维生素A对动物脂类代谢的调节作用与机制
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摘要
维生素A是影响动物组织脂类代谢和脂肪生成的关键因子。本文综述了维生素A对动物脂肪生成和脂类代谢的影响,并从脂类代谢相关基因的表达及其信号通路、脂肪细胞的数量与脂肪细胞因子分泌的角度综述了其可能的影响机制,为深入探究维生素A调控动物的脂肪代谢提供了理论依据。
引文
[1]Puigserver P,Vazquez F,Bonet M L,et al..In vitro and in vivo induction of brown adipocyte uncoupling protein(thermogenin)by retinoic acid[J].Biochemical Journal,1996,317:827-833.
[2]Strom K,Gundersen T E.Hansson 0,et al..Hormone-sensitive lipase(HSL)is also a retinyl ester hydrolase.-evidence from mice lacking HSL[J].The FASEB Journal.Research Communication,2009,23:2307-2316.
[3]Jeyakumar S M,Sheril A,Vajreswari A.Chronic vitamin A-enriched diet feeding induces body weight gain and adiposity in lean and glucose-intolerant obese rats of WNIN/GR-Ob strain[J].Experimental Physiology.2015,100(11):1352-1361.
[4]Ribot J,Felipe F,Bonet M L,et al..Changes of adiposity in response to vitamin A status correlate with changes of PPAR gamma 2 expression[J].Obesity Research.2001,9(8),500-509.
[5]Goodwin K,Abrahamowicz M,Leonard G,et al..Dietary Vitamin A and Visceral Adiposity:A Modulating Role of the Retinol-Binding Protein 4 Gene[J].Journal of Journal of Nutrigenetics and Nutrigenomics.2015,8(4-6):164-173.
[6]Krone K G,Ward A K,Madder KM,et al..Interaction of vitamin A supplementation level with ADH1C genotype on intramuscular fat in beef steers.[J].Animal,2016,10(3):403-409.
[7]Felipe F,Bonet M L,Ribot J,et al..Modulation of resistin expression by retinoic acid and vitamin A status[J].Diabetes,2004,53(4):882-889.
[8]Mercaer J,Ribot J,Murano I,et al..Remodeling of white adipose tissue after retinoic acid administration in mice[J].Endocrinology,2006,147(11):5325-5332.
[9]Amengual J,Ribot J,Bonet M L,et al..Retinoic acid treatment enhances lipid oxidation and inhibits lipid biosynthesis capacities in the liver of mice[J].Cellular Physiology and Biochemistry,2010,25:657-666.
[10]Berry D C,Noy N.All-trans-retinoic acid represses obesity and insulin resistance by activating both peroxisome proliferation-activated receptor beta/delta and retinoic acid receptor[J].Molecular and Cellular Biology,2009,29(12):3286-3296.
[11]Ayuso M,Fernandez A,Isabel B,et al..Long term vitamin A restriction improves meat quality parameters and modifies gene expression in Iberian pigs[J].Journal of Animal Science,2015,93(6):2730-2744.
[12]Silva L D S V D.Veiga G V D.Ramalho R A.Association of serum concentrations of retinol and carotenoids with overweight in children and adolescents[J].Nutrition,2007,23(5):392-397.
[13]Villaca Chaves G,Pereira S E,Saboya C J,et al..Non-alcoholic fatty liver disease and its relationship with the nutritional status of vitamin A in individuals with classⅢobesity[J].Obesity Surgery,2008,18(4):378-385.
[14]Burrows T L,Warren J M,Colyvas K,et al..Validation of overweight children's fruit and vegetable intake using plasma carotenoids[J].Obesity,2009,17:162-168.
[15]Strauss R S.Comparison of serum concentrations of alpha-tocopherol and beta-carotene in a cross-sectional sample of obese and nonobese children(NHANES III)[J].Journal of Pediatrics,1999,134(2):160-165.
[16]Facchini F,Coulston A M,Reaven G M.Relation between dietary vitamin intake and resistance to insulin-mediated glucose disposal in healthy volunteers[J].American Journal of Clinical Nutrition,1996,63(6):946-949.
[17]Mercader J.Granados N,Bonet M L,et al..All-trans retinoic acid decreases murine adipose retinol binding protein 4production[J].Cellular Physiology and Biochemistry,2008,22(4)363-372.
[18]Tourniaire F.Musinovic H.Gouranton E,et al..All-trans retinoic acid induces oxidative phosphorylation and mitochondria biogenesis in adipocytes[J],Journal of Lipid Research,2015,56(6):1100-1109.
[19]Murano I,Morroni M,Zingaretti M C,et al..Morphology of ferret subcutaneous adipose tissue after 6-monthdaily supplementation with oral beta-carotene[J].Biochimica Et Biophysica Acta,2015,1740(2):305-312.
[20]Sanchez J.Fuster A,Oliver P,et al..Effects of beta-carotene supplementation on adipose tissue thermogenic capacity in ferrets(Mustela putorius furo)[J].British Journal of Nutrition,2009,102(11):1686-1694.
[21]Kumar M V,Sunvold G D,Scarpace P J.Dietary vitamin A supplementation in rats suppression of leptin and induction of UCP1 mRNA[J].Journal of Lipid Research,1999,40(5)824-829.
[22]Jeyakumar S M.Vajreswari A.Giridharan N V.Chronic dietary vitamin A supplementation regulates obesity in an obese mutant WNIN/Ob rat model[J].Obesity,2006,14(1):52-59.
[23]Yehya A.Baer J T,Smiley W,et al..Hypervitaminosis A altering the lipid profile in a hypercholesterolemic patient[J].Journal of Clinical Lipidology,2009,3(3):205-207.
[24]Khanna A.Reddy T S.Effect of undernutrition and vitamin A deficiency on the phospholipid composition of rat tissues at 21 days of age-I.Liver,spleen and kidney[J].International Journal for Vitamin and Nutrition Research,1983,53(1):3-8.
[25]Wiss O,Wiss V.Alterations of the lipid metabolism of rat liver as early symptoms of vitamin A deficiency[J].International Journal for Vitamin and Nutrition Research,1980,50(3):233-237.
[26]Tripathy S,Chapman J D,Han C Y,Hogarth C A,et al..All-Trans-Retinoic Acid Enhances Mitochondrial Function in Models of Human Liver[J].Molecular Pharmacology,2016,89(5):560-574.
[27]Mcclinintick J N,Crabb D W,Tian H,et al..Global effects of vitamin A deficiency on gene expression in rat liver:evidence for hypoandrogenism[J].Journal of Nutritional Biochemistry,2006,17(5):345-355.
[28]Oliveros L B,Domeniconi M A,Vega V A,et al..Vitamin A deficiency modifies lipid metabolism in rat liver[J].British Journal of Nutrition,2007,97(2):263-272.
[29]Serra F,Bonet M L.Puigserver P,et al..Stimulation of uncoupling protein 1 expression in brown adipocytes by naturally occurring carotenoids[J].International Journal of Obesity,1999,23(6):650-655.
[30]Felipe F,Bonet M L,Ribot J,et al..Up-regulation of muscle uncoupling protein 3 gene expression in mice following high fat diet,dietary vitamin A supplementation and acute retinoic acid-treatment[J].International Journal of Obesity,2003,27(1):60-69.
[31]Mercader J.Madsen L,Felipe F,et al..All-trans retinoic acid increases oxidative metabolism in mature adipocytes[J].Cellular Physiology and Biochemistry,2007,20(6):1061-1072.
[32]Repa J J,Liang G,Ou J,et al..Regulation of mouse sterol regulatory element-binding protein-lc gene(SREBP-lc)by oxysterol receptors,LXRalpha and LXRbeta[J].Genes and Development,2000,14(22).-2819-2830.
[33]Joseph S B,Laffitte B A.Patel P H,et al..Direct and indirect mechanisms for regulation of fatty acid synthase gene expression by liver X receptors[J].Journal of Biological Chemistry,2002,277(13):11019-11025.
[34]Feige J N,German L.Michalik B,et al..From molecular action to physiological outputs:peroxisome proliferator-activated receptors are nuclear receptors at the crossroads of key cellular functions[J].Progress in Lipid Research,2006,45(2):120-159.
[35]Lefterova M I,Lazar M A.New developments in adipogenesis[J].Medical Clinics of North America,2009,89(6):107-114.
[36]Mandard S,Muller M,Kersten S.Peroxisome proliferator-activated receptor alpha target genes[J].Ppar Research,2010,4:393-416.
[37]Ide T.Shimano H,Yoshikawa T,et al..Cross-talk between peroxisome proliferator-activated receptor(PPAR)alpha and liver X receptor(LXR)in nutritional regulation of fatty acid metabolism.TJ.LXRs suppress lipid degradation gene promoters through inhibition of PPAR signaling[J].Molecular Endocrinology,2003,17(7):1255-1267.
[38]Yoshikawa T,Ide T,Shimano H,et al..Cross-talk between peroxisome proliferator-activated receptor(PPAR)alpha and liver X receptor(LXR)in nutritional regulation of fatty acid metabolism.I.PPARs suppress sterol regulatory element binding protein-lc promoter through inhibition of LXR signaling[J].Molecular Endocrinology,2003,17(7):1240-1254.
[39]Blomhoff R.Blomhoff H K.Overview of retinoid metabolism and function[J].Journal of Neurobiology,2006,66(7):606-630.
[40]Cadoudal T,Glorian M,Massias A,et al..Retinoids upregulate phosphoenolpyruvate carboxykinase and glyceroneogenesis in human and rodent adipocytes[J].Journal of Nutrition,2008,138(6):1004-1009.
[41]Miller C W,Waters K M,Ntambi J M.Regulation of hepatic stearoyl-CoA desaturase gene 1 by vitamin A[J].Biochemical and Biophysical Research Communications,1997,231(1):206-210.
[42]Solanes G,Pedraza N,Iglesias R,et al..The human uncoupling protein-3 gene promoter requires MyoD and is induced by retinoic acid in muscle cells[J].Faseb Journal Official Publication of the Federation of American Societies for Experimental Biology,2000,14(14),2141-2143.
[43]Raisher B D.Gulick T,Zhang Z,et al..Identification of a novel retinoid-responsive element in the promoter region of the medium chain acyl-coenzyme A dehydrogenase gene[J].Journal of Biological Chemistry,1992,267(28):20264-20269.
[44]Yoshikawa T,Shimano H,Amemiya-Kudo M,et al..Identification of liver X receptor-retinoid X receptor as an activator of the sterol regulatory element-binding protein lc gene promoter[J].Molecular and Cellular Biology,2001,21(9):2991-3000.
[45]Taniguchi D,Mizzoguchi Y.Retinoic acids change gene expression profiles of bovine intramuscular adipocyte differentiation,based on microarray analysis.Animal Science Journal.2015,86(6):579-587.
[46]Shaw N,Elholm M,Noy N,et al..Retinoic acid is a high affinity selective ligand for the peroxisome proliferator-activated receptor beta/delta[J].Journal of Biological Chemistry,2003,278(43):41589-41592.
[47]Schug T T,Berry D CShaw N S.Opposing effects of retinoic acid on cell growth result from alternate activation of two different nuclear receptors[J].Cell,2007,129(4):723-733.
[48]Luquet S,Gaudel C,Hoist D,et al..Roles of PPAR delta in lipid absorption and metabolism:a new target for the treatment of type 2 diabetes[J].Biochimica Et Biophysica Acta,2005,1740(2):313-317.
[49]Wang Y X,Zhang C L,Yu R T,et al..Regulation of muscle fiber type and running endurance by PPARdelta[J].Plos Biology,2004,2(10):1532-1539.
[50]王大鹏,金永成,单安山,等.反式视黄酸对3T3-L1前脂肪细胞分化及PPAR7和C/EBPαmRNA表达的影响[J].中国兽医学报,2015(4):645-648.
[51]Marchildon F,St-Louis CAkter R,et al..Transcription factor Smad3 is required for the inhibition of adipogenesis by retinoic acid[J].Journal of Biological Chemistry,2010,285(17):13274-13284.
[52]Schwarz E J,Reginato M J,Shao D,et al..Retinoic acid blocks adipogenesis by inhibiting C/EBPbeta-mediated transcription[J].Molecular and Cellular Biology 17,1997,17(3):1552-1561.
[53]Steppan C M,Bailey S T,Bhat S,et al..The hormone resistin links obesity to diabetes[J].Nature,2001,409(6818):307-312.
[54]Ost A,Danielsson A.Liden M,et al..Retinolbinding protein-4 attenuates insulin-induced phosphorylation of IRS1and ERK1/2 in primary human adipocytes[J].Faseb Journal Official Publication of the Federation of American Societies for Experimental Biology,2007,21(13):3696-3704.
[55]Hollung K,Rise C P,Drevon C A,et al..Tissue-specific regulation of leptin expression and secretion by all-trans retinoic acid[J].Journal of Cellular Biochemistry,2004,92(2):307-315.
[56]Mercader J,Granados N,Bonet M L,et al..All-trans retinoic acid decreases murine adipose retinol binding protein 4production[J].Cellular Physiology and Biochemistry,2008,22(4):363-372.
[57]Song H.Shojima N.Sakoda H,et al..Resistin is regulated by C/EBPs,PPARs,and signal-transducing molecules[J].Biochemical and Biophysical Research Communications,2002,299(2):291-298.
[58]Mercader J.Palou A,Bonet M L.Induction of uncoupling protein-1 in mouse embryonic fibroblast-derived adipocytes by retinoic acid[J].Obesity,2010,18(4):655-662.
[59]Xiong Y,Collins Q F,An J,et al..p38 mitogen-activated protein kinase plays an inhibitory role in hepatic lipogenesis[J].Journal of Biological Chemistry,2007,282(7):4975-4982.
[60]Barger P M,Browning A C,Garner A N,et al..p38 mitogen-activated protein kinase activates peroxisome proliferator-activated receptor alpha:a potential role in the cardiac metabolic stress response[J],Journal of Biological Chemistry,2001,276(48):44495-44501.
[61]Puigserver P,Rhee J,Lin J,et al..Cytokine stimulation of energy expenditure through p38 MAP kinase activation of PPARgamma coactivator-l[J].Molecular Cell,2001,8(5):971-982.
[62]Dong M K,Choi H R,Park A,et al..Retinoic acid inhibits adipogenesis via activation of Wnt signaling pathway in3T3-L1 preadipocytes[J].Biochemical and Biophysical Research Communications,2013,434(3):455-459.
[2]Strom K,Gundersen T E.Hansson 0,et al..Hormone-sensitive lipase(HSL)is also a retinyl ester hydrolase.-evidence from mice lacking HSL[J].The FASEB Journal.Research Communication,2009,23:2307-2316.
[3]Jeyakumar S M,Sheril A,Vajreswari A.Chronic vitamin A-enriched diet feeding induces body weight gain and adiposity in lean and glucose-intolerant obese rats of WNIN/GR-Ob strain[J].Experimental Physiology.2015,100(11):1352-1361.
[4]Ribot J,Felipe F,Bonet M L,et al..Changes of adiposity in response to vitamin A status correlate with changes of PPAR gamma 2 expression[J].Obesity Research.2001,9(8),500-509.
[5]Goodwin K,Abrahamowicz M,Leonard G,et al..Dietary Vitamin A and Visceral Adiposity:A Modulating Role of the Retinol-Binding Protein 4 Gene[J].Journal of Journal of Nutrigenetics and Nutrigenomics.2015,8(4-6):164-173.
[6]Krone K G,Ward A K,Madder KM,et al..Interaction of vitamin A supplementation level with ADH1C genotype on intramuscular fat in beef steers.[J].Animal,2016,10(3):403-409.
[7]Felipe F,Bonet M L,Ribot J,et al..Modulation of resistin expression by retinoic acid and vitamin A status[J].Diabetes,2004,53(4):882-889.
[8]Mercaer J,Ribot J,Murano I,et al..Remodeling of white adipose tissue after retinoic acid administration in mice[J].Endocrinology,2006,147(11):5325-5332.
[9]Amengual J,Ribot J,Bonet M L,et al..Retinoic acid treatment enhances lipid oxidation and inhibits lipid biosynthesis capacities in the liver of mice[J].Cellular Physiology and Biochemistry,2010,25:657-666.
[10]Berry D C,Noy N.All-trans-retinoic acid represses obesity and insulin resistance by activating both peroxisome proliferation-activated receptor beta/delta and retinoic acid receptor[J].Molecular and Cellular Biology,2009,29(12):3286-3296.
[11]Ayuso M,Fernandez A,Isabel B,et al..Long term vitamin A restriction improves meat quality parameters and modifies gene expression in Iberian pigs[J].Journal of Animal Science,2015,93(6):2730-2744.
[12]Silva L D S V D.Veiga G V D.Ramalho R A.Association of serum concentrations of retinol and carotenoids with overweight in children and adolescents[J].Nutrition,2007,23(5):392-397.
[13]Villaca Chaves G,Pereira S E,Saboya C J,et al..Non-alcoholic fatty liver disease and its relationship with the nutritional status of vitamin A in individuals with classⅢobesity[J].Obesity Surgery,2008,18(4):378-385.
[14]Burrows T L,Warren J M,Colyvas K,et al..Validation of overweight children's fruit and vegetable intake using plasma carotenoids[J].Obesity,2009,17:162-168.
[15]Strauss R S.Comparison of serum concentrations of alpha-tocopherol and beta-carotene in a cross-sectional sample of obese and nonobese children(NHANES III)[J].Journal of Pediatrics,1999,134(2):160-165.
[16]Facchini F,Coulston A M,Reaven G M.Relation between dietary vitamin intake and resistance to insulin-mediated glucose disposal in healthy volunteers[J].American Journal of Clinical Nutrition,1996,63(6):946-949.
[17]Mercader J.Granados N,Bonet M L,et al..All-trans retinoic acid decreases murine adipose retinol binding protein 4production[J].Cellular Physiology and Biochemistry,2008,22(4)363-372.
[18]Tourniaire F.Musinovic H.Gouranton E,et al..All-trans retinoic acid induces oxidative phosphorylation and mitochondria biogenesis in adipocytes[J],Journal of Lipid Research,2015,56(6):1100-1109.
[19]Murano I,Morroni M,Zingaretti M C,et al..Morphology of ferret subcutaneous adipose tissue after 6-monthdaily supplementation with oral beta-carotene[J].Biochimica Et Biophysica Acta,2015,1740(2):305-312.
[20]Sanchez J.Fuster A,Oliver P,et al..Effects of beta-carotene supplementation on adipose tissue thermogenic capacity in ferrets(Mustela putorius furo)[J].British Journal of Nutrition,2009,102(11):1686-1694.
[21]Kumar M V,Sunvold G D,Scarpace P J.Dietary vitamin A supplementation in rats suppression of leptin and induction of UCP1 mRNA[J].Journal of Lipid Research,1999,40(5)824-829.
[22]Jeyakumar S M.Vajreswari A.Giridharan N V.Chronic dietary vitamin A supplementation regulates obesity in an obese mutant WNIN/Ob rat model[J].Obesity,2006,14(1):52-59.
[23]Yehya A.Baer J T,Smiley W,et al..Hypervitaminosis A altering the lipid profile in a hypercholesterolemic patient[J].Journal of Clinical Lipidology,2009,3(3):205-207.
[24]Khanna A.Reddy T S.Effect of undernutrition and vitamin A deficiency on the phospholipid composition of rat tissues at 21 days of age-I.Liver,spleen and kidney[J].International Journal for Vitamin and Nutrition Research,1983,53(1):3-8.
[25]Wiss O,Wiss V.Alterations of the lipid metabolism of rat liver as early symptoms of vitamin A deficiency[J].International Journal for Vitamin and Nutrition Research,1980,50(3):233-237.
[26]Tripathy S,Chapman J D,Han C Y,Hogarth C A,et al..All-Trans-Retinoic Acid Enhances Mitochondrial Function in Models of Human Liver[J].Molecular Pharmacology,2016,89(5):560-574.
[27]Mcclinintick J N,Crabb D W,Tian H,et al..Global effects of vitamin A deficiency on gene expression in rat liver:evidence for hypoandrogenism[J].Journal of Nutritional Biochemistry,2006,17(5):345-355.
[28]Oliveros L B,Domeniconi M A,Vega V A,et al..Vitamin A deficiency modifies lipid metabolism in rat liver[J].British Journal of Nutrition,2007,97(2):263-272.
[29]Serra F,Bonet M L.Puigserver P,et al..Stimulation of uncoupling protein 1 expression in brown adipocytes by naturally occurring carotenoids[J].International Journal of Obesity,1999,23(6):650-655.
[30]Felipe F,Bonet M L,Ribot J,et al..Up-regulation of muscle uncoupling protein 3 gene expression in mice following high fat diet,dietary vitamin A supplementation and acute retinoic acid-treatment[J].International Journal of Obesity,2003,27(1):60-69.
[31]Mercader J.Madsen L,Felipe F,et al..All-trans retinoic acid increases oxidative metabolism in mature adipocytes[J].Cellular Physiology and Biochemistry,2007,20(6):1061-1072.
[32]Repa J J,Liang G,Ou J,et al..Regulation of mouse sterol regulatory element-binding protein-lc gene(SREBP-lc)by oxysterol receptors,LXRalpha and LXRbeta[J].Genes and Development,2000,14(22).-2819-2830.
[33]Joseph S B,Laffitte B A.Patel P H,et al..Direct and indirect mechanisms for regulation of fatty acid synthase gene expression by liver X receptors[J].Journal of Biological Chemistry,2002,277(13):11019-11025.
[34]Feige J N,German L.Michalik B,et al..From molecular action to physiological outputs:peroxisome proliferator-activated receptors are nuclear receptors at the crossroads of key cellular functions[J].Progress in Lipid Research,2006,45(2):120-159.
[35]Lefterova M I,Lazar M A.New developments in adipogenesis[J].Medical Clinics of North America,2009,89(6):107-114.
[36]Mandard S,Muller M,Kersten S.Peroxisome proliferator-activated receptor alpha target genes[J].Ppar Research,2010,4:393-416.
[37]Ide T.Shimano H,Yoshikawa T,et al..Cross-talk between peroxisome proliferator-activated receptor(PPAR)alpha and liver X receptor(LXR)in nutritional regulation of fatty acid metabolism.TJ.LXRs suppress lipid degradation gene promoters through inhibition of PPAR signaling[J].Molecular Endocrinology,2003,17(7):1255-1267.
[38]Yoshikawa T,Ide T,Shimano H,et al..Cross-talk between peroxisome proliferator-activated receptor(PPAR)alpha and liver X receptor(LXR)in nutritional regulation of fatty acid metabolism.I.PPARs suppress sterol regulatory element binding protein-lc promoter through inhibition of LXR signaling[J].Molecular Endocrinology,2003,17(7):1240-1254.
[39]Blomhoff R.Blomhoff H K.Overview of retinoid metabolism and function[J].Journal of Neurobiology,2006,66(7):606-630.
[40]Cadoudal T,Glorian M,Massias A,et al..Retinoids upregulate phosphoenolpyruvate carboxykinase and glyceroneogenesis in human and rodent adipocytes[J].Journal of Nutrition,2008,138(6):1004-1009.
[41]Miller C W,Waters K M,Ntambi J M.Regulation of hepatic stearoyl-CoA desaturase gene 1 by vitamin A[J].Biochemical and Biophysical Research Communications,1997,231(1):206-210.
[42]Solanes G,Pedraza N,Iglesias R,et al..The human uncoupling protein-3 gene promoter requires MyoD and is induced by retinoic acid in muscle cells[J].Faseb Journal Official Publication of the Federation of American Societies for Experimental Biology,2000,14(14),2141-2143.
[43]Raisher B D.Gulick T,Zhang Z,et al..Identification of a novel retinoid-responsive element in the promoter region of the medium chain acyl-coenzyme A dehydrogenase gene[J].Journal of Biological Chemistry,1992,267(28):20264-20269.
[44]Yoshikawa T,Shimano H,Amemiya-Kudo M,et al..Identification of liver X receptor-retinoid X receptor as an activator of the sterol regulatory element-binding protein lc gene promoter[J].Molecular and Cellular Biology,2001,21(9):2991-3000.
[45]Taniguchi D,Mizzoguchi Y.Retinoic acids change gene expression profiles of bovine intramuscular adipocyte differentiation,based on microarray analysis.Animal Science Journal.2015,86(6):579-587.
[46]Shaw N,Elholm M,Noy N,et al..Retinoic acid is a high affinity selective ligand for the peroxisome proliferator-activated receptor beta/delta[J].Journal of Biological Chemistry,2003,278(43):41589-41592.
[47]Schug T T,Berry D CShaw N S.Opposing effects of retinoic acid on cell growth result from alternate activation of two different nuclear receptors[J].Cell,2007,129(4):723-733.
[48]Luquet S,Gaudel C,Hoist D,et al..Roles of PPAR delta in lipid absorption and metabolism:a new target for the treatment of type 2 diabetes[J].Biochimica Et Biophysica Acta,2005,1740(2):313-317.
[49]Wang Y X,Zhang C L,Yu R T,et al..Regulation of muscle fiber type and running endurance by PPARdelta[J].Plos Biology,2004,2(10):1532-1539.
[50]王大鹏,金永成,单安山,等.反式视黄酸对3T3-L1前脂肪细胞分化及PPAR7和C/EBPαmRNA表达的影响[J].中国兽医学报,2015(4):645-648.
[51]Marchildon F,St-Louis CAkter R,et al..Transcription factor Smad3 is required for the inhibition of adipogenesis by retinoic acid[J].Journal of Biological Chemistry,2010,285(17):13274-13284.
[52]Schwarz E J,Reginato M J,Shao D,et al..Retinoic acid blocks adipogenesis by inhibiting C/EBPbeta-mediated transcription[J].Molecular and Cellular Biology 17,1997,17(3):1552-1561.
[53]Steppan C M,Bailey S T,Bhat S,et al..The hormone resistin links obesity to diabetes[J].Nature,2001,409(6818):307-312.
[54]Ost A,Danielsson A.Liden M,et al..Retinolbinding protein-4 attenuates insulin-induced phosphorylation of IRS1and ERK1/2 in primary human adipocytes[J].Faseb Journal Official Publication of the Federation of American Societies for Experimental Biology,2007,21(13):3696-3704.
[55]Hollung K,Rise C P,Drevon C A,et al..Tissue-specific regulation of leptin expression and secretion by all-trans retinoic acid[J].Journal of Cellular Biochemistry,2004,92(2):307-315.
[56]Mercader J,Granados N,Bonet M L,et al..All-trans retinoic acid decreases murine adipose retinol binding protein 4production[J].Cellular Physiology and Biochemistry,2008,22(4):363-372.
[57]Song H.Shojima N.Sakoda H,et al..Resistin is regulated by C/EBPs,PPARs,and signal-transducing molecules[J].Biochemical and Biophysical Research Communications,2002,299(2):291-298.
[58]Mercader J.Palou A,Bonet M L.Induction of uncoupling protein-1 in mouse embryonic fibroblast-derived adipocytes by retinoic acid[J].Obesity,2010,18(4):655-662.
[59]Xiong Y,Collins Q F,An J,et al..p38 mitogen-activated protein kinase plays an inhibitory role in hepatic lipogenesis[J].Journal of Biological Chemistry,2007,282(7):4975-4982.
[60]Barger P M,Browning A C,Garner A N,et al..p38 mitogen-activated protein kinase activates peroxisome proliferator-activated receptor alpha:a potential role in the cardiac metabolic stress response[J],Journal of Biological Chemistry,2001,276(48):44495-44501.
[61]Puigserver P,Rhee J,Lin J,et al..Cytokine stimulation of energy expenditure through p38 MAP kinase activation of PPARgamma coactivator-l[J].Molecular Cell,2001,8(5):971-982.
[62]Dong M K,Choi H R,Park A,et al..Retinoic acid inhibits adipogenesis via activation of Wnt signaling pathway in3T3-L1 preadipocytes[J].Biochemical and Biophysical Research Communications,2013,434(3):455-459.