猪肉中脂肪酸组成的品种差异及脂肪酸沉积的调控机制
|
摘要
猪肉是我国消费者最主要的动物蛋白质来源,其品质及安全与人类健康息息相关。猪肉中脂肪酸的含量与组成直接影响着肉的品质,如pH、嫩度、风味和多汁性等;同时,脂肪酸含量及组成与心血管疾病、肥胖和糖尿病等代谢性疾病的发生发展密切相关。因此,如何调控猪肉中脂肪酸含量及组成已成为畜牧业研究的热点之一。脂肪酸含量及组成受遗传、营养等多方面因素调控。本文在简述了脂肪酸与肉品质的关系及沉积机制的基础上,论述了猪肉中脂肪酸组成的品种差异及其营养调控途径和转基因调控技术等方面的相关研究进展,为调控猪肉中脂肪酸沉积进而生产功能性猪肉产品提供理论参考。
Pork is the main source of animal protein for Chinese consumers. The quality and the safety of pork are closely related to human health. The contents and composition of fatty acids in pork directly affect the meat quality,such as pH,tenderness,flavor and juiciness. M oreover,fatty acids are closely related to the occurrence and development of cardiovascular diseases,obesity and diabetes. Thus,howto regulate the fatty acid content and composition in pork has become one of the hotspots in animal husbandry research. The contents and composition of fatty acid in pork is regulated by heredity,nutrition and other factors. In this review,we briefly described the deposition mechanism of fatty acids and the relationship with meat quality. Furthermore,we discussed the breed differences,the nutritional regulation and transgenic regulation of fatty acid composition in pork. This reviewcould provide theoretical reference for regulating fatty acid deposition in pork and subsequently producing functional pork products. [Chinese Journal of Animal Nutrition,2019,31( 6) : 2507-2514]
Pork is the main source of animal protein for Chinese consumers. The quality and the safety of pork are closely related to human health. The contents and composition of fatty acids in pork directly affect the meat quality,such as pH,tenderness,flavor and juiciness. M oreover,fatty acids are closely related to the occurrence and development of cardiovascular diseases,obesity and diabetes. Thus,howto regulate the fatty acid content and composition in pork has become one of the hotspots in animal husbandry research. The contents and composition of fatty acid in pork is regulated by heredity,nutrition and other factors. In this review,we briefly described the deposition mechanism of fatty acids and the relationship with meat quality. Furthermore,we discussed the breed differences,the nutritional regulation and transgenic regulation of fatty acid composition in pork. This reviewcould provide theoretical reference for regulating fatty acid deposition in pork and subsequently producing functional pork products. [Chinese Journal of Animal Nutrition,2019,31( 6) : 2507-2514]
引文
[1]MARTINS J M,NEVER J A,FREITAS A,et al.Rearing system and oleic acid supplementation effect on carcass and lipid characteristics of tw o muscles from an obese pig breed[J].Animal,2015,9(10):1721-1730.
[2]孙建广,张石蕊.猪肉品质研究进展[J].中国猪业,2008,3(7):58-59.
[3]KOUBA M,BENATMANE F,BLOCHET J E,et al.Effect of a linseed diet on lipid oxidation,fatty acid composition of muscle,perirenal fat,and raw and cooked rabbit meat[J].M eat Science,2008,80(3):829-834.
[4]KHAN M I,JO C,TARIQ M R.Meat flavor precursors and factors influencing flavor precursors-A systematic review[J].M eat Science,2015,110:278-284.
[5]ZHANG J,WANG M,XIE J C,et al.Volatile flavor constituents in the pork broth of black-pig[J].Food Chemistry,2017,226:51-60.
[6]CAMERON N D,ENSER M,NUTE G R,et al.Genotype w ith nutrition interaction on fatty acid composition of intramuscular fat and the relationship w ith flavour of pig meat[J].M eat Science,2000,55(2):187-195.
[7]PEIRETTI P G,MEINERI G.Effects on growth performance,carcass characteristics,and the fat and meat fatty acid profile of rabbits fed diets w ith chia(Salvia hispanica L.)seed supplements[J].M eat Science,2008,80(4):1116-1121.
[8]CALDER P C.Functional roles of fatty acids and their effects on human health[J].Journal of Parenteral and Enteral Nutrition,2015,39(Suppl.1):18S-32S.
[9]ZHANG J,CHAI J,LUO Z G,et al.Meat and nutritional quality comparison of purebred and crossbred pigs[J].Animal Science Journal,2018,89(1):202-210.
[10]WATANABE Y,TATSUNO I.Omega-3 polyunsaturated fatty acids for cardiovascular diseases:present,past and future[J].Expert Review of Clinical Pharmacology,2017,10(8):865-873.
[11]SIMOPOULOS A P.An increase in the omega-6/omega-3 fatty acid ratio increases the risk for obesity[J].Nutrients,2016,8(3):128.
[12]WU J H,LEMAITRE R N,KING I B,et al.Circulating omega-6 polyunsaturated fatty acids and total and cause-specific mortality:the cardiovascular health study[J].Circulation,2014,130(15):1245-1253.
[13]MARRIOTT B P,YU K,MAJCHRZAK-HONG S,et al.Understanding diet and modeling changes in the omega-3 and omega-6 fatty acid composition of U.S.garrison foods for active duty personnel[J].M ilitary M edicine,2014,179(11):168-175.
[14]JONES R,ADEL-ALVAREZ L A,ALVAREZ O R,et al.Arachidonic acid and colorectal carcinogenesis[J].Molecular and Cellular Biochemistry,2003,253(1/2):141-149.
[15]HARRIS W S.International recommendations for consumption of long-chain omega-3 fatty acids[J].Journal of Cardiovascular M edicine,2007,8:S50-S52.
[16]张秀英,皇甫海燕,陈菁菁,等.高油酸油菜的研究进展及前景[J].作物研究,2007(增刊):654-656.
[17]WHELAN J,FRITSCHE K.Linoleic acid[J].Advances in Nutrition,2013,4(3):311-312.
[18]吴永保,李琳,闻治国,等.动物体内极长链多不饱和脂肪酸代谢及其生理功能[J].中国畜牧杂志,2018,54(3):20-26.
[19]SCHENK S,SABERI M,OLEFSKY J M.Insulin sensitivity:modulation by nutrients and inflammation[J].Journal of Clinical Investigation,2008,118(9):2992-3002.
[20]TAKEUCHI K,REUE K.Biochemistry,physiology,and genetics of GPAT,AGPAT,and lipin enzymes in triglyceride synthesis[J].American Journal of Physiology:Endocrinology and M etabolism,2009,296(6):E1195-E1209.
[21]LINDN D,WILLIAM-OLSSON L,RHEDIN M,et al.Overexpression of mitochondrial GPAT in rat hepatocytes leads to decreased fatty acid oxidation and increased glycerolipid biosynthesis[J].Journal of Lipid Research,2004,45(7):1279-1288.
[22]WENDEL A A,COOPER D E,ILKAYEVA O R,et al.Glycerol-3-phosphate acyltransferase(GPAT)-1,but not GPAT4,incorporates new ly synthesized fatty acids into triacylglycerol and diminishes fatty acid oxidation[J].Journal of Biological Chemistry,2013,288(38):27299-27306.
[23]BONEN A,CHABOWSKI A,LUIKEN J J,et al.M echanisms and regulation of protein-mediated cellular fatty acid uptake:molecular,biochemical,and physiological evidence[J].Physiology,2007,22(1):15-28.
[24]LUIKEN J J F P,BONEN A,GLATZ J F C.Cellular fatty acid uptake is acutely regulated by membrane-associated fatty acid-binding proteins[J].Prostaglandins,Leukotrienes and Essential Fatty Acids,2002,67(2):73-78.
[25]SCHWENK R W,HOLLOWAY G P,LUIKEN J J FP,et al.Fatty acid transport across the cell membrane:regulation by fatty acid transporters[J].Prostaglandins,Leukotrienes and Essential Fatty Acids,2010,82(4/5/6):149-154.
[26]STORCH J,THUMSER A E A.The fatty acid transport function of fatty acid-binding proteins[J].Biochimica et Biophysica Acta:M olecular and Cell Biology of Lipids,2000,1486(1):28-44.
[27]姜延志,李学伟,杨光希.猪L-FABP基因的克隆、表达特征及遗传多态性研究[J].遗传学报,2006,33(7):598-606.
[28]ABUMRAD N A,GOLDBERG I J.CD36 actions in the heart:lipids,calcium,inflammation,repair and more?[J].Biochimica et Biophysica Acta:M olecular and Cell Biology of Lipids,2016,1861(10):1442-1449.
[29]KIM T T,DYCK J R B.The role of CD36 in the regulation of myocardial lipid metabolism[J].Biochimica et Biophysica Acta:M olecular and Cell Biology of Lipids,2016,1861(10):1450-1460.
[30]GLATZ J F C,NABBEN M,HEATHER L C,et al.Regulation of the subcellular trafficking of CD36,a major determinant of cardiac fatty acid utilization[J].Biochimica et Biophysica Acta:M olecular and Cell Biology of Lipids,2016,1861(10):1461-1471.
[31]GLATZ J F C,LUIKEN J J F P.From fat to FAT(CD36/SR-B2):understanding the regulation of cellular fatty acid uptake[J].Biochimie,2016,136:21-26.
[32]COBURN C T,HAJRI T,IBRAHIMI A,et al.Role of CD36 in membrane transport and utilization of longchain fatty acids by different tissues[J].Journal of M olecular Neuroscience,2001,16(2/3):117-121.
[33]BLACK P N,AHOWESSO C,MONTEFUSCO D,et al.Fatty acid transport proteins:targeting FATP2 as a gatekeeper involved in the transport of exogenous fatty acids[J].M edicinal Chemistry Communications,2016,7(4):612-622.
[34]YANG Y,FENG Y Y,ZHANG X W,et al.Activation of PPARαby fatty acid accumulation enhances fatty acid degradation and sulfatide synthesis[J].Tohoku Journal of Experimental M edicine,2015,240(2):113-122.
[35]NIELSEN R,PEDERSEN T,HAGENBEEK D,et al.Genome-w ide profiling of PPARγ:RXR and RNApolymeraseⅡoccupancy reveals temporal activation of distinct metabolic pathw ays and changes in RXRdimer composition during adipogenesis[J].Genes&Development,2008,22(21):2953-2967.
[36]LEFTEROVA M I,ZHANG Y,STEGER D J,et al.PPARγand C/EBP factors orchestrate adipocyte biology via adjacent binding on a genome-w ide scale[J].Genes&Development,2008,22(21):2941-2952.
[37]MIKKELSEN T S,XU Z,ZHANG X L,et al.Comparative epigenomic analysis of murine and human adipogenesis[J].Cell,2010,143(1):156-169.
[38]POULSEN L L C,SIERSBK M,MANDRUP S.PPARs:fatty acid sensors controlling metabolism[J].Seminars in Cell&Developmental Biology,2012,23(6):631-639.
[39]VITERBO V S,LOPEZ B I M,KANG H,et al.Genome w ide association study of fatty acid composition in Duroc sw ine[J].Asian-Australasian Journal of Animal Sciences,2018,31(8):1127-1133.
[40]ZHANG J,ZHANG Y,GONG H,et al.Genetic mapping using 1.4M SNP array refined loci for fatty acid composition traits in Chinese Erhualian and Bamaxiang pigs[J].Journal of Animal Breeding and Genetics,2017,134(6):472-483.
[41]SIMOPOULOS A P,CLELAND L G,CHRISTIANS-EN E N.Omega-6/omega-3 essential fatty acid ratio:the scientific evidence[J].Journal of Human Nutrition and Dietetics,2010,17(2):165-166.
[42]SPYCHALLA J P,KINNEY A J,BROWSE J.Identification of an animalω-3 fatty acid desaturase by heterologous expression in Arabidopsis[J].Proceedings of the National Academy of Sciences of the United States of America,1997,94(4):1142-1147.
[43]LI M J,OUYANG H S,YUAN H M,et al.Site-specific fat-1 knock-in enables significant decrease of n-6PUFAs/n-3PUFAs ratio in pigs[J].G3:Genes,Genomes,Genetics,2018,8(5):1747-1754.
[44]KANG J X,WANG J D,WU L,et al.Fat-1 mice convert n-6 to n-3 fatty acids[J].Nature,2004,427(6974):504.
[45]LAI L X,KANG J X,LI R F,et al.Generation of cloned transgenic pigs rich in omega-3 fatty acids[J].Nature Biotechnology,2006,24(4):435-436.
[46]宋倩倩,张金枝,刘健,等.不同品种猪肉质性状和脂肪酸含量的研究[J].家畜生态学报,2018,39(2):24-28.
[47]吴妹英,肖天放,张力.莆田黑猪及其杂种肌肉脂肪酸组成与含量的研究[J].福建畜牧兽医,2007(增刊):16-18.
[48]鲁云风,张晓娜,张征田.南阳黑猪和大白猪脂肪酸分析及其综合评价[J].中国畜牧兽医,2017,44(4):1032-1036.
[49]章杰,罗宗刚,陈磊,等.荣昌猪和杜洛克猪肉质及营养价值的比较分析[J].食品科学,2015,36(24):127-130.
[50]于永生,张志彬,罗晓彤,等.三个猪种肌肉脂肪酸含量分析[J].黑龙江畜牧兽医,2016(1):115-117.
[51]BOTHMA C,HUGO A,OSTHOFF G,et al.Effect of dietary conjugated linoleic acid supplementation on the technological quality of backfat of pigs[J].M eat Science,2014,97(2):277-286.
[52]SUN D Y,ZHU X P,QIAO S Y,et al.Effects of conjugated linoleic acid levels and feeding intervals on performance,carcass traits and fatty acid composition of finishing barrow s[J].Archives of Animal Nutrition,2004,58(4):277-286.
[53]WEBER T E,RICHERT B T,BELURY M A,et al.E-valuation of the effects of dietary fat,conjugated linoleic acid,and ractopamine on growth performance,pork quality,and fatty acid profiles in genetically lean gilts[J].Journal of Animal Science,2006,84(3):720-732.
[54]敖力格日玛,吉木斯.富含α-亚麻酸的添加剂对生长育肥猪肌肉及脂肪组织脂肪酸组成的影响[C]//中国畜牧兽医学会2006学术年会论文集(上册).北京:中国畜牧兽医学会,2006.
[55]KOUBA M,ENSER M,WHITTINGTON F M,et al.Effect of a high-linolenic acid diet on lipogenic enzyme activities,fatty acid composition,and meat quality in the grow ing pig[J].Journal of Animal Science,2003,81(8):1967-1979.
[56]SMINK W,VERSTEGEN M W A,GERRITS W J J.Effect of intake of linoleic acid andα-linolenic acid levels on conversion into long-chain polyunsaturated fatty acids in backfat and in intramuscular fat of growing pigs[J].Journal of Animal Physiology and Animal Nutrition,2013,97(3):558-565.
[57]MADEIRA M S,ALFAIA C M,COSTA P,et al.Effect of betaine and arginine in lysine-deficient diets on grow th,carcass traits,and pork quality[J].Journal of Animal Science,2015,93(10):4721-4733.
[58]MARTINS J M,ALBUQUERQUE A,NEVES J A,et al.Influence of outdoor rearing and oleic acid supplementation on lipid characteristics of muscle and adipose tissues from obese Alentejano pigs[J].Journal of Animal Physiology and Animal Nutrition,2018,102(2):e578-e590.
[59]陈国顺,高海霞.中草药饲料添加剂对香猪生长性能和肉质的影响[J].畜牧与兽医,2012,44(1):23-27.
[60]胡广英,曹暄雅,曹日亮,等.中草药添加剂对不同猪种肉质中脂肪酸和胆固醇含量的效应[J].养猪,2018(1):65-68.
[61]朱晓艳,吕先召,邱晓东,等.苜蓿草粉对育肥猪肉品质、肌肉氨基酸和脂肪酸含量的影响[J].动物营养学报,2018,30(9):3473-3482.
[62]WANG L S,SHI Z,GAO R,et al.Effects of conjugated linoleic acid or betaine on the grow th performance and fatty acid composition in backfat and belly fat of finishing pigs fed dried distillers grains w ith solubles[J].Animal,2015,9(4):569-575.
[63]MADEIRA M S,ROLO E A,ALFAIA C M,et al.Influence of betaine and arginine supplementation of reduced protein diets on fatty acid composition and gene expression in the muscle and subcutaneous adipose tissue of cross-bred pigs[J].The British Journal of Nutrition,2016,115(6):937-950.
[64]HU C J,JIANG Q Y,ZHANG T,et al.Dietary supplementation w ith arginine and glutamic acid modifies grow th performance,carcass traits,and meat quality in grow ing-finishing pigs[J].Journal of Animal Science,2017,95(6):2680-2689.
[65]刘瑞生,徐建峰,王珂,等.中草药添加剂对羊肉营养成分、脂肪酸含量和重金属残留的影响[J].畜牧兽医杂志,2017,36(6):14-17.
[66]SUN Y T,LUO J,ZHU J J,et al.Effect of short-chain fatty acids on triacylglycerol accumulation,lipid droplet formation and lipogenic gene expression in goat mammary epithelial cells[J].Animal Science Journal,2016,87(2):242-249.
[2]孙建广,张石蕊.猪肉品质研究进展[J].中国猪业,2008,3(7):58-59.
[3]KOUBA M,BENATMANE F,BLOCHET J E,et al.Effect of a linseed diet on lipid oxidation,fatty acid composition of muscle,perirenal fat,and raw and cooked rabbit meat[J].M eat Science,2008,80(3):829-834.
[4]KHAN M I,JO C,TARIQ M R.Meat flavor precursors and factors influencing flavor precursors-A systematic review[J].M eat Science,2015,110:278-284.
[5]ZHANG J,WANG M,XIE J C,et al.Volatile flavor constituents in the pork broth of black-pig[J].Food Chemistry,2017,226:51-60.
[6]CAMERON N D,ENSER M,NUTE G R,et al.Genotype w ith nutrition interaction on fatty acid composition of intramuscular fat and the relationship w ith flavour of pig meat[J].M eat Science,2000,55(2):187-195.
[7]PEIRETTI P G,MEINERI G.Effects on growth performance,carcass characteristics,and the fat and meat fatty acid profile of rabbits fed diets w ith chia(Salvia hispanica L.)seed supplements[J].M eat Science,2008,80(4):1116-1121.
[8]CALDER P C.Functional roles of fatty acids and their effects on human health[J].Journal of Parenteral and Enteral Nutrition,2015,39(Suppl.1):18S-32S.
[9]ZHANG J,CHAI J,LUO Z G,et al.Meat and nutritional quality comparison of purebred and crossbred pigs[J].Animal Science Journal,2018,89(1):202-210.
[10]WATANABE Y,TATSUNO I.Omega-3 polyunsaturated fatty acids for cardiovascular diseases:present,past and future[J].Expert Review of Clinical Pharmacology,2017,10(8):865-873.
[11]SIMOPOULOS A P.An increase in the omega-6/omega-3 fatty acid ratio increases the risk for obesity[J].Nutrients,2016,8(3):128.
[12]WU J H,LEMAITRE R N,KING I B,et al.Circulating omega-6 polyunsaturated fatty acids and total and cause-specific mortality:the cardiovascular health study[J].Circulation,2014,130(15):1245-1253.
[13]MARRIOTT B P,YU K,MAJCHRZAK-HONG S,et al.Understanding diet and modeling changes in the omega-3 and omega-6 fatty acid composition of U.S.garrison foods for active duty personnel[J].M ilitary M edicine,2014,179(11):168-175.
[14]JONES R,ADEL-ALVAREZ L A,ALVAREZ O R,et al.Arachidonic acid and colorectal carcinogenesis[J].Molecular and Cellular Biochemistry,2003,253(1/2):141-149.
[15]HARRIS W S.International recommendations for consumption of long-chain omega-3 fatty acids[J].Journal of Cardiovascular M edicine,2007,8:S50-S52.
[16]张秀英,皇甫海燕,陈菁菁,等.高油酸油菜的研究进展及前景[J].作物研究,2007(增刊):654-656.
[17]WHELAN J,FRITSCHE K.Linoleic acid[J].Advances in Nutrition,2013,4(3):311-312.
[18]吴永保,李琳,闻治国,等.动物体内极长链多不饱和脂肪酸代谢及其生理功能[J].中国畜牧杂志,2018,54(3):20-26.
[19]SCHENK S,SABERI M,OLEFSKY J M.Insulin sensitivity:modulation by nutrients and inflammation[J].Journal of Clinical Investigation,2008,118(9):2992-3002.
[20]TAKEUCHI K,REUE K.Biochemistry,physiology,and genetics of GPAT,AGPAT,and lipin enzymes in triglyceride synthesis[J].American Journal of Physiology:Endocrinology and M etabolism,2009,296(6):E1195-E1209.
[21]LINDN D,WILLIAM-OLSSON L,RHEDIN M,et al.Overexpression of mitochondrial GPAT in rat hepatocytes leads to decreased fatty acid oxidation and increased glycerolipid biosynthesis[J].Journal of Lipid Research,2004,45(7):1279-1288.
[22]WENDEL A A,COOPER D E,ILKAYEVA O R,et al.Glycerol-3-phosphate acyltransferase(GPAT)-1,but not GPAT4,incorporates new ly synthesized fatty acids into triacylglycerol and diminishes fatty acid oxidation[J].Journal of Biological Chemistry,2013,288(38):27299-27306.
[23]BONEN A,CHABOWSKI A,LUIKEN J J,et al.M echanisms and regulation of protein-mediated cellular fatty acid uptake:molecular,biochemical,and physiological evidence[J].Physiology,2007,22(1):15-28.
[24]LUIKEN J J F P,BONEN A,GLATZ J F C.Cellular fatty acid uptake is acutely regulated by membrane-associated fatty acid-binding proteins[J].Prostaglandins,Leukotrienes and Essential Fatty Acids,2002,67(2):73-78.
[25]SCHWENK R W,HOLLOWAY G P,LUIKEN J J FP,et al.Fatty acid transport across the cell membrane:regulation by fatty acid transporters[J].Prostaglandins,Leukotrienes and Essential Fatty Acids,2010,82(4/5/6):149-154.
[26]STORCH J,THUMSER A E A.The fatty acid transport function of fatty acid-binding proteins[J].Biochimica et Biophysica Acta:M olecular and Cell Biology of Lipids,2000,1486(1):28-44.
[27]姜延志,李学伟,杨光希.猪L-FABP基因的克隆、表达特征及遗传多态性研究[J].遗传学报,2006,33(7):598-606.
[28]ABUMRAD N A,GOLDBERG I J.CD36 actions in the heart:lipids,calcium,inflammation,repair and more?[J].Biochimica et Biophysica Acta:M olecular and Cell Biology of Lipids,2016,1861(10):1442-1449.
[29]KIM T T,DYCK J R B.The role of CD36 in the regulation of myocardial lipid metabolism[J].Biochimica et Biophysica Acta:M olecular and Cell Biology of Lipids,2016,1861(10):1450-1460.
[30]GLATZ J F C,NABBEN M,HEATHER L C,et al.Regulation of the subcellular trafficking of CD36,a major determinant of cardiac fatty acid utilization[J].Biochimica et Biophysica Acta:M olecular and Cell Biology of Lipids,2016,1861(10):1461-1471.
[31]GLATZ J F C,LUIKEN J J F P.From fat to FAT(CD36/SR-B2):understanding the regulation of cellular fatty acid uptake[J].Biochimie,2016,136:21-26.
[32]COBURN C T,HAJRI T,IBRAHIMI A,et al.Role of CD36 in membrane transport and utilization of longchain fatty acids by different tissues[J].Journal of M olecular Neuroscience,2001,16(2/3):117-121.
[33]BLACK P N,AHOWESSO C,MONTEFUSCO D,et al.Fatty acid transport proteins:targeting FATP2 as a gatekeeper involved in the transport of exogenous fatty acids[J].M edicinal Chemistry Communications,2016,7(4):612-622.
[34]YANG Y,FENG Y Y,ZHANG X W,et al.Activation of PPARαby fatty acid accumulation enhances fatty acid degradation and sulfatide synthesis[J].Tohoku Journal of Experimental M edicine,2015,240(2):113-122.
[35]NIELSEN R,PEDERSEN T,HAGENBEEK D,et al.Genome-w ide profiling of PPARγ:RXR and RNApolymeraseⅡoccupancy reveals temporal activation of distinct metabolic pathw ays and changes in RXRdimer composition during adipogenesis[J].Genes&Development,2008,22(21):2953-2967.
[36]LEFTEROVA M I,ZHANG Y,STEGER D J,et al.PPARγand C/EBP factors orchestrate adipocyte biology via adjacent binding on a genome-w ide scale[J].Genes&Development,2008,22(21):2941-2952.
[37]MIKKELSEN T S,XU Z,ZHANG X L,et al.Comparative epigenomic analysis of murine and human adipogenesis[J].Cell,2010,143(1):156-169.
[38]POULSEN L L C,SIERSBK M,MANDRUP S.PPARs:fatty acid sensors controlling metabolism[J].Seminars in Cell&Developmental Biology,2012,23(6):631-639.
[39]VITERBO V S,LOPEZ B I M,KANG H,et al.Genome w ide association study of fatty acid composition in Duroc sw ine[J].Asian-Australasian Journal of Animal Sciences,2018,31(8):1127-1133.
[40]ZHANG J,ZHANG Y,GONG H,et al.Genetic mapping using 1.4M SNP array refined loci for fatty acid composition traits in Chinese Erhualian and Bamaxiang pigs[J].Journal of Animal Breeding and Genetics,2017,134(6):472-483.
[41]SIMOPOULOS A P,CLELAND L G,CHRISTIANS-EN E N.Omega-6/omega-3 essential fatty acid ratio:the scientific evidence[J].Journal of Human Nutrition and Dietetics,2010,17(2):165-166.
[42]SPYCHALLA J P,KINNEY A J,BROWSE J.Identification of an animalω-3 fatty acid desaturase by heterologous expression in Arabidopsis[J].Proceedings of the National Academy of Sciences of the United States of America,1997,94(4):1142-1147.
[43]LI M J,OUYANG H S,YUAN H M,et al.Site-specific fat-1 knock-in enables significant decrease of n-6PUFAs/n-3PUFAs ratio in pigs[J].G3:Genes,Genomes,Genetics,2018,8(5):1747-1754.
[44]KANG J X,WANG J D,WU L,et al.Fat-1 mice convert n-6 to n-3 fatty acids[J].Nature,2004,427(6974):504.
[45]LAI L X,KANG J X,LI R F,et al.Generation of cloned transgenic pigs rich in omega-3 fatty acids[J].Nature Biotechnology,2006,24(4):435-436.
[46]宋倩倩,张金枝,刘健,等.不同品种猪肉质性状和脂肪酸含量的研究[J].家畜生态学报,2018,39(2):24-28.
[47]吴妹英,肖天放,张力.莆田黑猪及其杂种肌肉脂肪酸组成与含量的研究[J].福建畜牧兽医,2007(增刊):16-18.
[48]鲁云风,张晓娜,张征田.南阳黑猪和大白猪脂肪酸分析及其综合评价[J].中国畜牧兽医,2017,44(4):1032-1036.
[49]章杰,罗宗刚,陈磊,等.荣昌猪和杜洛克猪肉质及营养价值的比较分析[J].食品科学,2015,36(24):127-130.
[50]于永生,张志彬,罗晓彤,等.三个猪种肌肉脂肪酸含量分析[J].黑龙江畜牧兽医,2016(1):115-117.
[51]BOTHMA C,HUGO A,OSTHOFF G,et al.Effect of dietary conjugated linoleic acid supplementation on the technological quality of backfat of pigs[J].M eat Science,2014,97(2):277-286.
[52]SUN D Y,ZHU X P,QIAO S Y,et al.Effects of conjugated linoleic acid levels and feeding intervals on performance,carcass traits and fatty acid composition of finishing barrow s[J].Archives of Animal Nutrition,2004,58(4):277-286.
[53]WEBER T E,RICHERT B T,BELURY M A,et al.E-valuation of the effects of dietary fat,conjugated linoleic acid,and ractopamine on growth performance,pork quality,and fatty acid profiles in genetically lean gilts[J].Journal of Animal Science,2006,84(3):720-732.
[54]敖力格日玛,吉木斯.富含α-亚麻酸的添加剂对生长育肥猪肌肉及脂肪组织脂肪酸组成的影响[C]//中国畜牧兽医学会2006学术年会论文集(上册).北京:中国畜牧兽医学会,2006.
[55]KOUBA M,ENSER M,WHITTINGTON F M,et al.Effect of a high-linolenic acid diet on lipogenic enzyme activities,fatty acid composition,and meat quality in the grow ing pig[J].Journal of Animal Science,2003,81(8):1967-1979.
[56]SMINK W,VERSTEGEN M W A,GERRITS W J J.Effect of intake of linoleic acid andα-linolenic acid levels on conversion into long-chain polyunsaturated fatty acids in backfat and in intramuscular fat of growing pigs[J].Journal of Animal Physiology and Animal Nutrition,2013,97(3):558-565.
[57]MADEIRA M S,ALFAIA C M,COSTA P,et al.Effect of betaine and arginine in lysine-deficient diets on grow th,carcass traits,and pork quality[J].Journal of Animal Science,2015,93(10):4721-4733.
[58]MARTINS J M,ALBUQUERQUE A,NEVES J A,et al.Influence of outdoor rearing and oleic acid supplementation on lipid characteristics of muscle and adipose tissues from obese Alentejano pigs[J].Journal of Animal Physiology and Animal Nutrition,2018,102(2):e578-e590.
[59]陈国顺,高海霞.中草药饲料添加剂对香猪生长性能和肉质的影响[J].畜牧与兽医,2012,44(1):23-27.
[60]胡广英,曹暄雅,曹日亮,等.中草药添加剂对不同猪种肉质中脂肪酸和胆固醇含量的效应[J].养猪,2018(1):65-68.
[61]朱晓艳,吕先召,邱晓东,等.苜蓿草粉对育肥猪肉品质、肌肉氨基酸和脂肪酸含量的影响[J].动物营养学报,2018,30(9):3473-3482.
[62]WANG L S,SHI Z,GAO R,et al.Effects of conjugated linoleic acid or betaine on the grow th performance and fatty acid composition in backfat and belly fat of finishing pigs fed dried distillers grains w ith solubles[J].Animal,2015,9(4):569-575.
[63]MADEIRA M S,ROLO E A,ALFAIA C M,et al.Influence of betaine and arginine supplementation of reduced protein diets on fatty acid composition and gene expression in the muscle and subcutaneous adipose tissue of cross-bred pigs[J].The British Journal of Nutrition,2016,115(6):937-950.
[64]HU C J,JIANG Q Y,ZHANG T,et al.Dietary supplementation w ith arginine and glutamic acid modifies grow th performance,carcass traits,and meat quality in grow ing-finishing pigs[J].Journal of Animal Science,2017,95(6):2680-2689.
[65]刘瑞生,徐建峰,王珂,等.中草药添加剂对羊肉营养成分、脂肪酸含量和重金属残留的影响[J].畜牧兽医杂志,2017,36(6):14-17.
[66]SUN Y T,LUO J,ZHU J J,et al.Effect of short-chain fatty acids on triacylglycerol accumulation,lipid droplet formation and lipogenic gene expression in goat mammary epithelial cells[J].Animal Science Journal,2016,87(2):242-249.