饲料中n-3/n-6多不饱和脂肪酸水平对黄河鲤幼鱼生长性能及生长相关基因mRNA表达的影响
|
摘要
为了分析饲料n-3/n-6多不饱和脂肪酸(PUFAs)水平对黄河鲤幼鱼生长性能和生长相关基因的影响,实验以鱼油和混合植物油(花生油和紫苏籽油)为脂肪源配制5组等氮等能饲料。对照组(T1)以鱼油为唯一脂肪源,18:3 n-3/18:2 n-6为0.97,第二组(T2)以花生油为唯一脂肪源,18:3 n-3/18:2 n-6为0.02,其他3组实验饲料以花生油和紫苏籽油为脂肪源,且n-3/n-6比值分别为0.46(T3)、1.09(T4)和1.53(T5)。10周养殖实验结束后,分析各组鱼体的生长性能、血清中生长激素(GH)含量及不同部位肌肉中生长相关基因的表达水平。结果显示,与对照组相比,n-3/n-6对鱼体的脏体比(VSI)、饲料系数(FCR)、摄食率(FI)无显著影响,增重率(WGR)随n-3/n-6比值的增加先升后降,n-3/n-6比值等于1.09时,WGR达到峰值,为218.53%±24.32%。T2组血清GH含量显著高于其他处理组,随着n-3/n-6比值增大GH含量逐渐降低。背部肌肉中生长相关基因gh、ghr、igf-1和myod均表现为先升后降的趋势,igf-1r为先降后升,mstn无显著差异。红肌中gh、myod含量逐渐降低,ghr、mstn不存在显著差异,igf-1与背部肌肉相似,先升后降,igf-1r先降后升。腹部肌肉中gh、ghr和igf-1表达情况与背部肌肉一致,igf-1r和myod的表达不存在显著差异。T2组和T4组背部肌肉肌纤维数目存在显著差异,且T4组肌纤维较T2组粗。研究表明,n-3/n-6不仅在一定程度上提高鱼体的WGR,且不同比例的n-3/n-6水平会影响血清GH的含量,不同部位肌肉组织中GH/IGF轴生长相关基因的水平也受饲料中n-3/n-6比例的影响。
To analyze the influence of n-3/n-6 PUFAs levels on the growth performance and growth-related genes mRNA expression in common carp, five isonitrogenous and isoenergetic experimental diets based on fish oil and mixed vegetable oil were fed to the Cyprinus carpio haematopterus for 10 weeks. The control diet(T1) was only supplemented with fish oil as lipid source, n-3/n-6 ratio was 0.97. the second group(T2) was only supplemented with peanut oil, n-3/n-6 ratio was 0.02. n-3/n-6 ratio of the other three experiments groups were 0.46(T3), 1.09(T4) and 1.53(T5) with peanut oil and perilla seed oil as feed source. After 10 weeks cultivation experiments, we checked each group for fish growth performance, serum growth hormone and growth-related gene expression level in different parts of muscle tissue. The results indicated that there were no significant differences in the viscerosomatic index(VSI), feed coefficient(FCR), feed intake(FI)among the five tested groups. Weight gain rate of the fish(WGR) was raised first and subsequently declined with the increase of the n-3/n-6 ratio. WGR climbed the peak at T4 group. The serum GH levels of T2 group were significantly higher than those of others, and the level of GH in serum gradually reduced with the increase of of n-3/n-6 ratio. The expression levels of growthrelated genes in the dorsal muscles showed that gh, ghr, igf-1 and myod were enhanced first and subsequently declined. However, igf-1r displayed the opposite expressive trend, while there were no significant differences in mstn level among the five treated groups. In red muscle, the expression levels of gh and myod were gradually decreased with the increase of n-3/n-6 ratio, and the expressions of ghr and mstn exhibited no significant differences in five groups. Besides, the expressive trends of igf-1 and igf-1r were similar to that of dorsal muscles.Interestingly, the expressions of gh, ghr and igf-1 in the abdominal muscles were consistent with the dorsal muscles. But, the levels of igf-1r and myod manifested no significant difference in five tested groups. There were significant differences in the number of muscle fibers in the T2 group and T4 group, and the muscle fibers in T4 group were thicker than those of T2. This study showed that vegetable oil contributed to improve the WGR of the fish and the n-3/n-6 ratio of the diets not only affected the serum GH level, but also influenced the growth related genes of GH/IGF axis in different parts of the muscle tissues.
To analyze the influence of n-3/n-6 PUFAs levels on the growth performance and growth-related genes mRNA expression in common carp, five isonitrogenous and isoenergetic experimental diets based on fish oil and mixed vegetable oil were fed to the Cyprinus carpio haematopterus for 10 weeks. The control diet(T1) was only supplemented with fish oil as lipid source, n-3/n-6 ratio was 0.97. the second group(T2) was only supplemented with peanut oil, n-3/n-6 ratio was 0.02. n-3/n-6 ratio of the other three experiments groups were 0.46(T3), 1.09(T4) and 1.53(T5) with peanut oil and perilla seed oil as feed source. After 10 weeks cultivation experiments, we checked each group for fish growth performance, serum growth hormone and growth-related gene expression level in different parts of muscle tissue. The results indicated that there were no significant differences in the viscerosomatic index(VSI), feed coefficient(FCR), feed intake(FI)among the five tested groups. Weight gain rate of the fish(WGR) was raised first and subsequently declined with the increase of the n-3/n-6 ratio. WGR climbed the peak at T4 group. The serum GH levels of T2 group were significantly higher than those of others, and the level of GH in serum gradually reduced with the increase of of n-3/n-6 ratio. The expression levels of growthrelated genes in the dorsal muscles showed that gh, ghr, igf-1 and myod were enhanced first and subsequently declined. However, igf-1r displayed the opposite expressive trend, while there were no significant differences in mstn level among the five treated groups. In red muscle, the expression levels of gh and myod were gradually decreased with the increase of n-3/n-6 ratio, and the expressions of ghr and mstn exhibited no significant differences in five groups. Besides, the expressive trends of igf-1 and igf-1r were similar to that of dorsal muscles.Interestingly, the expressions of gh, ghr and igf-1 in the abdominal muscles were consistent with the dorsal muscles. But, the levels of igf-1r and myod manifested no significant difference in five tested groups. There were significant differences in the number of muscle fibers in the T2 group and T4 group, and the muscle fibers in T4 group were thicker than those of T2. This study showed that vegetable oil contributed to improve the WGR of the fish and the n-3/n-6 ratio of the diets not only affected the serum GH level, but also influenced the growth related genes of GH/IGF axis in different parts of the muscle tissues.
引文
[1]Trépanier M O,Hopperton K E,Orr S K,et al.N-3polyunsaturated fatty acids in animal models with neuroinflammation:an update[J].European Journal of Pharmacology,2016,785:187-206.
[2]周继术,吉红,王建华,等.鱼油对鲤生长及脂质代谢的影响[J].中国海洋大学学报,2008,38(2):275-280.Zhou J S,Ji H,Wang J H,et al.Influence of fish oil on growth and lipid metabolism in common carp(Cyprinus carpio)[J].Periodical of Ocean University of China,2008,38(2):275-280(in Chinese).
[3]潘瑜,毛述宏,关勇,等.饲料中不同脂肪源对鲤鱼生长性能、脂质代谢和抗氧化能力的影响[J].动物营养学报,2012,24(7):1368-1375.Pan Y,Mao S H,Guan Y,et al.Effects of different lipid sources in diets on growth performance,lipid metabolism and antioxidant ability of common carp(Cyprinus carpio)[J].Chinese Journal of Animal Nutrition,2012,24(7):1368-1375(in Chinese).
[4]Bell M V,Henderson R J,Sargent J R.The role of polyunsaturated fatty acids in fish[J].Comparative Biochemistry and Physiology-Part B:Comparative Biochemistry,1986,83(4):711-719.
[5]Turchini G M,Torstensen B E,Ng W K.Fish oil replacement in finfish nutrition[J].Reviews in Aquaculture,2010,1(1):10-57.
[6]王道尊,潘兆龙,梅志平.不同脂肪源饲料对青鱼生长的影响[J].水产学报,1989,13(4):370-374.Wang D Z,Pan Z L,Mei Z P.Effect of diets containing different lipid sources on growth of black carp(Mylopharyneodor piceus)[J].Journal of Fisheries of China,1989,13(4):370-374(in Chinese).
[7]刘玮,徐萍,任本根,等.不同脂肪源饲料对草鱼稚鱼生长的影响[J].水产学报,1995,19(4):362-365.Liu W,Xu P,Ren B G,et al.Effects of diets containing different lipids on growth of juvenile grass carp(Ctenopharyngodon idellus)[J].Journal of Fisheries of China,1995,19(4):362-365(in Chinese).
[8]卓丽欣,赵红霞,黄燕华,等.氧化鱼油对黄颡鱼幼鱼肠道健康的影响及精氨酸的干预作用[J].水产学报,2018,42(1):100-111.Zhuo L X,Zhao H X,Huang Y H,et al.Influence of oxidized fish oil on the intestinal health of juvenile yellow catfish(Pelteobagrus fulvidraco)and the use of arginine as an intervention measure[J].Journal of Fisheries of China,2018,42(1):100-111(in Chinese).
[9]Yu T C,Sinnhuber R O.Effect of dietary ω3 and ω6 fatty acids on growth and feed conversion efficiency of coho salmon(Oncorhynchus kisutch)[J].Aquaculture,1979,16(1):31-38.
[10]Glencross B D,Smith D M,Thomas M R,et al.The effect of dietary n-3 and n-6 fatty acid balance on the growth of the prawn Penaeus monodon[J].Aquaculture Nutrition,2002,8(1):43-51.
[11]Izquierdo M S,Obach A,Arantzamendi L,et al.Dietary lipid sources for seabream and seabass:growth performance,tissue composition and flesh quality[J].Aquaculture Nutrition,2003,9(6):397-407.
[12]Tan X Y,Luo Z,Xie P,et al.Effect of dietary linolenic acid/linoleic acid ratio on growth performance,hepatic fatty acid profiles and intermediary metabolism of juvenile yellow catfish Pelteobagrus fulvidraco[J].Aquaculture,2009,296(1-2):96-101.
[13]Caballero M J,Obach A,Rosenlund G,et al.Impact of different dietary lipid sources on growth,lipid digestibility,tissue fatty acid composition and histology of rainbow trout,Oncorhynchus mykiss[J].Aquaculture,2002,214(1-4):253-271.
[14]Grisdale Helland B,Ruyter B,Rosenlund G,et al.Influence of high contents of dietary soybean oil on growth,feed utilization,tissue fatty acid composition,heart histology and standard oxygen consumption of Atlantic salmon(Salmo salar)raised at two temperatures[J].Aquaculture,2002,207(3-4):311-329.
[15]Takeuchi T.Essential fatty acid requirements in carp[J].Archiv Für Tierern?hrung,1996,49(1):23-32.
[16]Kennish J M,Sharp-Dahl J L,Chambers K A,et al.The effect of a herring diet on lipid composition,fatty acid composition,and cholesterol levels in the muscle tissue of pen-reared chinook salmon(Oncorhynchus tshawytscha)[J].Aquaculture,1992,108(3-4):309-322.
[17]张媛媛,刘波,戈贤平,等.不同脂肪源对异育银鲫生长性能、机体成分、血清生化指标、体组织脂肪酸组成及脂质代谢的影响[J].水产学报,2012,36(7):1111-1118.Zhang Y Y,Liu B,Ge X P,et al.Effect of dietary oil sources on growth performance,body composition,the serum biochemical indices,fatty acids composition and lipid metabolism of Carassius auratus gibelio[J].Journal of Fisheries of China,2012,36(7):1111-1118(in Chinese).
[18]李新,谭晓晨,张健,等.不同植物油对大规格吉富罗非鱼生长性能、组织脂肪酸组成的影响[J].动物营养学报,2016,28(10):3327-3336.Li X,Tan X C,Zhang J,et al.Effects of different vegetable oils on growth performance and tissue fatty acid composition of large size genetically improved farmed tilapia(Oreochromis niloticus)[J].Chinese Journal of Animal Nutrition,2016,28(10):3327-3336(in Chinese).
[19]李新,杨瑞斌,钱雪桥,等.不同植物脂肪源对吉富罗非鱼生长性能、肌肉脂肪酸和生理生化指标的影响[J].水生生物学报,2016,40(4):672-680.Li X,Yang R B,Qian X Q,et al.Effect of dietary vegetable oils when fed to genetically improved farmed tilapia strain of Nile tilapia Oreochromis niloticus[J].Acta Hydrobiologica Sinica,2016,40(4):672-680(in Chinese).
[20]彭祥和,李法见,林仕梅,等.亚麻籽油对罗非鱼生长性能及肉品质的影响[J].动物营养学报,2015,27(1):147-153.Peng X H,Li F J,Lin S M,et al.Effects of linseed oil on growth performance and meat quality of tilapia(Oreochromis niloticus)[J].Chinese Journal of Animal Nutrition,2015,27(1):147-153(in Chinese).
[21]左然涛,麦康森,徐玮,等.脂肪酸对鱼类免疫系统的影响及调控机制研究进展[J].水产学报,2015,39(7):1079-1088.Zuo R T,Mai K S,Xu W,et al.Advance of studies on the effects of fatty acids on immune responses and nutritional regulation mechanism in fish species[J].Journal of Fisheries of China,2015,39(7):1079-1088(in Chinese).
[22]张洪涛,单雷,毕玉平.n-6和n-3多不饱和脂肪酸在人和动物体内的功能关系[J].山东农业科学,2006(2):115-120.Zhang H T,Shan L,Bi Y P.Functional relationship between n-6 and n-3 polyunsaturated fatty acids in the bodies of human and animal[J].Shandong Agricultural Sciences,2006(2):115-120(in Chinese).
[23]Hu W,Zhu Z Y.Integration mechanisms of transgenes and population fitness of GH transgenic fish[J].Science China Life Sciences,2010,53(4):401-408.
[24]谢帝芝,于若梦,陈芳,等.饲料LNA/LA比对鲤幼鱼生长性能,肝脏脂肪酸组成及Δ6 fad,elovl5 mRNA表达的影响[J].水产学报,2017,41(5):757-765.Xie D Z,Yu R M,Chen F,et al.Effects of dietary LNA/LA ratio on growth,hepatic fatty acid profiles and△6fad,elovl5 mRNA expression in common carp(Cyprinus carpio)[J].Journal of Fisheries of China,2017,41(5):757-765(in Chinese).
[25]Sargent J R,Tocher D R,Bell J G.The lipids[M]//Halver J E,Hardy R W.Fish Nutrition.3rd ed.San Diego,California:Elsevier,2002:181-257.
[26]Olsen R E,Henderson R J,McAndrew B J.The conversion of linoleic acid and linolenic acid to longer chain polyunsaturated fatty acids by tilapia(Oreochromis nilotica)in vivo[J].Fish Physiology and Biochemistry,1990,8(3):261-270.
[27]Francis D S,Peters D J,Turchini G M.Apparent in vivoΔ-6 desaturase activity,efficiency,and affinity are affected by total dietary C18 PUFA in the freshwater fish murray cod[J].Journal of Agricultural and Food Chemistry,2009,57(10):4381-4190.
[28]Blanchard G,Makombu J G,Kestemont P.Influence of different dietary 18:3n-3/18:2n-6 ratio on growth performance,fatty acid composition and hepatic ultrastructure in Eurasian perch,Perca fluviatilis[J].Aquaculture,2008,284(1-4):144-150.
[29]Biga P R,Cain K D,Hardy R W,et al.Growth hormone differentially regulates muscle myostatin1 and-2 and increases circulating cortisol in rainbow trout(Oncorhynchus mykiss)[J].General and Comparative Endocrinology,2004,138(1):32-41.
[30]李晶,沙长青,杨成辉.重组鲑鱼生长激素基因酵母对鲤鱼苗促生长作用研究[J].中国饲料,2004(18):30-33.Li J,Sha C Q,Yang C H.Recombinant salmon growth hormone gene yeast to study the growth effect of carp seedling[J].China Feed,2004(18):30-33(in Chinese).
[31]Moriyama S,Swanson P,Nishii M,et al.Development of a homologous radioimmunoassay for coho salmon insulin-like growth factor-I[J].General and Comparative Endocrinology,1994,96(1):149-161.
[32]刘月星,马洪雨,马春艳,等.水产动物重要经济性状相关功能基因的研究进展[J].生物技术通报,2014(2):30-40.Liu Y X,Ma H Y,Ma C Y,et al.Research progress of functional genes related with important economic traits for aquatic animals[J].Biotechnology Bulletin,2014(2):30-40(in Chinese).
[33]代向燕,张玮,卓子见,等.鱼类生长的神经内分泌调控[J].中国科学:生命科学,2015,58(2):137-147.Dai X Y,Zhang W,Zhuo Z J,et al.Neuroendocrine regulation of somatic growth in fishes[J].Science China Life Sciences,2015,58(2):137-147.
[34]De-Santis C,Jerry D R.Candidate growth genes in finfish-where should we be looking?[J].Aquaculture,2007,272(1-4):22-38.
[35]Tan X G,Rotllant J,Li H Q,et al.SmyD1,a histone methyltransferase,is required for myofibril organization and muscle contraction in zebrafish embryos[J].Proceedings of the National Academy of Sciences of the United States of America,2006,103(8):2713-2718.
[36]Johnston I A.Environment and plasticity of myogenesis in teleost fish[J].Journal of Experimental Biology,2006,209(12):2249-2264.
[37]Eppler E,Berishvili G,Mazel P,et al.Distinct organspecific up-and down-regulation of IGF-I and IGF-IImRNA in various organs of a GH-overexpressing transgenic Nile tilapia[J].Transgenic Research,2010,19(2):231-240.
[38]Yuan X N,Jiang X Y,Pu J W,et al.Functional conservation and divergence of duplicated insulin-like growth factor 2 genes in grass carp(Ctenopharyngodon idellus)[J].Gene,2011,470(1-2):46-52.
[39]Vong Q P,Chan K M,Cheng C H.Quantification of common carp(Cyprinus carpio)IGF-I and IGF-IImRNA by real-time PCR:differential regulation of expression by GH[J].Journal of Endocrinology,2003,178(3):513-521.
[40]S?Nger A M,Stoiber W.7-Muscle fiber diversity and plasticity[J].Fish Physiology,2001,18:187-250.
[41]赵浩斌,彭扣,王玉凤,等.鱼类肌肉生长抑制素研究进展[J].水生生物学报,2006,30(2):227-231.Zhao H B,Peng K,Wang Y F,et al.Progress of studies on myostatin of fish[J].Acta Hydrobiologica Sinica,2006,30(2):227-231(in Chinese).
[42]成嘉,褚武英,张建社.鱼类肌肉组织发生和分化相关基因的研究进展[J].生命科学研究,2010,14(4):355-362.Cheng J,Chu W Y,Zhang J S.Progresses and perspectives of the studies on fish muscle-related genes and their expression[J].Life Science Research,2010,14(4):355-362(in Chinese).
[43]Galt N J,Froehlich J M,Meyer B M,et al.High-fat diet reduces local myostatin-1,paralog expression and alters skeletal muscle lipid content in rainbow trout,Oncorhynchus mykiss[J].Fish Physiology and Biochemistry,2014,40(3):875-886.
[44]Mareco E A,de la Serrana D G,Johnston I A,et al.Characterization of the transcriptome of fast and slow muscle myotomal fibres in the pacu(Piaractus mesopotamicus)[J].BMC Genomics,2015,16(1):182.
[45]Bower N I,Johnston I A.Transcriptional regulation of the IGF signaling pathway by amino acids and insulinlike growth factors during myogenesis in Atlantic salmon[J].PLoS One,2010,5(6):e11100.
[46]Vélez E J,Lutfi E,Jiménez-Amilburu V,et al.IGF-I and amino acids effects through TOR signaling on proliferation and differentiation of gilthead sea bream cultured myocytes[J].General and Comparative Endocrinology,2014,205:296-304.
[47]Azizi S,Nematollahi M A,Mojazi A B,et al.IGF-I and IGF-II effects on local IGF system and signaling pathways in gilthead sea bream(Sparus aurata)cultured myocytes[J].General and Comparative Endocrinology,2016,232:7-16.
[48]Duarte-Guterman P,Navarro-Martín L,Trudeau V L.Mechanisms of crosstalk between endocrine systems:regulation of sex steroid hormone synthesis and action by thyroid hormones[J].General and Comparative Endocrinology,2014,203:69-85.
[2]周继术,吉红,王建华,等.鱼油对鲤生长及脂质代谢的影响[J].中国海洋大学学报,2008,38(2):275-280.Zhou J S,Ji H,Wang J H,et al.Influence of fish oil on growth and lipid metabolism in common carp(Cyprinus carpio)[J].Periodical of Ocean University of China,2008,38(2):275-280(in Chinese).
[3]潘瑜,毛述宏,关勇,等.饲料中不同脂肪源对鲤鱼生长性能、脂质代谢和抗氧化能力的影响[J].动物营养学报,2012,24(7):1368-1375.Pan Y,Mao S H,Guan Y,et al.Effects of different lipid sources in diets on growth performance,lipid metabolism and antioxidant ability of common carp(Cyprinus carpio)[J].Chinese Journal of Animal Nutrition,2012,24(7):1368-1375(in Chinese).
[4]Bell M V,Henderson R J,Sargent J R.The role of polyunsaturated fatty acids in fish[J].Comparative Biochemistry and Physiology-Part B:Comparative Biochemistry,1986,83(4):711-719.
[5]Turchini G M,Torstensen B E,Ng W K.Fish oil replacement in finfish nutrition[J].Reviews in Aquaculture,2010,1(1):10-57.
[6]王道尊,潘兆龙,梅志平.不同脂肪源饲料对青鱼生长的影响[J].水产学报,1989,13(4):370-374.Wang D Z,Pan Z L,Mei Z P.Effect of diets containing different lipid sources on growth of black carp(Mylopharyneodor piceus)[J].Journal of Fisheries of China,1989,13(4):370-374(in Chinese).
[7]刘玮,徐萍,任本根,等.不同脂肪源饲料对草鱼稚鱼生长的影响[J].水产学报,1995,19(4):362-365.Liu W,Xu P,Ren B G,et al.Effects of diets containing different lipids on growth of juvenile grass carp(Ctenopharyngodon idellus)[J].Journal of Fisheries of China,1995,19(4):362-365(in Chinese).
[8]卓丽欣,赵红霞,黄燕华,等.氧化鱼油对黄颡鱼幼鱼肠道健康的影响及精氨酸的干预作用[J].水产学报,2018,42(1):100-111.Zhuo L X,Zhao H X,Huang Y H,et al.Influence of oxidized fish oil on the intestinal health of juvenile yellow catfish(Pelteobagrus fulvidraco)and the use of arginine as an intervention measure[J].Journal of Fisheries of China,2018,42(1):100-111(in Chinese).
[9]Yu T C,Sinnhuber R O.Effect of dietary ω3 and ω6 fatty acids on growth and feed conversion efficiency of coho salmon(Oncorhynchus kisutch)[J].Aquaculture,1979,16(1):31-38.
[10]Glencross B D,Smith D M,Thomas M R,et al.The effect of dietary n-3 and n-6 fatty acid balance on the growth of the prawn Penaeus monodon[J].Aquaculture Nutrition,2002,8(1):43-51.
[11]Izquierdo M S,Obach A,Arantzamendi L,et al.Dietary lipid sources for seabream and seabass:growth performance,tissue composition and flesh quality[J].Aquaculture Nutrition,2003,9(6):397-407.
[12]Tan X Y,Luo Z,Xie P,et al.Effect of dietary linolenic acid/linoleic acid ratio on growth performance,hepatic fatty acid profiles and intermediary metabolism of juvenile yellow catfish Pelteobagrus fulvidraco[J].Aquaculture,2009,296(1-2):96-101.
[13]Caballero M J,Obach A,Rosenlund G,et al.Impact of different dietary lipid sources on growth,lipid digestibility,tissue fatty acid composition and histology of rainbow trout,Oncorhynchus mykiss[J].Aquaculture,2002,214(1-4):253-271.
[14]Grisdale Helland B,Ruyter B,Rosenlund G,et al.Influence of high contents of dietary soybean oil on growth,feed utilization,tissue fatty acid composition,heart histology and standard oxygen consumption of Atlantic salmon(Salmo salar)raised at two temperatures[J].Aquaculture,2002,207(3-4):311-329.
[15]Takeuchi T.Essential fatty acid requirements in carp[J].Archiv Für Tierern?hrung,1996,49(1):23-32.
[16]Kennish J M,Sharp-Dahl J L,Chambers K A,et al.The effect of a herring diet on lipid composition,fatty acid composition,and cholesterol levels in the muscle tissue of pen-reared chinook salmon(Oncorhynchus tshawytscha)[J].Aquaculture,1992,108(3-4):309-322.
[17]张媛媛,刘波,戈贤平,等.不同脂肪源对异育银鲫生长性能、机体成分、血清生化指标、体组织脂肪酸组成及脂质代谢的影响[J].水产学报,2012,36(7):1111-1118.Zhang Y Y,Liu B,Ge X P,et al.Effect of dietary oil sources on growth performance,body composition,the serum biochemical indices,fatty acids composition and lipid metabolism of Carassius auratus gibelio[J].Journal of Fisheries of China,2012,36(7):1111-1118(in Chinese).
[18]李新,谭晓晨,张健,等.不同植物油对大规格吉富罗非鱼生长性能、组织脂肪酸组成的影响[J].动物营养学报,2016,28(10):3327-3336.Li X,Tan X C,Zhang J,et al.Effects of different vegetable oils on growth performance and tissue fatty acid composition of large size genetically improved farmed tilapia(Oreochromis niloticus)[J].Chinese Journal of Animal Nutrition,2016,28(10):3327-3336(in Chinese).
[19]李新,杨瑞斌,钱雪桥,等.不同植物脂肪源对吉富罗非鱼生长性能、肌肉脂肪酸和生理生化指标的影响[J].水生生物学报,2016,40(4):672-680.Li X,Yang R B,Qian X Q,et al.Effect of dietary vegetable oils when fed to genetically improved farmed tilapia strain of Nile tilapia Oreochromis niloticus[J].Acta Hydrobiologica Sinica,2016,40(4):672-680(in Chinese).
[20]彭祥和,李法见,林仕梅,等.亚麻籽油对罗非鱼生长性能及肉品质的影响[J].动物营养学报,2015,27(1):147-153.Peng X H,Li F J,Lin S M,et al.Effects of linseed oil on growth performance and meat quality of tilapia(Oreochromis niloticus)[J].Chinese Journal of Animal Nutrition,2015,27(1):147-153(in Chinese).
[21]左然涛,麦康森,徐玮,等.脂肪酸对鱼类免疫系统的影响及调控机制研究进展[J].水产学报,2015,39(7):1079-1088.Zuo R T,Mai K S,Xu W,et al.Advance of studies on the effects of fatty acids on immune responses and nutritional regulation mechanism in fish species[J].Journal of Fisheries of China,2015,39(7):1079-1088(in Chinese).
[22]张洪涛,单雷,毕玉平.n-6和n-3多不饱和脂肪酸在人和动物体内的功能关系[J].山东农业科学,2006(2):115-120.Zhang H T,Shan L,Bi Y P.Functional relationship between n-6 and n-3 polyunsaturated fatty acids in the bodies of human and animal[J].Shandong Agricultural Sciences,2006(2):115-120(in Chinese).
[23]Hu W,Zhu Z Y.Integration mechanisms of transgenes and population fitness of GH transgenic fish[J].Science China Life Sciences,2010,53(4):401-408.
[24]谢帝芝,于若梦,陈芳,等.饲料LNA/LA比对鲤幼鱼生长性能,肝脏脂肪酸组成及Δ6 fad,elovl5 mRNA表达的影响[J].水产学报,2017,41(5):757-765.Xie D Z,Yu R M,Chen F,et al.Effects of dietary LNA/LA ratio on growth,hepatic fatty acid profiles and△6fad,elovl5 mRNA expression in common carp(Cyprinus carpio)[J].Journal of Fisheries of China,2017,41(5):757-765(in Chinese).
[25]Sargent J R,Tocher D R,Bell J G.The lipids[M]//Halver J E,Hardy R W.Fish Nutrition.3rd ed.San Diego,California:Elsevier,2002:181-257.
[26]Olsen R E,Henderson R J,McAndrew B J.The conversion of linoleic acid and linolenic acid to longer chain polyunsaturated fatty acids by tilapia(Oreochromis nilotica)in vivo[J].Fish Physiology and Biochemistry,1990,8(3):261-270.
[27]Francis D S,Peters D J,Turchini G M.Apparent in vivoΔ-6 desaturase activity,efficiency,and affinity are affected by total dietary C18 PUFA in the freshwater fish murray cod[J].Journal of Agricultural and Food Chemistry,2009,57(10):4381-4190.
[28]Blanchard G,Makombu J G,Kestemont P.Influence of different dietary 18:3n-3/18:2n-6 ratio on growth performance,fatty acid composition and hepatic ultrastructure in Eurasian perch,Perca fluviatilis[J].Aquaculture,2008,284(1-4):144-150.
[29]Biga P R,Cain K D,Hardy R W,et al.Growth hormone differentially regulates muscle myostatin1 and-2 and increases circulating cortisol in rainbow trout(Oncorhynchus mykiss)[J].General and Comparative Endocrinology,2004,138(1):32-41.
[30]李晶,沙长青,杨成辉.重组鲑鱼生长激素基因酵母对鲤鱼苗促生长作用研究[J].中国饲料,2004(18):30-33.Li J,Sha C Q,Yang C H.Recombinant salmon growth hormone gene yeast to study the growth effect of carp seedling[J].China Feed,2004(18):30-33(in Chinese).
[31]Moriyama S,Swanson P,Nishii M,et al.Development of a homologous radioimmunoassay for coho salmon insulin-like growth factor-I[J].General and Comparative Endocrinology,1994,96(1):149-161.
[32]刘月星,马洪雨,马春艳,等.水产动物重要经济性状相关功能基因的研究进展[J].生物技术通报,2014(2):30-40.Liu Y X,Ma H Y,Ma C Y,et al.Research progress of functional genes related with important economic traits for aquatic animals[J].Biotechnology Bulletin,2014(2):30-40(in Chinese).
[33]代向燕,张玮,卓子见,等.鱼类生长的神经内分泌调控[J].中国科学:生命科学,2015,58(2):137-147.Dai X Y,Zhang W,Zhuo Z J,et al.Neuroendocrine regulation of somatic growth in fishes[J].Science China Life Sciences,2015,58(2):137-147.
[34]De-Santis C,Jerry D R.Candidate growth genes in finfish-where should we be looking?[J].Aquaculture,2007,272(1-4):22-38.
[35]Tan X G,Rotllant J,Li H Q,et al.SmyD1,a histone methyltransferase,is required for myofibril organization and muscle contraction in zebrafish embryos[J].Proceedings of the National Academy of Sciences of the United States of America,2006,103(8):2713-2718.
[36]Johnston I A.Environment and plasticity of myogenesis in teleost fish[J].Journal of Experimental Biology,2006,209(12):2249-2264.
[37]Eppler E,Berishvili G,Mazel P,et al.Distinct organspecific up-and down-regulation of IGF-I and IGF-IImRNA in various organs of a GH-overexpressing transgenic Nile tilapia[J].Transgenic Research,2010,19(2):231-240.
[38]Yuan X N,Jiang X Y,Pu J W,et al.Functional conservation and divergence of duplicated insulin-like growth factor 2 genes in grass carp(Ctenopharyngodon idellus)[J].Gene,2011,470(1-2):46-52.
[39]Vong Q P,Chan K M,Cheng C H.Quantification of common carp(Cyprinus carpio)IGF-I and IGF-IImRNA by real-time PCR:differential regulation of expression by GH[J].Journal of Endocrinology,2003,178(3):513-521.
[40]S?Nger A M,Stoiber W.7-Muscle fiber diversity and plasticity[J].Fish Physiology,2001,18:187-250.
[41]赵浩斌,彭扣,王玉凤,等.鱼类肌肉生长抑制素研究进展[J].水生生物学报,2006,30(2):227-231.Zhao H B,Peng K,Wang Y F,et al.Progress of studies on myostatin of fish[J].Acta Hydrobiologica Sinica,2006,30(2):227-231(in Chinese).
[42]成嘉,褚武英,张建社.鱼类肌肉组织发生和分化相关基因的研究进展[J].生命科学研究,2010,14(4):355-362.Cheng J,Chu W Y,Zhang J S.Progresses and perspectives of the studies on fish muscle-related genes and their expression[J].Life Science Research,2010,14(4):355-362(in Chinese).
[43]Galt N J,Froehlich J M,Meyer B M,et al.High-fat diet reduces local myostatin-1,paralog expression and alters skeletal muscle lipid content in rainbow trout,Oncorhynchus mykiss[J].Fish Physiology and Biochemistry,2014,40(3):875-886.
[44]Mareco E A,de la Serrana D G,Johnston I A,et al.Characterization of the transcriptome of fast and slow muscle myotomal fibres in the pacu(Piaractus mesopotamicus)[J].BMC Genomics,2015,16(1):182.
[45]Bower N I,Johnston I A.Transcriptional regulation of the IGF signaling pathway by amino acids and insulinlike growth factors during myogenesis in Atlantic salmon[J].PLoS One,2010,5(6):e11100.
[46]Vélez E J,Lutfi E,Jiménez-Amilburu V,et al.IGF-I and amino acids effects through TOR signaling on proliferation and differentiation of gilthead sea bream cultured myocytes[J].General and Comparative Endocrinology,2014,205:296-304.
[47]Azizi S,Nematollahi M A,Mojazi A B,et al.IGF-I and IGF-II effects on local IGF system and signaling pathways in gilthead sea bream(Sparus aurata)cultured myocytes[J].General and Comparative Endocrinology,2016,232:7-16.
[48]Duarte-Guterman P,Navarro-Martín L,Trudeau V L.Mechanisms of crosstalk between endocrine systems:regulation of sex steroid hormone synthesis and action by thyroid hormones[J].General and Comparative Endocrinology,2014,203:69-85.