth and expression of related lipid metabolism gene of lipid of darkbarbel catfish, Pelteobagrus vachelli
An experiment was conducted to determine the effect of dietary lipid levels on growth performance and expression regulation of genes of darkbarbel catfish, P. vachelli juveniles cultured in plastic boxes. Three isonitrogenous diets (42% dietary protein) with increasing dietary lipid concentration (5%,11%,20% of dry material, DM) were fed to satiation to triplicate groups of 30 fish (mean weight:1.0±0.1 g) for 70 days. Fish weight gains of 5% lipid and 11% lipid groups were similar, but weight gains of 20%lipid group significantly reduced (P<0.05). Furthermore, intraperitoneal fat ratio (IPF) significantly increased with lipid level (P<0.05), but no significant differences were found for hepatosomatic index (HSI) between different lipid level diets (P<0.05). Furthermore, in liver, the mRNA expression of LPL, HL and L-FABP significantly increased, FAS and CAT significantly reduced with the augment of diet lipid level (P<0.05). In visceral adipose tissue, gene expression characterization of lipid metabolism was multiplex. Both LPL and L-FABP mRNA expression were the highest in 5% lipid group. The 20% lipid group significantly increased the expression of CAT mRNA, but there was no significant difference between 5% lipid and 11% lipid groups. CAT mRNA was the highest in 20% lipid group. However, the FAS transcripts was decreased with increasing of dietary lipid level (P<0.05), but there was no significant difference between 11% and 20% lipid groups. These results suggested that increasing dietary lipid level could induce the metabolism of lipid in liver and visceral adipose tissue, and reduce the lipid biosynthesis.
4 Effect of murine leptin injections on expression of the related metabolism gene of lipid of darkbarbel catfish, Pelteobagrus vachelli
An experiment was conducted to determine the effect of leptin on regulation of genes related to lipid metabolism in darkbarkbel catfish, Pelteobagrus vachelli, were evaluated. The darkbarbel catfish, P. vachelli, (30±0.5g) were injected daily with higher murine leptin (20μg/100μl), lower murine leptin (5μg/100μl), phosphate-buffered saline (PBS), or simply handled without injection for 15 days. At the end of the experiment, fish were assayed for expression regulation of genes related to lipid metabolism in liver. Injecting leptin could remarkably increase the mRNA expression of CAT and L-FABP, but no significant difference was found between high and low concentration. Though there were increasing trends with leptin injection for HL and LPL mRNA expression, but no significant differences were found in all groups (P<0.05). However, the FAS mRNA expression in leptin treated groups was significantly decreased. These results suggest that fish respond to murine leptin injections by increasing fat transport, and depressing the lipid biosynthesis.
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