共轭亚油酸对草鱼肝脏和肌肉组织的细胞学形态、抗氧化能力以及脂肪代谢相关基因表达的影响
|
摘要
采用65d的生长实验研究共轭亚油酸(CLA)对草鱼肝脏和肌肉细胞学形态、抗氧化能力以及脂肪代谢相关基因表达的影响。共配制7种近似等氮(粗蛋白:36 g/100 g)等脂(粗脂肪:4.5 g/100 g)的饲料:0 (对照组)、0.5 (CLA0.5)、1 (CLA1)、1.5 (CLA1.5)、2 (CLA2)、2.5 (CLA2.5)和3 g/100 g CLA (CLA3)。每个饲料组均设置3个生物学重复,实验鱼初始体重约为(5.08±0.08) g。实验结果显示:与对照组相比, CLA0.5-CLA2.5组草鱼肝脏和肌肉组织细胞学形态均无明显异常,但CLA3组肝细胞线粒体空泡化,内质网肿胀,排列过于紧密;肌细胞肌节结构松散,肌原纤维降解;在CLA1.5-CLA2.5组肝脏和肌肉中超氧化物歧化酶(SOD)、总抗氧化能力(T-AOC)和谷胱甘肽过氧化物酶(GPx)活力均显著上升(P<0.05);在CLA1.5-CLA3组中肝脏过氧化氢酶(CAT)活力均显著高于对照组(P<0.05),但谷胱甘肽还原酶(GR)活力无显著变化(P>0.05),且在CLA0.5-CLA3组中肌肉CAT和GR活力均无显著变化(P>0.05)。CLA1.5-CLA2组肝脏中MDA含量显著低于对照组(P<0.05),而CLA2.5-CLA3组MDA含量显著高于对照组(P<0.05)。CLA3组肌肉中MDA含量显著高于对照组(P<0.05)。与对照组相比,在CLA1.5-CLA2.5组肝脏和肌肉中乙酰辅酶A羧化酶(ACC) mRNA表达显著下调(P<0.05),脂蛋白脂酶(LPL)、激素敏感性脂肪酶(HSL)和过氧化物酶体增殖物激活受体α(PPARα)均显著上调表达(P<0.05)。在CLA2-CLA3组肝脏以及CLA1.5-CLA2和CLA3组肌肉中脂肪酸去饱和酶2(FAD2)mRNA显著上调表达(P<0.05)。过氧化物酶体增殖物激活受体γ(PPARγ)mRNA在CLA1-CLA3组肝脏中显著下调表达(P<0.05),而其在肌肉中均无显著变化(P>0.05)。综上所述,结合2%CLA在不影响草鱼的生长和饲料利用,且显著降低肝脏脂肪积累,研究结果建议, 2%CLA在不影响草鱼肝脏和肌肉组织细胞学形态的前提下,可有效提高其肝脏和肌肉组织抗氧化能力并促进其脂肪分解代谢以及抑制脂肪合成代谢相关基因表达。
A 65-day trial was conducted to investigate effects of CLA on cytological morphology, activity of antioxidant enzymes, and gene expressions of lipid metabolism in the liver and muscle of grass carp. Seven isonitrogenous(crude protein: 36 g/100 g) and isolipidic(crude lipid: 4.5 g/100 g) diets were formulated: 0 CLA(control), 0.5%(CLA0.5), 1%(CLA1), 1.5(CLA1.5), 2(CLA2), 2.5(CLA2.5), and 3% CLA(CLA3). Three biological replicates of fish were assigned to each diet with the initial body weight at(5.08±0.08) g. Endoplasmic reticulum in the liver, and loose sarcolemma of myocyte and degradation of myofibril in the muscle when compared with the control fish. The activities of superoxide dismutase(SOD) and glutathione peroxidase(GPx), and the total antioxidant capacity(T-AOC)both in the liver and muscle of CLA1.5-CLA2.5 groups significantly(P<0.05) increased compared with control group.The activities of catalase(CAT) in the liver of CLA1.5-CLA3 groups were significantly(P<0.05) higher than that of control group, however, there was no significant(P>0.05) change in the glutathione reductase(GR) activity in the liver.The MDA contents in the liver of fish fed with CLA1.5-CLA2 diets were significantly(P<0.05) lower than that of control group, however, the MDA contents in the liver of fish fed with CLA2.5-CLA3 diets significantly(P<0.05) increased compared with control group. In contrast, fish fed with CLA3 had significantly higher MDA contents in the muscle(P<0.05) compared with control group. Compared with the control fish, the mRNA level of acetyl-CoA carboxylase(ACC) both in the liver and muscle of fish fed with CLA1.5-CLA2.5 diets were significantly(P<0.05)down-regulated; the mRNA levels of lipoprotein lipase(LPL), hormone-sensitive lipase(HSL), and peroxisome proliferator-activated receptor alpha(PPARα) of fish fed with CLA1.5-CLA2.5 diets were significantly(P<0.05) up-regulated; the mRNA levels of fatty acid desaturase 2(FAD2) in the liver of fish fed with CLA2-CLA3 diets and in the muscle of fish fed with CLA1.5-CLA2 and CLA3 diets were significantly(P<0.05) up-regulated; the mRNA level of peroxisome proliferator-activated receptor gamma(PPARγ) in the liver of fish fed with CLA1-CLA3 diets was significantly(P<0.05) down-regulated. Conversely, there was no significant(P>0.05) difference for the PPARγ mRNA level in the muscle of fish fed with CLA0.5-CLA3 diets. In conclusion, supplementation of 2% CLA could significantly decrease the lipid contents in the liver and muscle without affecting cytological morphology of liver and muscle, the growth and feed utilization of grass carp by improving the antioxidant capacities in the liver and muscle.
A 65-day trial was conducted to investigate effects of CLA on cytological morphology, activity of antioxidant enzymes, and gene expressions of lipid metabolism in the liver and muscle of grass carp. Seven isonitrogenous(crude protein: 36 g/100 g) and isolipidic(crude lipid: 4.5 g/100 g) diets were formulated: 0 CLA(control), 0.5%(CLA0.5), 1%(CLA1), 1.5(CLA1.5), 2(CLA2), 2.5(CLA2.5), and 3% CLA(CLA3). Three biological replicates of fish were assigned to each diet with the initial body weight at(5.08±0.08) g. Endoplasmic reticulum in the liver, and loose sarcolemma of myocyte and degradation of myofibril in the muscle when compared with the control fish. The activities of superoxide dismutase(SOD) and glutathione peroxidase(GPx), and the total antioxidant capacity(T-AOC)both in the liver and muscle of CLA1.5-CLA2.5 groups significantly(P<0.05) increased compared with control group.The activities of catalase(CAT) in the liver of CLA1.5-CLA3 groups were significantly(P<0.05) higher than that of control group, however, there was no significant(P>0.05) change in the glutathione reductase(GR) activity in the liver.The MDA contents in the liver of fish fed with CLA1.5-CLA2 diets were significantly(P<0.05) lower than that of control group, however, the MDA contents in the liver of fish fed with CLA2.5-CLA3 diets significantly(P<0.05) increased compared with control group. In contrast, fish fed with CLA3 had significantly higher MDA contents in the muscle(P<0.05) compared with control group. Compared with the control fish, the mRNA level of acetyl-CoA carboxylase(ACC) both in the liver and muscle of fish fed with CLA1.5-CLA2.5 diets were significantly(P<0.05)down-regulated; the mRNA levels of lipoprotein lipase(LPL), hormone-sensitive lipase(HSL), and peroxisome proliferator-activated receptor alpha(PPARα) of fish fed with CLA1.5-CLA2.5 diets were significantly(P<0.05) up-regulated; the mRNA levels of fatty acid desaturase 2(FAD2) in the liver of fish fed with CLA2-CLA3 diets and in the muscle of fish fed with CLA1.5-CLA2 and CLA3 diets were significantly(P<0.05) up-regulated; the mRNA level of peroxisome proliferator-activated receptor gamma(PPARγ) in the liver of fish fed with CLA1-CLA3 diets was significantly(P<0.05) down-regulated. Conversely, there was no significant(P>0.05) difference for the PPARγ mRNA level in the muscle of fish fed with CLA0.5-CLA3 diets. In conclusion, supplementation of 2% CLA could significantly decrease the lipid contents in the liver and muscle without affecting cytological morphology of liver and muscle, the growth and feed utilization of grass carp by improving the antioxidant capacities in the liver and muscle.
引文
[1]Chinese Fishery Statistical Yearbook[M].Beijing:Chinese Agricultural Press.2017,25[中国渔业统计年鉴.北京:中国农业出版社.2017,25]
[2]Wang K Y,Miao C H,Huang J L,et al.Changes in main biochemical indices and Acetyl-CoA Carboxylase 1mRNA expression of crass carp fed a high-fat diet[J].Chinese Journal of Animal Nutrition,2012,24(12):2375-2383[汪开毓,苗常鸿,黄锦炉,等.投喂高脂饲料后草鱼主要生化指标和乙酰辅酶A羧化酶1 mRNA表达的变化.动物营养学报,2012,24(12):2375-2383]
[3]Wu C Y.The pharmacodynamic function study of Sedum sarmentosum extract in grass carp fatty liver model[D].Thesis for Master of Veterinary.Sichuan Agricultural University,Chengdu.2013[吴春艳.垂盆草提取物对草鱼脂肪肝模型药效作用研究.四川农业大学硕士论文,成都.2013]
[4]Banni S.Conjugated linoleic acid metabolism[J].Current Opinion in Lipidology,2002,13(3):261-266
[5]Yang B,Chen H,Stanton C,et al.Review of the roles of conjugated linoleic acid in health and disease[J].Journal of Functional Foods,2015,15:314-325
[6]Twibell R G,Watkins B A,Rogers L,et al.Effects of dietary conjugated linoleic acids on hepatic and muscle lipids in hybrid striped bass[J].Lipids,2000,35(2):155-161
[7]Twibell R G,Watkins B A,Brown P B.Dietary conjugated linoleic acids and lipid source alter fatty acid composition of juvenile yellow perch,Perca flavescens[J].The Journal of Nutrition,2001,131(9):2322-2328
[8]Tan X Y,Luo Z,Xie P,et al.Effect of dietary conjugated linoleic acid(CLA)on growth performance,body composition and hepatic intermediary metabolism in juvenile yellow catfish Pelteobagrus fulvidraco[J].Aquaculture,2010,310(1-2):186-191
[9]Choi B D,Kang S J,Ha Y L,et al.Accumulation of Conjugated Linoleic Acid(CLA)in Tissues of Fish Fed Diets Containing Various Levels of CLA[M].Quality Attributes of Muscle Foods.1999,61-67
[10]Twibell R G,Wilson R P.Effects of dietary conjugated linoleic acids and total dietary lipid concentrations on growth responses of juvenile channel catfish,Ictalurus punctatus[J].Aquaculture,2003,221(1):621-628
[11]Figueiredo-Silva A C,Rema P,Bandarra N M,et al.Effects of dietary conjugated linoleic acid on growth,nutrient utilization,body composition,and hepatic lipogenesis in rainbow trout juveniles(Oncorhynchus mykiss)[J].Aquaculture,2005,248(1):163-172
[12]Makol A,Torrecillas S,Fernándezvaquero A,et al.Effect of conjugated linoleic acid on dietary lipids utilization,liver morphology and selected immune parameters in sea bass juveniles(Dicentrarchus labrax)[J].Comparative Biochemistry&Physiology,2009,154(2):179-187
[13]Makol A,Torrecillas S,Caballero M J,et al.Effect of long term feeding with conjugated linoleic acid(CLA)in growth performance and lipid metabolism of European sea bass(Dicentrarchus labrax)[J].Aquaculture,2012,368-369(1):129-137
[14]Zuo R T,Ai Q H,Mai K S,et al.Effects of conjugated linoleic acid on growth,non-specific immunity,antioxidant capacity,lipid deposition and related gene expression in juvenile large yellow croaker(Larmichthys crocea)fed soyabean oil-based diets[J].British Journal of Nutrition,2013,110(7):1220-1232
[15]Zhao Z Y.Effects of conjugated linoleic acid on lipid metabolism,immunity,flesh quality and PPAR gene expression in large Yellow croaker[D].Thesis for Doctor of Science.Zhejiang University,Hangzhou.2008[赵占宇.共轭亚油酸(CLA)对大黄鱼脂肪代谢、免疫、肉品质及PPAR基因表达的影响.浙江大学博士论文,杭州.2008]
[16]Kennedy S R,Campbell P J,Porter A,et al.Influence of dietary conjugated linoleic acid(CLA)on lipid and fatty acid composition in liver and flesh of Atlantic salmon(Salmo salar)[J].Comparative Biochemistry&Physiology,2005,141(2):168-178
[17]Leaver M J,Tocher D R,Obach A,et al.Effect of dietary conjugated linoleic acid(CLA)on lipid composition,metabolism and gene expression in Atlantic salmon(Salmo salar)tissues[J].Comparative Biochemistry and Physiology,2006,145(2):258-267
[18]Dong G F,Zou Q,Wang H,et al.Conjugated linoleic acid differentially modulates growth,tissue lipid deposi tion,and gene expression involved in the lipid metabo lism of grass carp[J].Aquaculture,2014,432(2):181-191
[19]Zou Q,Yang Y O,Wei B H,et al.Effects of dietary conjugated linoleic acid on growth performance,tissue adipocytokine levels,and lipid metabolism of grass carp[J].Aquaculture Nutrition,2018.DOI:10.1111/ANU.12,815
[20]Bradford,M.A rapid sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein dye binding[J].Analytical Biochemistry,1976,72(1-2):248-254
[21]National Research Council(NRC)[M].Nutrients Requirements of Fish and Shrimp.Washington DC:The National Academies Press.2011,109
[22]Du Z Y,Liu Y J,Tian L X,et al.Effect of dietary lipid level on growth,feed utilization and body composition by juvenile grass carp(Ctenopharyngodon idella)[J].Aquaculture Nutrition,2005,11(2):139-146
[23]Ding Z L,Kong Y Q,Zhang Y X,et al.Effects of dietary conjugated linoleic acid on growth,antioxidant capacity and lipid metabolism of Macrobrachium nipponense[J].Journal of Fisheries of China,2017,41(1):123-133[丁志丽,孔有琴,张易祥,等.共轭亚油酸对日本沼虾生长、抗氧化及脂质代谢的影响.水产学报,2017,41(1):123-133]
[24]Nogueira C W,Quinhones E B,Jung E A C,et al.Antiinflammatory and antinociceptive activity of diphenyl diselenide[J].Inflammation Research,2003,52(2):56-63
[25]Tao L,Fu S X.Progress on arachidonic acid and oxidative stress[J].Chinese Journal of Pathophysiology,2011,27(11):2233-2236[陶磊,傅淑霞.花生四烯酸与氧化应激的研究进展.中国病理生理杂志,2011,27(11):2233-2236]
[26]Rudneva I I.Blood antioxidant system of Black Sea elasmobranch and teleost[J].Comparative Biochemistry and Physiology,1997,118(2):255-260
[27]Martínez-álvarez R M,Morales A E,Sanz A.Antioxi dant defenses in fish:biotic and abiotic factors[J].Reviews in Fish Biology&Fisheries,2005,15(1-2):75-88
[28]Zhang K F,Zhang Z P,Chen Y,et al.Antioxidant defense system in animals[J].Chinese Journal of Zoology,2007,42(2):153-160[张克烽,张子平,陈芸,等.动物抗氧化系统中主要抗氧化酶基因的研究进展.动物学杂志,2007,42(2):153-160]
[29]Fridovich I.Superoxide Radical and Superoxide Dismutases[J].Autoxidation in Food and Biological Systems,1980,64(1):97-112
[30]Fang C H,Qiao K,Liu Z Y,et al.The research progress of antioxidant enzymes in marine organisms[J].Journal of Fisheries Research,2016,38(4):331-342[方春华,乔琨,刘智禹,等.海洋生物中抗氧化酶的研究进展.渔业研究,2016,38(4):331-342]
[31]Huang Y S,Lin Z D,Rong H,et al.Effects of conjugated linoleic acid on growth,body composition,antioxidant status,lipid metabolism and immunity parameters of juvenile Chu’s croaker,Nibea coibor[J].Aquaculture Research,2017,49(1):546-556
[32]Fracalossi D M,Lovell R T.Dietary lipid sources influence responses of channel catfish(Ictalurus punctatus)to challenge with the pathogen Edwardsiella ictaluri[J].Aquaculture,1994,119(2):287-298
[33]Cui W D,Leng X J,Li X Q,et al.Effects of astaxanthin and canthaxanthin on pigmentation of Muscle and total antioxidant capacity of liver in rainbow trout(Oncorhynchus mykiss)[J].Journal of Fisheries of China,2009,33(6):987-995[崔惟东,冷向军,李小勤,等.虾青素和角黄素对虹鳟肌肉着色和肝脏总抗氧化能力的影响.水产学报,2009,33(6):987-995]
[34]Funai K,Semenkovich C F.Skeletal muscle lipid flux:running water carries no poison[J].American Journal of Physiology Endocrinology&Metabolism,2011,301(2):E245-E251
[35]Dong X J.Comparison study on regulation ofΔ6 fatty acyl desaturase among rainbow trout,Japanese seabass and large yellow[D].Thesis for Doctor of Agriculture.Ocean University of China,Qingdao.2015[董小敬.虹鳟、花鲈和大黄鱼Δ6脂肪酸去饱和酶调控差异的研究.中国海洋大学博士论文,青岛.2015]
[36]Kennedy S R,Leaver M J,Campbell P J,et al.Influence of dietary oil content and conjugated linoleic acid(CLA)on lipid metabolism enzyme activities and gene expression in tissues of Atlantic salmon(Salmo salar L.)[J].Lipids,2006,41(5):423-436
[37]Takahashi Y,Kushiro M,Shinohara K,et al.Activity and mRNA levels of enzymes involved in hepatic fatty acid synthesis and oxidation in mice fed conjugated linoleic acid[J].Biochimica et Biophysica Acta,2003,1631(3):265-273
[38]Lass A,Zimmermann R,Oberer M,et al.Lipolysis-Ahighly regulated multi-enzyme complex mediates the catabolism of cellular fat stores[J].Progress in Lipid Research,2011,50(1):14-27
[39]Larosa P C,Miner J,Xia Y,et al.Trans-10,cis-12 conjugated linoleic acid causes inflammation and delipidation of white adipose tissue in mice:a microarray and histological analysis[J].Physiological Genomics,2006,27(3):282-294
[40]Pyper S R,Viswakarma N,Yu S,et al.PPARα:energy combustion,hypolipidemia,inflammation and cancer[J].Nuclear Receptor Signaling,2010,8:e002
[41]Rosa R,Andrade A M,Bandarra N M,et al.Physiological and biochemical effects of conjugated linoleic acid and its use in aquaculture[J].Reviews in Aquaculture,2010,2(2):59-72
[42]Peters J M,Park Y,Gonzalez F J,et al.Influence of conjugated linoleic acid on body composition and target gene expression in peroxisome proliferator-activated receptorα-null mice[J].Biochimica et Biophysica Acta,2001,1533(3):233-242
[43]Ide T.Interaction of fish oil and conjugated linoleic acid in affecting hepatic activity of lipogenic enzymes and gene expression in liver and adipose tissue[J].Diabetes,2005,54(2):412-423
[2]Wang K Y,Miao C H,Huang J L,et al.Changes in main biochemical indices and Acetyl-CoA Carboxylase 1mRNA expression of crass carp fed a high-fat diet[J].Chinese Journal of Animal Nutrition,2012,24(12):2375-2383[汪开毓,苗常鸿,黄锦炉,等.投喂高脂饲料后草鱼主要生化指标和乙酰辅酶A羧化酶1 mRNA表达的变化.动物营养学报,2012,24(12):2375-2383]
[3]Wu C Y.The pharmacodynamic function study of Sedum sarmentosum extract in grass carp fatty liver model[D].Thesis for Master of Veterinary.Sichuan Agricultural University,Chengdu.2013[吴春艳.垂盆草提取物对草鱼脂肪肝模型药效作用研究.四川农业大学硕士论文,成都.2013]
[4]Banni S.Conjugated linoleic acid metabolism[J].Current Opinion in Lipidology,2002,13(3):261-266
[5]Yang B,Chen H,Stanton C,et al.Review of the roles of conjugated linoleic acid in health and disease[J].Journal of Functional Foods,2015,15:314-325
[6]Twibell R G,Watkins B A,Rogers L,et al.Effects of dietary conjugated linoleic acids on hepatic and muscle lipids in hybrid striped bass[J].Lipids,2000,35(2):155-161
[7]Twibell R G,Watkins B A,Brown P B.Dietary conjugated linoleic acids and lipid source alter fatty acid composition of juvenile yellow perch,Perca flavescens[J].The Journal of Nutrition,2001,131(9):2322-2328
[8]Tan X Y,Luo Z,Xie P,et al.Effect of dietary conjugated linoleic acid(CLA)on growth performance,body composition and hepatic intermediary metabolism in juvenile yellow catfish Pelteobagrus fulvidraco[J].Aquaculture,2010,310(1-2):186-191
[9]Choi B D,Kang S J,Ha Y L,et al.Accumulation of Conjugated Linoleic Acid(CLA)in Tissues of Fish Fed Diets Containing Various Levels of CLA[M].Quality Attributes of Muscle Foods.1999,61-67
[10]Twibell R G,Wilson R P.Effects of dietary conjugated linoleic acids and total dietary lipid concentrations on growth responses of juvenile channel catfish,Ictalurus punctatus[J].Aquaculture,2003,221(1):621-628
[11]Figueiredo-Silva A C,Rema P,Bandarra N M,et al.Effects of dietary conjugated linoleic acid on growth,nutrient utilization,body composition,and hepatic lipogenesis in rainbow trout juveniles(Oncorhynchus mykiss)[J].Aquaculture,2005,248(1):163-172
[12]Makol A,Torrecillas S,Fernándezvaquero A,et al.Effect of conjugated linoleic acid on dietary lipids utilization,liver morphology and selected immune parameters in sea bass juveniles(Dicentrarchus labrax)[J].Comparative Biochemistry&Physiology,2009,154(2):179-187
[13]Makol A,Torrecillas S,Caballero M J,et al.Effect of long term feeding with conjugated linoleic acid(CLA)in growth performance and lipid metabolism of European sea bass(Dicentrarchus labrax)[J].Aquaculture,2012,368-369(1):129-137
[14]Zuo R T,Ai Q H,Mai K S,et al.Effects of conjugated linoleic acid on growth,non-specific immunity,antioxidant capacity,lipid deposition and related gene expression in juvenile large yellow croaker(Larmichthys crocea)fed soyabean oil-based diets[J].British Journal of Nutrition,2013,110(7):1220-1232
[15]Zhao Z Y.Effects of conjugated linoleic acid on lipid metabolism,immunity,flesh quality and PPAR gene expression in large Yellow croaker[D].Thesis for Doctor of Science.Zhejiang University,Hangzhou.2008[赵占宇.共轭亚油酸(CLA)对大黄鱼脂肪代谢、免疫、肉品质及PPAR基因表达的影响.浙江大学博士论文,杭州.2008]
[16]Kennedy S R,Campbell P J,Porter A,et al.Influence of dietary conjugated linoleic acid(CLA)on lipid and fatty acid composition in liver and flesh of Atlantic salmon(Salmo salar)[J].Comparative Biochemistry&Physiology,2005,141(2):168-178
[17]Leaver M J,Tocher D R,Obach A,et al.Effect of dietary conjugated linoleic acid(CLA)on lipid composition,metabolism and gene expression in Atlantic salmon(Salmo salar)tissues[J].Comparative Biochemistry and Physiology,2006,145(2):258-267
[18]Dong G F,Zou Q,Wang H,et al.Conjugated linoleic acid differentially modulates growth,tissue lipid deposi tion,and gene expression involved in the lipid metabo lism of grass carp[J].Aquaculture,2014,432(2):181-191
[19]Zou Q,Yang Y O,Wei B H,et al.Effects of dietary conjugated linoleic acid on growth performance,tissue adipocytokine levels,and lipid metabolism of grass carp[J].Aquaculture Nutrition,2018.DOI:10.1111/ANU.12,815
[20]Bradford,M.A rapid sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein dye binding[J].Analytical Biochemistry,1976,72(1-2):248-254
[21]National Research Council(NRC)[M].Nutrients Requirements of Fish and Shrimp.Washington DC:The National Academies Press.2011,109
[22]Du Z Y,Liu Y J,Tian L X,et al.Effect of dietary lipid level on growth,feed utilization and body composition by juvenile grass carp(Ctenopharyngodon idella)[J].Aquaculture Nutrition,2005,11(2):139-146
[23]Ding Z L,Kong Y Q,Zhang Y X,et al.Effects of dietary conjugated linoleic acid on growth,antioxidant capacity and lipid metabolism of Macrobrachium nipponense[J].Journal of Fisheries of China,2017,41(1):123-133[丁志丽,孔有琴,张易祥,等.共轭亚油酸对日本沼虾生长、抗氧化及脂质代谢的影响.水产学报,2017,41(1):123-133]
[24]Nogueira C W,Quinhones E B,Jung E A C,et al.Antiinflammatory and antinociceptive activity of diphenyl diselenide[J].Inflammation Research,2003,52(2):56-63
[25]Tao L,Fu S X.Progress on arachidonic acid and oxidative stress[J].Chinese Journal of Pathophysiology,2011,27(11):2233-2236[陶磊,傅淑霞.花生四烯酸与氧化应激的研究进展.中国病理生理杂志,2011,27(11):2233-2236]
[26]Rudneva I I.Blood antioxidant system of Black Sea elasmobranch and teleost[J].Comparative Biochemistry and Physiology,1997,118(2):255-260
[27]Martínez-álvarez R M,Morales A E,Sanz A.Antioxi dant defenses in fish:biotic and abiotic factors[J].Reviews in Fish Biology&Fisheries,2005,15(1-2):75-88
[28]Zhang K F,Zhang Z P,Chen Y,et al.Antioxidant defense system in animals[J].Chinese Journal of Zoology,2007,42(2):153-160[张克烽,张子平,陈芸,等.动物抗氧化系统中主要抗氧化酶基因的研究进展.动物学杂志,2007,42(2):153-160]
[29]Fridovich I.Superoxide Radical and Superoxide Dismutases[J].Autoxidation in Food and Biological Systems,1980,64(1):97-112
[30]Fang C H,Qiao K,Liu Z Y,et al.The research progress of antioxidant enzymes in marine organisms[J].Journal of Fisheries Research,2016,38(4):331-342[方春华,乔琨,刘智禹,等.海洋生物中抗氧化酶的研究进展.渔业研究,2016,38(4):331-342]
[31]Huang Y S,Lin Z D,Rong H,et al.Effects of conjugated linoleic acid on growth,body composition,antioxidant status,lipid metabolism and immunity parameters of juvenile Chu’s croaker,Nibea coibor[J].Aquaculture Research,2017,49(1):546-556
[32]Fracalossi D M,Lovell R T.Dietary lipid sources influence responses of channel catfish(Ictalurus punctatus)to challenge with the pathogen Edwardsiella ictaluri[J].Aquaculture,1994,119(2):287-298
[33]Cui W D,Leng X J,Li X Q,et al.Effects of astaxanthin and canthaxanthin on pigmentation of Muscle and total antioxidant capacity of liver in rainbow trout(Oncorhynchus mykiss)[J].Journal of Fisheries of China,2009,33(6):987-995[崔惟东,冷向军,李小勤,等.虾青素和角黄素对虹鳟肌肉着色和肝脏总抗氧化能力的影响.水产学报,2009,33(6):987-995]
[34]Funai K,Semenkovich C F.Skeletal muscle lipid flux:running water carries no poison[J].American Journal of Physiology Endocrinology&Metabolism,2011,301(2):E245-E251
[35]Dong X J.Comparison study on regulation ofΔ6 fatty acyl desaturase among rainbow trout,Japanese seabass and large yellow[D].Thesis for Doctor of Agriculture.Ocean University of China,Qingdao.2015[董小敬.虹鳟、花鲈和大黄鱼Δ6脂肪酸去饱和酶调控差异的研究.中国海洋大学博士论文,青岛.2015]
[36]Kennedy S R,Leaver M J,Campbell P J,et al.Influence of dietary oil content and conjugated linoleic acid(CLA)on lipid metabolism enzyme activities and gene expression in tissues of Atlantic salmon(Salmo salar L.)[J].Lipids,2006,41(5):423-436
[37]Takahashi Y,Kushiro M,Shinohara K,et al.Activity and mRNA levels of enzymes involved in hepatic fatty acid synthesis and oxidation in mice fed conjugated linoleic acid[J].Biochimica et Biophysica Acta,2003,1631(3):265-273
[38]Lass A,Zimmermann R,Oberer M,et al.Lipolysis-Ahighly regulated multi-enzyme complex mediates the catabolism of cellular fat stores[J].Progress in Lipid Research,2011,50(1):14-27
[39]Larosa P C,Miner J,Xia Y,et al.Trans-10,cis-12 conjugated linoleic acid causes inflammation and delipidation of white adipose tissue in mice:a microarray and histological analysis[J].Physiological Genomics,2006,27(3):282-294
[40]Pyper S R,Viswakarma N,Yu S,et al.PPARα:energy combustion,hypolipidemia,inflammation and cancer[J].Nuclear Receptor Signaling,2010,8:e002
[41]Rosa R,Andrade A M,Bandarra N M,et al.Physiological and biochemical effects of conjugated linoleic acid and its use in aquaculture[J].Reviews in Aquaculture,2010,2(2):59-72
[42]Peters J M,Park Y,Gonzalez F J,et al.Influence of conjugated linoleic acid on body composition and target gene expression in peroxisome proliferator-activated receptorα-null mice[J].Biochimica et Biophysica Acta,2001,1533(3):233-242
[43]Ide T.Interaction of fish oil and conjugated linoleic acid in affecting hepatic activity of lipogenic enzymes and gene expression in liver and adipose tissue[J].Diabetes,2005,54(2):412-423