摘要
本研究通过活体药物处理、构建超表达及干扰载体并处理鸡原代脂肪细胞、球状脂联素处理细胞等技术手段,研究脂联素对肉鸡活体脂肪沉积、肌肉品质和鸡脂肪细胞中脂代谢的影响及其调控机制。试验获得以下结果：
1.为了研究肉鸡脂联素表达变化对活体脂肪沉积及肌肉品质的影响,试验采用罗格列酮及地塞米松处理肉鸡,连续处理三周,检测血清中脂联素水平,脂联素及其受体基因的表达规律,同时研究肉鸡体脂沉积,检测脂肪组织中脂代谢关键基因的表达。结果表明：(1)罗格列酮显著上调肉鸡血清中脂联素水平,地塞米松显著抑制血清中脂联索水平,并呈时间依赖性；(2)罗格列酮显著上调腹脂、皮脂、胸肌、腿肌和肝脏中脂联素及其受体AdipoR1基因的表达；(3)罗格列酮显著降低血清中TG和LDL-C水平,上调HDL-C水平,地塞米松显著上调血清中TG和LDL-C水平,下调HDL-C水平；(4)罗格列酮显著抑制腹脂和颈部皮脂以及胸肌和腿肌中的脂质沉积,而地塞米松显著增加肉鸡腹脂、腹部皮脂、颈部皮脂和腿部皮脂以及胸肌和腿肌中脂质沉积；(5)罗格列酮显著下调胸肌和腿肌中总FA、SFA和MUFA的含量,地塞米松则显著促进肌肉中脂肪酸的沉积；(6)罗格列酮处理组肌肉剪切力上升,滴水损失上升,肉色下降,而地塞米松组则相反。(7)罗格列酮显著上调脂肪、肝脏和胸肌中ATGL基因的表达,显著抑制脂肪及肝脏组织中CEBPα和FAS的表达。地塞米松显著上调CEBPα和FAS的表达,而显著抑制ATGL基因的表达。
2.为了研究全长脂联素表达变化对鸡脂肪细胞分化的影响,试验构建了鸡脂联素基因超表达及干扰真核质粒载体,处理鸡原代脂肪细胞细胞。结果表明：(1)超表达脂联素抑制脂肪细胞分化及脂滴生成,而沉默脂联素基因则促进鸡脂肪细胞分化和脂滴生成；(2)超表达脂联素抑制了CEBPα和FAS的表达水平,而促进了ATGL的表达。干扰脂联素基因则促进了CEBPα和FAS的表达,抑制了ATGL的表达；(3)超表达脂联素激活了p38MAPK/ATF-2信号通路,SB253580能够阻断这种活化作用,而干扰脂联素则抑制了p38MAPK/ATF-2通路;(4)超表达脂联素抑制了TOR/p70S6Kinase信号通路,雷帕霉素则能够加强这种抑制作用,而干扰脂联素则激活了TOR/p70S6Kinase信号通路。
3.为了研究全长脂联素的水解产物-球状脂联素对鸡脂肪细胞分化鸡脂质蓄积的影响,试验采用200μg/mL人重组球状脂联素处理鸡原代脂肪细胞。结果表明：(1)球状脂联素能够促进鸡脂肪细胞内源脂联素及AdipoR1基因表达；(2)球状脂联素抑制了鸡脂肪细胞分化和脂质沉积(3)球状脂联素上调CEBPα和FAS的表达,抑制了FAS表达；(4)球状脂联素上调p38MAPK/ATF-2蛋白磷酸化水平,并且能够修复SB253580介导的p38MAPK/ATF-2通路抑制作用；(5)球状脂联素抑制TOR/p70S6Kinase蛋白磷酸化水平,并且能够协同雷帕霉素进一步降低TOR/p70S6Kinase蛋白磷酸化。
4.为了研究鸡脂联素对脂肪细胞中线粒体发育及功能的影响,我们构建了鸡脂联素腺病毒载体处理鸡原代脂肪细胞,并采用干扰真核载体处理细胞。结果表明：(1)超表达脂联素负调控鸡脂肪细胞分化和脂质沉积,超表达脂联素促进CEBPα、LPL和ATGL的表达,抑制FAS、A-FABP和ACC的表达；(2)超表达脂联素促进鸡脂肪细胞24h葡萄糖摄取和培养基中FFA的释放,而干扰脂联素则抑制脂肪细胞24h葡萄糖摄取和培养基中FFA的释放；(3)脂联素促进脂肪细胞中线粒体发育及线粒体内糖脂代谢功能基因的表达；(4)脂联素上调脂肪细胞线粒体膜电位；(5)超表达脂联素促进p38MAPK和ERK的磷酸化水平,抑制TOR蛋白磷酸化。
综上所述,本研究证明了罗格列酮和地塞米松分别上调或下调肉鸡活体内脂联素的表达,并且脂联素表达的变化与肉鸡体脂沉积以及肌肉品质成负相关关系；超表达脂联素以及重组球状脂联素显著抑制了脂肪细胞分化,而干扰脂联素基因则显著促进脂肪细胞分化,p38MAPK和TOR通路参与脂联素对鸡脂肪细胞分化的调节；超表达脂联素促进了鸡脂肪细胞中线粒体的发育并激活线粒体中糖脂代谢功能,并且上调了线粒体膜电位；ERK和TOR信号通路参与了脂联素对线粒体功能的调节。
The present study was conducted to investigate the effect of adiponectin on fat deposition and muscle quality of broiler chicken in vivo and lipid metabolism of chicken adipocyte in vitro, mean while, the mechanism of the action of adiponectin was also observed. Rosiglitazone and dexamethasone were used to treat broilers in vivo for three weeks; over-expression and interference vectors targeting of chicken adiponectin gene were constructed and transfected into chicken preadipocytes;200μg/mL recombinant globular adiponectin was used to treat chicken preadipocyte. At last, recombinant adenovirus vector targeting chicken adiponectin was constructed and treated chicken preadipocytes to clarify the effect of adiponectin on mitochondria development and its function in chicken adipocyte. Results showed that:
1.(1) Rosiglitazone increased serum adiponectin concentration in a time-dependent manner, while dexamethasone increased serum adiponectin;(2) Rosiglitazone up-regulated the expression levels of adiponectin and adipoRl in tissues, while dexamethasone decreased adiponectin and adipoRl mRNA levels;(3) Rosiglitazone decreased the levels of serum lipids, while dexamethasone increased serum lipids;(4) Rosiglitazone decreased abdominal fat, neck subcutaneous fat deposition and lipids in muscle, while dexamethasone increased abdominal fat, neck subcutaneous fat, leg subcutaneous fat, abdominal subcutaneous fat and lipids in muscle;(5) Rosiglitazone down-regulated total FA, SFA and MUFA in muscle, while dexamethasone increased the deposition of total FA,SFA and MUFA in muscle;(6) Rosiglitazone increased the expression of ATGL in tissues and inhibited CEBPa and FAS, while dexamethasone have the opposite effect.
2.(1) overexpression adiponectin inhibited the differentiation of chicken preadipocyte;(2) adiponectin decreased the expression of CEBPa and FAS, while increased ATGL expression;(3) over-expression of adiponectin could activate p38MAPK/ATF-2pathway and also repaired the suppression of p38MAPK and ATF-2activation by SB253580;(4) over-expression of adiponectin suppressed TOR/p70S6Kinase pathway in chicken adipocytes and may further reduce the phosphorylation level of the TOR and p70S6Kinase based on the inhibitory effect of TOR.
3.(1) globular adiponectin inhibited the differentiation of chicken preadipocyte;(2) globular adiponectin decreased the expression of CEBPa and FAS, while increased ATGL expression;(3) globular adiponectin could activate p38MAPK/ATF-2pathway and also repaired the suppression of p38MAPK and ATF-2activation by SB253580;(4) globular adiponectin suppressed TOR/p70S6Kinase pathway in chicken adipocytes and may further reduce the phosphorylation level of the TOR and p70S6Kinase based on the inhibitory effect of TOR.
4.(1) over-expression of adiponectin inhibited chicken adipocyte differentiation, increased the expression of CEBPa, LPL and ATGL, while decreased the expression of FAS, A-FABP and ACC;(2) over-expression of adiponectin promoted glucose uptake and FFA release in medium during24h;(3) over-expression of adiponectin promoted mitochondria development and increased the expression of glucose and lipid metabolism marker genes in the mitochondria;(4) over-expression of adiponectin increased mitochondrial membrane potential;(5) over-expression of adiponectin activated p38MAPK and ERK pathway, while inhibited the activation of TOR pathway.
In summary, in our studies, we proved that rosiglitazone and dexamethasone could regulate the levels of serum adiponectin and analyzed the impact of serum adiponectin on fat deposition and meat quality. Over-expression of full length of adiponectin and globular adiponectin had the same impact on the inhibitory of chicken preadipocyte differentiation. Over-expression of adiponectin promoted mitochondria development and also stimulated glucose and lipid metabolism in mitochondria. p38MAPK, ERK and TOR signaling pathways involved in the regulation of adiponectin on mitochondrial function.
引文
Accorsi PA, Gamberoni M, Isani G, et al.2005. Leptin does not seem to influence glucose uptake by bovine mammary explants. J Physiol Pharmacol,56(4):689-698
Addy CL, Gavrila A, Tsiodras S, et al.2003. Hypoadiponectinemia is associated with insulin resistance, hypertriglyceridemia, and fat redistribution in human immunodeficiency virus-infected patients treated with highly active antiretroviral therapy. J Clin Endocrino Metab,88:62-636
Anderson S, Bankier AT, Barrell BG, de Bruijn MH, Coulson AR, et al.1981. Sequence and organization of the human mitochondrial genome. Nature,410 (5806):457-465
Andrews RC, Walker BR.1999. Glucocorticoids and insulin resistance:old hormones, new targets. Clin Sci, 96:513-523
Arita Y, S Kihara, N Ouchi, K Maeda and H Kuriyama et al.2002. Adipocyte-derived plasma protein adiponectin acts as a platelet-derived growth factor-BB-binding protein and regulates growth factor-induced common postreceptor signal in vascular smooth muscle cell. Circulation,105:2893-2898
Bauche I B, El Mkadem S A, Pottier A M, et al.2007. Overexpression of adiponectin targeted to adipose tissue in transgenic mice:impaired adipocyte differentiation. Endoctrinology,148:1539-1549
Berndt J, Kovacs P, Ruschke K, et al.2007. Fatty acid synthase gene expression in human adipose tissue: association with obesity and type 2 diabetes.Diabetologia,50:1472-1480
Bernlohr, DA, Bolanowski, et al.1985. Evidence for an increase in transcription of specific mRNAs during differen-tiation of 3T3-L1 preadipocytes. J Biol Chem,260:5563-5567
Billon N, Iannarelli P, Monteiro MC, Glavieux-Pardanaud C,Richardson WD, Kessaris N, et al.2007. The generation of adipocytes by the neural crest. Development,134(12):2283-92
Brinkman K, Smeitink JA, Romijn JA.1999. Mitochondrial toxicity induced by nucleoside-analogue reverse-transcriptase inhibitors is a key factor in the pathogenesis of antiretroviral-therapy-related lipodystrophy. Lancet,354:1112-1115
Ceddia RB, Somwar R, Maida A, et al.2005. Globular adiponectin increases GLUT4 translocation and glucose uptake but reduces glycogen synthesis in rat skeletal muscle cells. Diabetologia,48(1):132-9
Chapman AB, Knight DM, Dieckmann BS, et al.1984. Analysis of gene expression during differentiation of adipogenic cells in culture and hormonal control of the developmental program. J Biol Chem,259:15548-15555
Chedid P, Hurtado-Nedelec M, Marion-Gaber B.2012. Adiponectin and its globular fragment differentially modulate the oxidative burst of primary human phagocytes. Am J Pathol,180(2):682-92
Cheng SW, Fryer LG, Carling D, Shepherd PR.2004. Thr2446 is a novel mammalian target of rapamycin (mTOR) phosphorylation site regulated by nutrient status. J Biol Chem,279:15719-15722
Choo HJ, Kim JH, Kwon OB, et al.2006. Mitochondria are impaired in the adipocytes of type 2 diabetic mice. Diabetologia,49:784-791
Christy RJ, Kaestner KH, Geiman DE, et al.1991. CCAAT/enhancer binding protein gene promoter: binding of nuclear factors during differentiation of 3T3-L1 preadipocytes. Proc Natl Acad Sci USA, 88:2593-2597
Clarke SL, Robinson CE, Gimble JM.1997. CAAT/enhancer binding proteins directly modulate transcription from the peroxisome proliferator-activated receptor gamma 2 promoter. Biochem Biophys Res Commun,240:99-103
Clayton DA.1991. Replication and transcription of vertebrate mitochondrial DNA. Annu Rev Cell Bio, 17:453-478
Coletta DK, Sriwijitkamol A, Wajcberg E, et al.2009. Pioglitazone stimulates AMP-activated protein kinase signalling and increases the expression of genes involved in adiponectin signalling, mitochondrial function and fat oxidation in human skeletal muscle in vivo:a randomised trial. Diabetologia,52(4): 723-732
Cornelius P, Enerback S, Bjursell G, et al.1988. Regulation of lipoprotein lipase mRNA content in 3T3-L1 cells by tumor necrosis factor. Biochem J,249:765-769
Cornelius P, MacDougald OA, Lane MD.1994. Regulation of adipocyte development. Annu Rev Nutr,14: 99-129
Dani C, Amri EZ, Bertrand B, et al.1990. Expression and regulation of pOb24 and lipoprotein lipase genes during adipose conversion. J Cell Biochem,43:103-110
Edmondson DG, Olson EN.1993. Helix-loop-helix proteins as regulators of muscle-specific transcription. J Biol Chem,268:755-758
Ekstrand MI, Falkenberg M, Rantanen A, et al.2004. Mitochondrial transcription factor A regulates mtDNA copy number in mammals. Hum Mol Genet,13(9):935-44
Erickson RL, Longo KA, Ross SE, et al.2000. Structure and function of C/EBPa. In:Adipocyte Biology and Hormone Signaling (Ntambi, J M, ed). IOS Press, Inc. Burke, VA
Freytag SO.1988. Enforced expression of the c-myc oncogene inhibits cell differentiation by precluding entry into a distinct predifferentiation state in G0/G1. Mol Cell Biol,8:1614-1624
Fruebis J, Tsao Ts, Javorschi S, et al.2001. Proteolytic cleavage product of 30-kDa adipocyte complement-related protein increases fatty acid oxidation in muscle and causes weight loss in mice. Proc Natl Acad Sci USA,98:2005-2010
Fu Y, Luo N, Klein R L, et al.2005. Adiponectin promotes adiopocyte differentiation, insulin sensitivity, and lipid accumulation. J Lipid Res,46:1369-1379
Fukuhara H, Sor F, Drissi R, et al.1993. Linear mitochondrial DNAs of yeasts:frequency of occurrence and general features. Mol Cell Biol,13 (4):2309-2314
Gagnon A, Lau S, Sorisky A.2001. Rapamycin-sensitive phase of 3T3-L1 preadipocyte differentiation after clonal expansion. J Cell Physiol,189(1):14-22
Ge'loen A, Roy PE, Bukowecki LJ.1989. Regression of white adipose tissue in diabetic rats. Am J Physiol, 257:E547-E553
Green H, Kehinde O.1974. An established pre-adipose cell line and its differentiation in culture. Cell,1: 113-116
Green H, Kehinde O.1979. Formation of normally differentiated subcutaneous fat pads by an established preadipose cell line. J Cell Physiol,101:169-172
Gregoire FM, Smas CM, Sul HS.1998. Understanding adipocyte differentiation. Physiol Rev,78:783-809 Gwartney BL, Jones SJ, Calkins CR, et al.1992. Response time of broiler chickens to cimaterol:Meat tenderness, muscle composition, fiber size, and carcass characteristics. J Anim Sci,70:2144-2150
Haemmerle G, Lass A, Zimmermann R, et al.2006. Defective lipolysis and altered energy metabolism in mice lacking adipose triglyceride lipase. Science,312 (5774):734-737
Hirabayashi M, D Ijiri, Y Kamei, et al.2005. Transformation of skeletal muscle from fast-to slowtwitch during acquisition of cold tolerance in the chick. Endocrinology,146:399-405
Hotta K, Funahashi T, Bodkin NL, et al.2001. Circulating concentrations of the adipocyte protein adiponectin are decreased in parallel with reduced insulin sensitivity during the progression to type 2 diabetes in rhesus monkeys. Diabetes,50(5):1126-33
Hu E, P Liang Spieglman BM.1996. AdipoQ is a novel adipose-specific gene dysregulated in obesity. J Biol Chem,271:10697-10703
Huang PH, Chen JS, HY Tsai, et al.2011. Globular adiponectin improves high glucose-suppressed endothelial progenitor cell function through endothelial nitric oxide synthase dependent mechanisms. J Mol CellCardiol,51(1):109-119
Iwamoto N, Yokoyama S.2011. Protein kinase D regulates the adiponectin gene expression through phosphorylation of AP-2:A common pathway to the ABCA1 gene regulation. Atherosclerosis,216:90-96
Iwabu M, Yamauchi T, Okada-Iwabu M, et al.2010. Adiponectin and AdipoRl regulate PGC-lalpha and mitochondria by Ca (2+) and AMPK/SIRT1. Nature,464(7293):1313-1319
Jacobi SK, Ajuwon KM, Weber TE, et al.2004. Dyer and M.E. Spurlock, Cloning and expression of porcine adiponectin, and its relationship to adiposity, lipogenesis and the acute phase response. J Endocrinol,182:133-144
Jukes TH, Osawa S.1990. The genetic code in mitochondria and chloroplasts. Experientia,46 (11-12): 1117-1126
Kadowaki T, Yamauchi T.2005. Adiponectin and adiponectin receptors. Endocrine Rev,26:439-451
Kelly DP, Scarpullar RC.2004. Transcriptional regulatory circuits controlling mitochondrial biogenesis and function. Genes Dev,18:357-368
Kershaw EE, Hamm JK, Verhagen LA, et al.2006. Adipose triglyceride lipase:function, regulation by insulin, and comparison with adiponutrin. Diabetes,55:148-157
Kim T, Wayne Leitner J, Adochio R, et al.2009. Knockdown of JNK rescues 3T3-L1 adipocytes from insulin resistance induced by mitochondrial dysfunction. Biochem Biophys Res Commun,378 (4):772-776
Koh EH, Park JY, Park HS, et al.2007. Essential role of mitochondrial function in adiponectin synthesis in adipocytes. Diabetes,56(12):2973-2981
Konieczny SF, Emerson CP.1984.5-Azacytidine induction of stable mesodermal stem cell lineages from 10T1/2 cells:evidence for regulatory genes controlling determination. Cell,38:791-800
Lee TS, Lin CY, Tsai JY, et al.2009. Resistin increases lipid accumulation by affecting class A scavenger receptor, CD36 and ATP-binding cassette transporter-Al in macrophages. Life Sci,84:97-104
Lenaz G, DAurelio M, Merlo PM et al.2000. Mitochondrial bioenergetics in aging. Biochim Biophys Acta, 1459(2-3):397-404
Lin FT, Lane MD.1994. CCAAT/enhancer binding protein alpha is sufficient to initiate the 3T3-L1 adipocyte differentiation program. Proc Natl Acad Sci USA,91:8757-8761
Liu B, Wang Y, Wu S, et al.2008. Insulin regulates the expression of adiponectin and adiponectin receptors in porcine adipocytes. Domest Anim Endocrin,34(4):352-359
Long Q, Lei T, Feng B, et al.2010. Peroxisome proliferator-activated receptor-gamma increases adiponectin secretion via transcriptional repression of endoplasmic reticulum chaperone protein ERp44. Endocrinology,51(7):3195-3203
Luo XH, Guo LJ, Yuan LQ, Xie H, Zhou HD, et al.2005. Adiponectin stimulates human osteoblasts proliferation and differentiation via the MAPK signaling pathway. Exp Cell Res,309:99-109
MacDougald OA, Lane MD.1995. Transcriptional regulation of gene expression during adipocyte differentiation. Annu Rev Biochem,64:345-373
Maddineni S, Metzger S, Ocon O, et al.2005. Adiponectin gene is expressed in multiple tissues in the chicken:Food deprivation influences adiponectin messenger ribonucleic acid expression. Endocrinology, 146,4250-4256
Maeda K, K Okubo, I Shimomura, et al.1996. cDNA cloning and expression of a novel adipose specific collagen-like factore, apM1 (Adipose most abundant gene transcript 1). Bioch Biophy Res Commun,221: 286-289
Maeda N, Takahashi M, Funahashi T, et al.2001. PPAR gamma ligands increase expression and plasma concentrations of adiponectin,and adipose derived protein. Diabetes,50:2094-9
Mao X, Kikani CK, Riojas RA, Langlais P, Wang L, et al.2006. APPL1 binds to adiponectin receptors and mediates adiponectin signalling and function. Nat Cell Biol,8:516-523
Masaki T, Chiba S, Yasuda T, et al.2003. Peripheral, but not central, administration of adiponectin reduces visceral adiposity and upregulates the expression of uncoupling protein in agouti yellow (Ay/a) obese mice. Diabetes,52 (9):2266-2273
Mottram DS, Edwards RA.1983. The role of triglycerides and phospholipids in the aroma of cooked beef. J Sci Food Agric,34:517-522
Muhlhausler BS, Morrison JL, Mcmillen IC.2009. Rosiglitazone increases the expression of peroxisome proliferator-activated receptor-gamma target genes in adipose tissue, liver, and skeletal muscle in the sheep fetus in late gestation. Endocrinology,150(9):4287-4294
Nakano YT, Tobe NH, Choi-Miura T, et al.1996. Isolation and characterization of GBP28, a novel gelatin-binding protein purified from human plasma. J Biochem,120:803-812
Nbtsubara M, Namioka K, Katayose S.2003. Decreased plasma adiponectin concentrations in women with low grade c reactive protein elevation. Eur J Endocrinol,148(6):657-62
Negrel R, Grimaldi P, Ailhaud G.1978. Establishment of preadipocyte clonal line from epididymal fat pad of ob/obmouse that responds to insulin and to lipolytic hormones. Proc Natl Acad Sci USA,75:6054-6058
Nishizawa H, Shimomura L, Kishida K, et al.2002. Androgens decrease plasma adiponectin and insulin sensitizing adipocyte derived protein. Diabetes,51:2734-41
Novikoff AB, Novikoff PM, Rosen OM, et al.1980. Organelle relationships in cultured 3T3-L1 preadipocytes. J Cell Biol,87:180-196
Ocon-Grove OM, Krzysik-Walker SM, Maddineni SR, et al.2008. Adiponectin and its receptors are expressed in the chicken testis:influence of sexual maturation on testicular ADIPOR1 and ADIPOR2 mRNA abundance. Reproduction,136,627-638
Ono K, Han J.2000. The p38 signal transduction pathway:activation and function. Cell Signal,12:1-13
Pajvani UB, Du X, Combs TP, et al.2003. Structure-function studies of the adipocyte-secreted hormone Acrp30/adiponectin. Implications fpr metabolic regulation and bioactivity. J Biol Chem,278(11):9073-85.
Qiao L, Schaack J, Shao J.2006. Suppression of adiponectin gene expression by histone deacetylase inhibitor valproic acid. Endocrinology,147 (2):865-874
Qiao L, Kinney B, Schaack J, Shao J.2011. Adiponectin Inhibits Lipolysis in Mouse Adipocytes. Diabetes, 60:1519-1527
Qi Y, N Takahashi, HilemanSM, et al.2004. Adiponectin acts in the brain to decrease body weight. Nat Med,10:524-529
Ramachandran R, O Ocon-Grove and SL Metzger.2007. Molecular cloning and tissue expression of chicken AdipoRl and AdipoR2 complementary deoxyribonucleic acids. Domest Anim Endocrinol,33: 19-31
Raingeaud J, Gupta S, Rogers JS, et al.1995. Pro-inflammatory cytokines and environmental stress cause p38 mitogen-activated protein kinase activation by dual phosphorylation on tyrosine and threonine. J Biol Chem,270:7420-7426
Rei Shibata, Kaon Sato, David R Pimentel, et al.2005. Adiponectin protects against myocardial ischemia-reperfusion injury through AMPK-and COX-2-dependent mechanisms. Nat Med,11(10): 1096-1103
Reznikoff CA, Brankow DW, Heidelberger C.1973. Establishment and characterization of a cloned line of C3H mouse embryo cells sensitive to post-confluence inhibition of division. Cancer Res,33:3231-3238
Roth J, Qiang X, Marban SL, et al.2004. The obesity pandemic:where have we been and where are we going? Obes Res,2:88S-101S
Saito K, Tobe T, Minoshima S, et al.1999. Organization of the gene for gelatin-binding protein (GBP28).Gene,229(1-2):67-73
Salas-Marco J, Fan-Minogue H, Kallmeyer AK, et al.2006. Distinct paths to stop codon reassignment by the variant-code organisms Tetrahymena and Euplotes. Mol Cell Biol,26 (2):438-447
Sato K, Akiba Y, Chida Y, et al.1999. Lipoprotein hydrolysis and fat accumulation in chicken adipose tissues are reduced by chronic administration of lipoprotein lipase monoclonal antibodies. Poult Sci,78: 1286-1291
Sattar AA, Sattar R.2012. Globular adiponectin activates Akt in cultured myocytes. Biochem Biophys Res Commun,424(4):753-757
Scherer PE, S Williams, M Fgliano, et al.1995. A novel serum protein similar to Clq, produced exclusively in adipocytes. J Biol Chem,27:26746-26749
Scott RE, FlorineDL, Wille JJ, et al.1982. Coupling of growth arrest and differentiation at a distinct state in the G1 phase of the cell cycle:GD. Proc Natl Acad Sci USA,79:845-849
Serrero G, Lepak NM, Goodrich SP.1992. Paracrine regulation of adipose differentiation by arachidonate metabolites:prostaglandin F2 alpha inhibits early and late markers of differentiation in the adipogenic cell line 1246. Endocrinology,131:2545-2551
Shao D, Lazar MA.1997. Peroxisome proliferator activated receptor gamma, CCAAT/enhancer-binding protein alpha, and cell cycle status regulate the commitment to adipocyte differentiation. J Biol Chem,272: 21473-21478
Shao R, Kirkness EF, Barker SC.2009. The single mitochondrial chromosome typical of animals has evolved into 18 minichromosomes in the human body louse, Pediculns humanus. Geno Res,19 (5): 904-912
Shi X, Burkart A, Nicoloro S M, et al.2008. Paradoxical effect of mitochondrial respiratory chain impairment on insulin signaling and glucose transport in adipose cells. J Biol Chem,283 (45): 30658-30667
Shuman JD, Cheong J, Coligan JE.1997. ATF-2 and C/EBPalpha can form a heterodimeric DNA binding complex in vitro. Functional implications for transcriptional regulation. J Biol Chem,272:12793-12800
Skpzypski M, Let, Kaczarek P, et al.2011. Orexin A stimulates glucose uptake, lipid accumulation and adiponectin secretion from 3T3-L1 adipocytes and isolated primary rat adipocytes. Diabetologia, 54(7):1841-52
SM Khan, RM Smigrodzki, R Swerdlow.2007. Cell and animal models of mtDNA biology:progress and prospects. Ameri J Physiol Cell Physiol,292 (2):C658-C669
Smas CM, Sul HS.1995. Control of adipocyte differentiation. Biochem J,309:697-710
Staiger H, S Kaltenbach, K Staiger, et al.2004. Expression of adiponectin receptor mRNA in human skeletal muscle cells is related to in vivo parameters of glucose and lipid metabolism. Diabetes,53: 2195-2201
Sukhija PS, Palmquist DL.1988. Rapid method for determination of total fatty acid content and composition of feedstuffs and feces, J Agri Food Chem,36:1202-1206
Tahmoorespur M, Ghazanfari S, Nobari K.2010. Evaluation of adiponectin gene expression in the abdominal adipose tissue of broiler chickens:Feed restriction, dietary energy, and protein influences adiponectin messenger ribonucleic acid expression. Poult Sci,89 (10):2092-2100
Takahashi M, Afita Y, Yamagata K, et al.2000. Genomic structure and mutations in adipose-specific gene, adiponectin. J Obes Relat Metab,24(7):861-868
Tan GD, Debard C, Funahashi T, et al.2005. Changes in adiponectin receptor expression in muscle and adipose tissue of type 2 diabetic patients during rosiglitazone therapy. Diabetologia,48:1585-1589.
Taylor SM, Jones PA.1979. Multiple new phenotypes induced in 10T1/2 and 3T3 cells treated with 5-azacytidine. Cell,17:771-779
Thompson JM.2004. The effects of marbling on flavour and juiciness scores of cooked beef after adjusting to a constant tenderness. Aust J Exp Agri,44:645-652
Tsao TS, Murrey HE, Hug C, Lee DH, Lodish HF.2002. Oligomerization state-dependent activation of NF-kappa B signaling pathway by adipocyte complement-related protein of 30 kDa (Acrp30). J Biol Chem, 277(33):29359-62
Tsuchida TA, Takekawa S, Hada Y, et al.2005. Peroxisome proliferator-activated receptor (PPAR)-a activation increases adiponectin receptors and reduces obesity-related inflammation in adipose tissue: comparison of activation of PPAR, Diabetes,54:33-58
Vankoningsloo S, De Pauw A, Houbion A, et al.2006. CREB activation induced by mitochondrial dysfunction triggers triglyceride accumulation in 3T3-L1 preadipocytes. J Cell Sci,119:1266-1282
Wang YX, Zhan XA, Yuan D, et al.2011. Effects of selenomethionine and sodium selenite supplementation on meat quality, selenium distribution and antioxidant status in broilers, Czech J Anim Sci,56:305-313
Weyer C, Funahashi T, Tanaka S, et al.2001. Hypoadiponectinemia in obesity and type 2 diabetes:close association with insulin resistance and hyperinsulinemia. J Clin Endocrinol Metab,86(5):1930-1935
Wilson-Fritch L, Burkart A, Bell G, et al.2003. Mitochondrial biogenesis and remodeling during adipogenesis and in response to the insulin sensitizer rosiglitazone. Mol Cell Bio,123:1085-1094
Wu Z, Xie Y, Bucher NLR, et al.1995. Conditional ectopic expression of C/EBPb in NIH3T3 cells induces PPAR g and stimulates adipogenesis. Genes Dev,9:2350-2363
Xu A, Tso AWK, Cheung BMY, et al.2007. Circulating adipocyte-fatty acid binding protein levels predict the development of the metabolic syndrome-A 5-year prospective study. Circulation,115:1537-1543
Xu AM, Wang Y, Keshaw H, et al.2003. The fat-derived hormone adiponectin alleviates alcoholic and nonalcoholic fatty liver diseases in mice. J Clin Invest,112:91-100
Yamao F, Muto A, Kawauchi Y, et al.1985. UGA is read as tryptophan in Mycoplasma capricolum. Proc Natl Acad Sci USA,82 (8):2306-2309
Yamauchi T, J Kamon, H Waki, et al.2001.The fat-derived hormone adiponectin reverses insulin resistance associated with both lipoatrophy and obesity. Nat Med,7:941-946
Yamauchi T, J Kamon, H Waki, et al.2003. Globular adiponectin protected ob/ob mice from diabetes and ApoE-deficient mice from atherosclerosis. J Biol Chem,278:2461-2468
Yamauchi T, J Kamon, Y Minokoshi, et al.2002. Adiponectin stimulates glucose utilization and fatty acid oxidation by activating AMP-activated protein kinase. Nat Med,8:1288-1295
Yamauchi T, Kamon J, Ito Y, et al.2003. Cloning of adiponectin receptors that mediate antidiabetic metabolic effects. Nature,423:762-769
Yeh WC, Bierer BE, McKnight SL.1995. Rapamycin inhibits clonal expansion and adipogenic differentiation of 3T3-L1 cells. Proc Natl Acad Sci USA,92:11086-11090
Yuan J, W Liu, ZL Liu et al.2006. cDNA cloning, genomic structure, chromosomal mapping and expression analysis of ADIPOQ (adiponectin) in chicken. Cytogenet Genome Res,112:148-151
Yoon MJ, Lee GY, YH Chung, et al.2006. Adiponectin increases fatty acid oxidation in skeletal muscle cells by sequential activation of AMP-activated protein kinase, p38 mitogen-activated protein kinase, and peroxisome proliferator-activated receptor alpha. Diabetes,55:2562-2570
Zimmermann R, Strauss JG, Haemmerle G, et al.2004. Fat mobilization in adipose tissue is promoted by adipose triglyceride lipase. Science,306 (5700):1383-1386
董飚,孟和,顾志良等.2007.脂联素与脂联素受体的研究进展.中国畜牧兽医,34(2)：44-48
党永明、房亚东、胡炯宇等.2010.成体大鼠心肌细胞微管解聚对线粒体分布及能量代谢的影响.中华烧伤杂志,26(1)：18-22
胡小波,张育坚,陈霞等.2005.胰岛素、Dex、肿瘤坏死因子α能调节脂联素的表达和分泌.医学分子生物学杂志,2(2)：90-93
李晓南,陈荣华,潘亚等.2006.人类皮下和网膜脂肪组织中脂联素基因的表达和调控的研究.南京医科大学学报,26(1)：6-9
马泰,朱启星.2006.线粒体形态学改变与细胞凋亡.细胞生物学杂志,28：671-675
曲杨燕,刘云海,郭勇等.2009.绵羊卵母细胞体外成熟过程中的线粒体分布变化.畜牧兽医学报,(8)：1258-1261
吴家睿.2011.脂肪细胞分化：一个故事、两个章节.中国科学,56(17)：1327-1334
舒相华,刘勇,宋春莲等.2008.添加共轭亚油酸对肉鸡肌肉脂肪酸影响的研究.中国畜牧兽医,35(10)：27-28
肖静,张英,于炜婷等.2010.微囊化HepG2细胞脂代谢与线粒体功能和蛋白的合成.中国组织工程研究与临床康复,14(38)：7082-7086
徐秋良,吴运香.畜禽肉嫩度及其影响因素.2010.家畜生态学报,31(6)：100-103
张力莉,徐晓锋.2012.动物脂肪酸合成酶基因表达调控的研究进展.畜牧与兽医,44(4)：101-103