山羊Kruppel样转录因子家族在前体脂肪细胞分化中的表达模式及相关性分析
|
摘要
【目的】优化山羊前体脂肪细胞体外培养体系,揭示Kruppel样转录因子家族(KLFs)成员在前体脂肪细胞分化过程中的表达模式。【方法】于四川省简阳大哥大牧业有限公司随机选取3只7日龄简州大耳羊为研究对象。放血致死并清洗后,在无菌环境中取其皮下脂肪组织。利用Ⅰ型胶原酶消化法获得其皮下前体脂肪细胞,待细胞长至细胞密度为80%时传代至6孔培养板中(F3代),以"150μmol·L~(-1)油酸+10 mg·L~(-1)胰岛素"前体脂肪细胞进行成脂诱导,并在诱导分化后0 d、3 d、5 d、7 d收集细胞,Trizol法提取细胞总RNA,利用实时荧光定量PCR(RT-qPCR)检测KLF2~(-1)0, KLF 12和KLF15共11个KLF转录因子家族成员在不同分化阶段细胞中的表达水平,并利用皮尔逊系数分析各个成员mRNA总体表达水平之间的相关性。【结果】150μmol·L~(-1)油酸+10 mg·L~(-1)胰岛素处理前体脂肪细胞24 h后细胞中开始出现小脂滴,48 h后脂滴数量增多、体积增大,诱导120 h后脂肪细胞分化程度显著增加。RT-qPCR结果显示,脂肪分化标志基因过氧化物酶体增殖物活化受体γ(PPARγ)基因随着诱导分化的进行其表达水平持续上升,其中在第7天时其表达水平极显著高于其他各组(P<0.01)。KLF2、KLF3、KLF4、KLF6、KLF7和KLF15在脂肪细胞中表达水平显著(P<0.05)或极显著(P<0.01)高于前体脂肪细胞中表达水平,其中KLF3、KLF6和KLF15在分化5 d时表达水平最高,KLF2和KLF4表达水平在7 d时最高,而KLF7则在3 d时表达水平最高;KLF5、KLF8、KLF9、KLF10和KLF12在前体脂肪细胞中的表达水平极显著高于分化后脂肪细胞中的表达水平(P<0.01),其中KLF8、KLF10和KLF12的表达水平均在5 d时达到最低,而KLF5和KLF9则在7 d时表达水平最低;相关性分析结果显示KLF家族各成员mRNA在山羊前体脂肪细胞分化过程中总体表达水平存在相关性,KLF12与6个基因具有较强的相关性,其次为KLF4和KLF9,分别与5个成员具有较强的相关性,而KLF2与其他基因均没有较强的相关性。【结论】明确了KLF家族在山羊前体脂肪细胞分化过程中的表达模式,解析了各个成员mRNA表达之间的相关性,为阐明山羊前体脂肪细胞分化的分子机制提供重要的基础数据。
【Objective】The aim of this study was to optimize the culture system of goat preadipocytes in vitro and elucidate the expression patterns of Kruppel like factors(KLFs) during preadipocytes differentiation. 【Method】Three 7-day-old Jianzhou Big-Eared goats were selected randomly from Sichuan Jianyang DAGEDA Animal Husbandry Co. Ltd. as experimental animal.After bloodletting death of goats, their subcutaneous fat tissues were collected in a sterile room. The goat subcutaneous preadipocytes were collected using collagenase I digestion method. The cells at Generation Three were then passaged to a 6-well plate when the cells were grown to 80% density. The combined use of were used for preadipocytes differentiate inducing. The cells were then collected after cultivation for 0 d, 3 d, 5 d and 7 d. The total RNA was extracted by using Trizol method. Real-time fluorescent quantitative PCR(RT-qPCR) was performed to determine the expression levels of 11 members of KLFs, including KLF2~(-1)0, KLF12 and KLF15, during goat preadipocytes differentiation. Pearson correlation coefficient was used for expression correlation analysis between KLF individuals.【Result】The results showed that the combined use of 150 μmol·L~(-1) oleic acid and 10 mg·L~(~(-1)) insulin promoted the formation of small lipid droplets after 24 h cultivation. After 48 h, the lipid droplets became bigger and much more.The differentiation of the preadipocytes were enhanced significantly after 120 h comparing with preadipocytes at 0 h. The results of RT-qPCR showed that the combined use of oleic acid and insulin stimulated the expression of peroxisome proliferator-activated receptor-gamma(PPARγ) continuously companying the differentiation of preadipocytes, and toped at 7 d which significantly higher than other groups(P<0.01). The expression levels of KLF2, KLF3, KLF4, KLF6, KLF7 and KLF15 in adipocytes were significantly(P<0.05) or extremely significantly(P<0.01) higher than those in preadipocytes. While the expression of KLF3, KLF6 and KLF15 were topped at 5 d, the KLF2 and KLF4 toped at 7 d. However, the KLF7 expressed most at 3 d. For the other KLF individuals, the expression levels of KLF5, KLF8, KLF9, KLF10 and KLF12 in preadipocytes were extremely significantly higher(P<0.01) than those in adipocytes. The expression of KLF8, KLF10 and KLF12 bottomed at 5 d, while that of the KLF5 and KLF9 bottomed at 7 d.Correlation analysis showed that KLF individual significantly(P<0.05) correlated with each other across cell differentiation. KLF12 was strongly correlated with 6 individuals, followed by KLF4 and KLF9 both of which were significantly correlated with 5 individuals. However, KLF2 was not significantly correlated with other individuals. 【Conclusion】In this study, the expression patterns of KLFs during goat preadipocytes differentiation were clarified, and also the correlations among KLF individuals. These results provided important basic data for elucidating the molecular mechanism of goat preadipocytes differentiation.
【Objective】The aim of this study was to optimize the culture system of goat preadipocytes in vitro and elucidate the expression patterns of Kruppel like factors(KLFs) during preadipocytes differentiation. 【Method】Three 7-day-old Jianzhou Big-Eared goats were selected randomly from Sichuan Jianyang DAGEDA Animal Husbandry Co. Ltd. as experimental animal.After bloodletting death of goats, their subcutaneous fat tissues were collected in a sterile room. The goat subcutaneous preadipocytes were collected using collagenase I digestion method. The cells at Generation Three were then passaged to a 6-well plate when the cells were grown to 80% density. The combined use of were used for preadipocytes differentiate inducing. The cells were then collected after cultivation for 0 d, 3 d, 5 d and 7 d. The total RNA was extracted by using Trizol method. Real-time fluorescent quantitative PCR(RT-qPCR) was performed to determine the expression levels of 11 members of KLFs, including KLF2~(-1)0, KLF12 and KLF15, during goat preadipocytes differentiation. Pearson correlation coefficient was used for expression correlation analysis between KLF individuals.【Result】The results showed that the combined use of 150 μmol·L~(-1) oleic acid and 10 mg·L~(~(-1)) insulin promoted the formation of small lipid droplets after 24 h cultivation. After 48 h, the lipid droplets became bigger and much more.The differentiation of the preadipocytes were enhanced significantly after 120 h comparing with preadipocytes at 0 h. The results of RT-qPCR showed that the combined use of oleic acid and insulin stimulated the expression of peroxisome proliferator-activated receptor-gamma(PPARγ) continuously companying the differentiation of preadipocytes, and toped at 7 d which significantly higher than other groups(P<0.01). The expression levels of KLF2, KLF3, KLF4, KLF6, KLF7 and KLF15 in adipocytes were significantly(P<0.05) or extremely significantly(P<0.01) higher than those in preadipocytes. While the expression of KLF3, KLF6 and KLF15 were topped at 5 d, the KLF2 and KLF4 toped at 7 d. However, the KLF7 expressed most at 3 d. For the other KLF individuals, the expression levels of KLF5, KLF8, KLF9, KLF10 and KLF12 in preadipocytes were extremely significantly higher(P<0.01) than those in adipocytes. The expression of KLF8, KLF10 and KLF12 bottomed at 5 d, while that of the KLF5 and KLF9 bottomed at 7 d.Correlation analysis showed that KLF individual significantly(P<0.05) correlated with each other across cell differentiation. KLF12 was strongly correlated with 6 individuals, followed by KLF4 and KLF9 both of which were significantly correlated with 5 individuals. However, KLF2 was not significantly correlated with other individuals. 【Conclusion】In this study, the expression patterns of KLFs during goat preadipocytes differentiation were clarified, and also the correlations among KLF individuals. These results provided important basic data for elucidating the molecular mechanism of goat preadipocytes differentiation.
引文
[1]STEPHENS J M.The fat controller:adipocyte development.PLoSBiology,2012,10(11):e1001436.
[2]KACZYNSKI J,COOK T,URRUTIA R.Sp1 and Kruppel-like transcription factors.Genome Biology,2003,4(2):206.
[3]LEE D S,CHOI H,HAN B S,KIM W K,LEE S C,OH K J,BAE KH.c-Jun regulates adipocyte differentiation via the KLF15-mediated mode.Biochemical and Biophysical Research Communications,2016,469(3):552-558.
[4]WU Z N,WANG S Q.Role of kruppel-like transcription factors in adipogenesis.Developmental Biology,2013,373(2):235-243.
[5]PEI J,GRISHIN N V.A new family of predicted Krüppel-like factor genes and pseudogenes in placental mammals.PLoS ONE,2013,8(11):e81109.
[6]WEI S J,ZHANG L F,ZHOU X,DU M,JIANG Z H,HAUSMAN GJ,BERGEN W G,ZAN L S,DODSON M V.Emerging roles of zinc finger proteins in regulating adipogenesis.Cellular and Molecular Life Sciences,2013,70(23):4569-4584.
[7]BIRSOY K,CHEN Z,FRIEDMAN J.Transcriptional regulation of adipogenesis by KLF4.Cell Metabolism,2008,7(4):339-347.
[8]PARK Y K,WANG L M,GIAMPIETRO A,LAI B B,LEE J E,GE K.Distinct roles of transcription factors KLF4,Krox20 and PPARγin adipogenesis.Molecular and Cellular Biology,2017,37(2):e00554-16.
[9]JIANG S Z,WEI H K,SONG T X,YANG Y,ZHAGN F,ZHOU Y F,PENG J,JIANG S W.KLF13 promotes porcine adipocyte differentiation through PPARγactivation.Cell&Bioscience,2015,5:28.
[10]GUO H F,KHAN R,RAZA S H A,NING Y,WEI,D W,HOSSEINIS M,ULLA I,GARCIA M D,ZAN L S.KLF15 promotes transcription of KLF3 gene in bovine adipocytes.GENE,2018,659(15):77-83.
[11]池永东,王永,朱江江,林森,赵越,林亚秋.藏山羊KLF8基因克隆及组织表达分析.基因组学与应用生物学,2019,38(2):478-502.CHI Y D,WANG Y,ZHU J J,LIN S,ZHAO Y,LIN Y Q.Cloning and tissue expression analysis of klf8 gene in Tibetan goat.Genomics and Applied Biology,2019,38(2):478-502.(in Chinese)
[12]田苗,左璐璐,张小玉,柏雪,江明锋,林亚秋.牦牛KLF10基因克隆及组织表达分析.黑龙江畜牧兽医,2017,17(09/1):234-236.TIAN M,ZUO L L,ZHANG X Y,BAI X,JIANG M F,LIN Y Q.Cloning and tissue expression analysis of Klf10 gene in Yak.Heilongjiang Animal Science and Veterinary Medicine,2017,17(09/1):234-236.(in Chinese)
[13]朱江江,林亚秋,王永,李倩,林森,熊朝瑞,俄木曲者.山羊KLF13基因的克隆及表达分析.农业生物技术学报.2017,25(1):102-109.ZHU J J,LIN Y Q,WANG Y,LI Q,LIN S,XIONG C R,EMU Q Z.Cloning and expression analysis of goat(Capra hircus)KLF13 gene.Journal of Agricultural Biotechnology,2017,25(1):102-109.(in Chinese)
[14]NEVE B,FERNANDEZ-ZAPICO M E,ASHKENAZI-KATALAN V,DINA C,HAMID Y H,JOLY E,VAILLANT E,BENMEZROUA Y,DURAND E,BAKAHER N,DELANNOY V,VAXILLAIRE M,COOK T,DALLINGA-THIE G M,JANSEN H,CHARLES M A,CLEMENT K,GALAN P,HERCBERG S,HELBECQUE N,CHARPENTIER G,PRENTKI M,HANSEN T,PEDERSEN O,URRUTIA R,MELLOUL D,FROGUEL P.Role of transcription factor KLF11 and its diabetes-associated gene variants in pancreatic beta cell function.Proceedings of the National Academy of Sciences of the United States of America,2005,102(13):4807-4812.
[15]WU G,YUAN S,CHEN Z,CHEN G,FAN Q,DONG H,YE F,LI J,ZHU X.The KLF14 transcription factor regulates glycolysis by downregulating LDHB in colorectal cancer.International Journal of Biological Sciences,2019,15(3):628-635.
[16]WANG J,GALVAO J,BEACH K M,LUO W,URRUTIA R A,GOLDBERG J L,OTTESON D C.Novel roles and mechanism for Kruppel-like Factor 16(KLF16)regulation of neurite outgrowth and ephrin receptor A5(EphA5)expression in retinal ganglion cells.Journal of Biological Chemistry,2016,291(35):18084-18095.
[17]ALI A,ZHANG P,LIANGFANG Y,WENSHE S,WANG H,LIN X,DAI Y,FENG X H,MOSES R,WANG D,LI X,XIAO J.KLF17empowers TGF-β/Smad signaling by targeting Smad3-dependent pathway to suppress tumor growth and metastasis during cancer progression.Cell Death&Disease,2015,6(3):e1681.
[18]PEARSON R,FLEETWOOD J,EATON S,CROSSLEY M,BAO S.Krüppel-like transcription factors:A functional family.International Journal of Biochemistry and Cell Biology,2008,40(10):1996-2001.
[19]FUNNELL A P W,MALONEY C A,THOMPSON L J,KEYS J,TALLACK M,PERKINS A C,CROSSLEY M.Erythroid Kruppellike factor directly activates the basic Kruppel-like factor gene in erythroid cells.Molecular and Cellular Biology,2007,27(7):2777-2790.
[20]EATON S A,FUNNELL A P W,SUE N,NICHOLAS H,RICHARDC M,CROSSLEY M.A network of Krüppel-like factors(Klfs).Journal of Biological Chemistry,2008,283(40):26937-26947.
[21]郭红芳,昝林森,孙永刚.牛前体脂肪细胞的分离培养及诱导分化.西北农林科技大学学报(自然科学版),2012,42(2):1-6.GUO H F,ZAN L S,SUN Y G.Primary culture and differentiation of bovine preadipocytes.Journal of Northwest A&F University,2012,42(2):1-6.(in Chinese)
[22]SHANG Z C,GUO L,WANG N,SHI H,WANG Y X,LI H.Oleate promotes differentiation of chicken primary preadipocytes in vitro.Bioscience Reports,2014,34(1):e00093.
[23]吉红,曹艳姿,林亚秋,刘品,卢荣华,苏尚顺,杨公社,奥宏海.草鱼前体脂肪细胞的原代培养.水生生物学报,2009,33(6):1226-1230.JI H,CAO Y Z,LIN Y Q,LIU P,LU R H,SU S S,YANG G S,AO HH.Primary culture of grass carp preadipocyte in vitro.Acta Hydrobiologica Sinica,2009,33(6):1226-1230.(in Chinese)
[24]蔡勇,阿依木古丽,杨具田,马忠仁,卢建雄,臧荣鑫,吴建平.绵羊前体脂肪细胞的原代培养及分化.动物营养学报,2010,22(6):1768-1774.CAI Y,AYIMUGULI,YANG J T,MA Z R,LU J X,ZANG R X,WUJ P.Primary culture and differentiation of Ovine preadipocytes.Chinese Journal of Animal Nutrition,2010,22(6):1768-1774.(in Chinese)
[25]曲贞晓,刘金凤,姜颖,潘庆杰.山羊前体脂肪细胞体外培养试验.青岛农业大学学报(自然科学版),2013,30(1):1-5.QU Z X,LIU J F,JIANG Y,PAN Q J.The culture experiment of goats preadipocytes in vitro.Journal of Qingdao Agricultural University(Natural Science),2013,30(1):1-5.(in Chinese)
[26]杜琛,付绍印,韩志玲,孟丽云,爱伦高娃,高丽霞,成立新,张文广,李金泉.绒山羊肌内前体脂肪细胞的基因表达分析.畜牧兽医学报,2013,44(10):1532-1538.DU C,FU S Y,HAN Z L,MENG L Y,AILUN G W,GAO L X,CHENG L X,ZHANG W G,LI J Q.Gene expression analysis on intramuscular preadipocytes in the cashmere goats.Acta Veterinaria et Zootechnica Sinica,2013,44(10):1532-1538.(in Chinese)
[27]CHEGURU P,CHAPALAMADUGU K C,DOUMIT M E,MURDOCH G K,HILL R A.Adipocyte differentiation-specific gene transcriptional response to C18 unsaturated fatty acids plus insulin.Pflugers Archiv-European Journal of Physiologyl,2012,463(3):429-447.
[28]DING S T,WANG J C,MERSMANN H J.Effect of unsaturated fatty acids on porcine adipocyte differentiation.Nutrition Research,2003,23(8):1059-1069.
[29]ADINA K S,ROBERT Y H,M.DANIEL L.Induction of fatty acid synthetase synthesis in differentiai 3T3-Ll preadipocytes.The Journal of Biological Chemistry,1980,255(25):4745-4750.
[30]BANERJEE S S,FEINBERG M W,WATANABE M,GRAY S,HASPEL R L,DENKINGER D J,KAWAHARA R,HAUNER H,JAIN M K.The Kruppel-like factor KLF2 inhibits peroxisome proliferator-activated receptor-gamma expression and adipogenesis.Journal of Biological Chemistry,2003,278(4):2581-2584.
[31]LEE H,KANG R,KIM Y S,CHUANG S I,YOON Y.Platycodin Dinhibits adipogenesis of 3T3-L 1 cells by modulating kruppel-like factor 2 and peroxisome proliferator-activated receptorγ.Phytotherapy Research,2010,24(S2):S161-S167.
[32]SUE N,JACK B H A,EATON S A,PEARSON R C M,FUNNELLA P W,TURNER J,CZOLIJ R,DENYER G,BAO S S,MOLERO-NAVAJAS J C,PERKINS A,FUJIWARA Y,ORKIN S H,BELL-ANDERSON K,CROSSLEY M.Targeted disruption ofthe basic Kriippel-like factor gene(Klf3)reveals a role in adipogenesis.Molecular and Cellular Biology,2008,28(12):3967-3978.
[33]EISENSTEIN A,CARROLL S H,JOHNSTON-COX H,FARB M,GOKCE N,RAVID K.An adenosine receptor-Krüppel-like factor 4protein axis inhibits adipogenesis.Journal of Biological Chemistry,2014,289(30):21071-21081.
[34]ZHANG Z W,WANG H X,SUN Y N,LI H,WANG N.Klf7modulates the differentiation and proliferation of chicken preadipocyte.Acta Biochimical et Biophysica Sinica(Shanghai),2013,45(4):280-288.
[35]LI D,YEA S,LI S,CHEN Z,NORLA G,BANCK M,LABORDA J,TAN S,FRIEDMAN J M,FRIEDMAN S L,WALSH M J.Kruppellike factor-6 promotes preadipocyte differentiation through histone deacetylase 3-dependent repression of DLK1.Journal of Biological Chemistry,2005,280(29):26941-26952.
[36]MATSUBARA Y,AOKI M,ENDO T,SATO K.Characterization of the expression profiles of adipogenesis-related factors,ZNF423,KLFs and FGF10,during preadipocyte differentiation and abdominal adipose tissue development in chickens.Comparative Biochemistry and Physiology B-Biochemistry&Molecular Biology,2013,165(3):189-195.
[37]AHMED M,GAFFEN S L.IL-17 inhibits adipogenesis in part via C/EBPα,PPARγand Krüppel-like factors.Cytokine,2013,61(3):898-905.
[38]OISHI Y,MANABE I,TOBE K,SHINDO T,FUJIU K,NISHIMURAG,MAEMURA K,YAMAUCHI T,KUBOTA N,SUZUKI R,KITAMURA T,AKIRA S,KADOWAKI T,NAGAI R.Kruppel-like transcription factor KLF5 is a key regulator of adipocyte differentiation.Cell Metabolism,2005,1(1):27-39.
[39]CERVANTES-CAMACHO C,BELTRáN-LANGARICA A,OCHOA-URIBE A K,MARSCH-MORENO M,AYALA-SUMUANO JT,VELEZ-DELVALLE C,KURI-HARCUCH W.The transient expression of Klf4 and Klf5 during adipogenesis depends on GSK3βactivity.Adipocyte,2015,4(4):248-255.
[40]LEE H,KIM H J,LEE Y J,LEE M Y,CHOI H,LEE H,KIM J W.Krüppel-like factor KLF8 plays a critical role in adipocyte differentiation.PLoS ONE,2012,7(12):e52474.
[41]PEI H,YAO Y,YANG Y,LIAO K,WU J R.Krüppel-like factor KLF9regulates PPARγtransactivation at the middle stage of adipogenesis.Cell Death and Differentition,2011,18(2):315-327.
[42]ILSLEY M D,GILLINDER K R,MAGOR G W,HUANG S,BAILEY T L,CROSSLEY M,PERKINS A C.Krüppel-like factors compete for promoters and enhancers to fine-tune transcription.Nucleic Acids Research,2017,45(11):6572-6588.
[43]FUNNELL A P,MAK K S,TWINE N A,PELKA G J,NORTON L J,RADZIEWIC T,POWER M,WIKINS M R,BELL-ANDERSON K S,FRASER S T,PERKINS A C,TAM P P,PEARSON R C M,CROSSLEY M.Generation of mice deficient in both KLF3/BKLFand KLF8 reveals a genetic interaction and a role for these factors in embryonic globin gene silencing.Molecular and Cellular Biology,2013,33(15):2976-2987.
[2]KACZYNSKI J,COOK T,URRUTIA R.Sp1 and Kruppel-like transcription factors.Genome Biology,2003,4(2):206.
[3]LEE D S,CHOI H,HAN B S,KIM W K,LEE S C,OH K J,BAE KH.c-Jun regulates adipocyte differentiation via the KLF15-mediated mode.Biochemical and Biophysical Research Communications,2016,469(3):552-558.
[4]WU Z N,WANG S Q.Role of kruppel-like transcription factors in adipogenesis.Developmental Biology,2013,373(2):235-243.
[5]PEI J,GRISHIN N V.A new family of predicted Krüppel-like factor genes and pseudogenes in placental mammals.PLoS ONE,2013,8(11):e81109.
[6]WEI S J,ZHANG L F,ZHOU X,DU M,JIANG Z H,HAUSMAN GJ,BERGEN W G,ZAN L S,DODSON M V.Emerging roles of zinc finger proteins in regulating adipogenesis.Cellular and Molecular Life Sciences,2013,70(23):4569-4584.
[7]BIRSOY K,CHEN Z,FRIEDMAN J.Transcriptional regulation of adipogenesis by KLF4.Cell Metabolism,2008,7(4):339-347.
[8]PARK Y K,WANG L M,GIAMPIETRO A,LAI B B,LEE J E,GE K.Distinct roles of transcription factors KLF4,Krox20 and PPARγin adipogenesis.Molecular and Cellular Biology,2017,37(2):e00554-16.
[9]JIANG S Z,WEI H K,SONG T X,YANG Y,ZHAGN F,ZHOU Y F,PENG J,JIANG S W.KLF13 promotes porcine adipocyte differentiation through PPARγactivation.Cell&Bioscience,2015,5:28.
[10]GUO H F,KHAN R,RAZA S H A,NING Y,WEI,D W,HOSSEINIS M,ULLA I,GARCIA M D,ZAN L S.KLF15 promotes transcription of KLF3 gene in bovine adipocytes.GENE,2018,659(15):77-83.
[11]池永东,王永,朱江江,林森,赵越,林亚秋.藏山羊KLF8基因克隆及组织表达分析.基因组学与应用生物学,2019,38(2):478-502.CHI Y D,WANG Y,ZHU J J,LIN S,ZHAO Y,LIN Y Q.Cloning and tissue expression analysis of klf8 gene in Tibetan goat.Genomics and Applied Biology,2019,38(2):478-502.(in Chinese)
[12]田苗,左璐璐,张小玉,柏雪,江明锋,林亚秋.牦牛KLF10基因克隆及组织表达分析.黑龙江畜牧兽医,2017,17(09/1):234-236.TIAN M,ZUO L L,ZHANG X Y,BAI X,JIANG M F,LIN Y Q.Cloning and tissue expression analysis of Klf10 gene in Yak.Heilongjiang Animal Science and Veterinary Medicine,2017,17(09/1):234-236.(in Chinese)
[13]朱江江,林亚秋,王永,李倩,林森,熊朝瑞,俄木曲者.山羊KLF13基因的克隆及表达分析.农业生物技术学报.2017,25(1):102-109.ZHU J J,LIN Y Q,WANG Y,LI Q,LIN S,XIONG C R,EMU Q Z.Cloning and expression analysis of goat(Capra hircus)KLF13 gene.Journal of Agricultural Biotechnology,2017,25(1):102-109.(in Chinese)
[14]NEVE B,FERNANDEZ-ZAPICO M E,ASHKENAZI-KATALAN V,DINA C,HAMID Y H,JOLY E,VAILLANT E,BENMEZROUA Y,DURAND E,BAKAHER N,DELANNOY V,VAXILLAIRE M,COOK T,DALLINGA-THIE G M,JANSEN H,CHARLES M A,CLEMENT K,GALAN P,HERCBERG S,HELBECQUE N,CHARPENTIER G,PRENTKI M,HANSEN T,PEDERSEN O,URRUTIA R,MELLOUL D,FROGUEL P.Role of transcription factor KLF11 and its diabetes-associated gene variants in pancreatic beta cell function.Proceedings of the National Academy of Sciences of the United States of America,2005,102(13):4807-4812.
[15]WU G,YUAN S,CHEN Z,CHEN G,FAN Q,DONG H,YE F,LI J,ZHU X.The KLF14 transcription factor regulates glycolysis by downregulating LDHB in colorectal cancer.International Journal of Biological Sciences,2019,15(3):628-635.
[16]WANG J,GALVAO J,BEACH K M,LUO W,URRUTIA R A,GOLDBERG J L,OTTESON D C.Novel roles and mechanism for Kruppel-like Factor 16(KLF16)regulation of neurite outgrowth and ephrin receptor A5(EphA5)expression in retinal ganglion cells.Journal of Biological Chemistry,2016,291(35):18084-18095.
[17]ALI A,ZHANG P,LIANGFANG Y,WENSHE S,WANG H,LIN X,DAI Y,FENG X H,MOSES R,WANG D,LI X,XIAO J.KLF17empowers TGF-β/Smad signaling by targeting Smad3-dependent pathway to suppress tumor growth and metastasis during cancer progression.Cell Death&Disease,2015,6(3):e1681.
[18]PEARSON R,FLEETWOOD J,EATON S,CROSSLEY M,BAO S.Krüppel-like transcription factors:A functional family.International Journal of Biochemistry and Cell Biology,2008,40(10):1996-2001.
[19]FUNNELL A P W,MALONEY C A,THOMPSON L J,KEYS J,TALLACK M,PERKINS A C,CROSSLEY M.Erythroid Kruppellike factor directly activates the basic Kruppel-like factor gene in erythroid cells.Molecular and Cellular Biology,2007,27(7):2777-2790.
[20]EATON S A,FUNNELL A P W,SUE N,NICHOLAS H,RICHARDC M,CROSSLEY M.A network of Krüppel-like factors(Klfs).Journal of Biological Chemistry,2008,283(40):26937-26947.
[21]郭红芳,昝林森,孙永刚.牛前体脂肪细胞的分离培养及诱导分化.西北农林科技大学学报(自然科学版),2012,42(2):1-6.GUO H F,ZAN L S,SUN Y G.Primary culture and differentiation of bovine preadipocytes.Journal of Northwest A&F University,2012,42(2):1-6.(in Chinese)
[22]SHANG Z C,GUO L,WANG N,SHI H,WANG Y X,LI H.Oleate promotes differentiation of chicken primary preadipocytes in vitro.Bioscience Reports,2014,34(1):e00093.
[23]吉红,曹艳姿,林亚秋,刘品,卢荣华,苏尚顺,杨公社,奥宏海.草鱼前体脂肪细胞的原代培养.水生生物学报,2009,33(6):1226-1230.JI H,CAO Y Z,LIN Y Q,LIU P,LU R H,SU S S,YANG G S,AO HH.Primary culture of grass carp preadipocyte in vitro.Acta Hydrobiologica Sinica,2009,33(6):1226-1230.(in Chinese)
[24]蔡勇,阿依木古丽,杨具田,马忠仁,卢建雄,臧荣鑫,吴建平.绵羊前体脂肪细胞的原代培养及分化.动物营养学报,2010,22(6):1768-1774.CAI Y,AYIMUGULI,YANG J T,MA Z R,LU J X,ZANG R X,WUJ P.Primary culture and differentiation of Ovine preadipocytes.Chinese Journal of Animal Nutrition,2010,22(6):1768-1774.(in Chinese)
[25]曲贞晓,刘金凤,姜颖,潘庆杰.山羊前体脂肪细胞体外培养试验.青岛农业大学学报(自然科学版),2013,30(1):1-5.QU Z X,LIU J F,JIANG Y,PAN Q J.The culture experiment of goats preadipocytes in vitro.Journal of Qingdao Agricultural University(Natural Science),2013,30(1):1-5.(in Chinese)
[26]杜琛,付绍印,韩志玲,孟丽云,爱伦高娃,高丽霞,成立新,张文广,李金泉.绒山羊肌内前体脂肪细胞的基因表达分析.畜牧兽医学报,2013,44(10):1532-1538.DU C,FU S Y,HAN Z L,MENG L Y,AILUN G W,GAO L X,CHENG L X,ZHANG W G,LI J Q.Gene expression analysis on intramuscular preadipocytes in the cashmere goats.Acta Veterinaria et Zootechnica Sinica,2013,44(10):1532-1538.(in Chinese)
[27]CHEGURU P,CHAPALAMADUGU K C,DOUMIT M E,MURDOCH G K,HILL R A.Adipocyte differentiation-specific gene transcriptional response to C18 unsaturated fatty acids plus insulin.Pflugers Archiv-European Journal of Physiologyl,2012,463(3):429-447.
[28]DING S T,WANG J C,MERSMANN H J.Effect of unsaturated fatty acids on porcine adipocyte differentiation.Nutrition Research,2003,23(8):1059-1069.
[29]ADINA K S,ROBERT Y H,M.DANIEL L.Induction of fatty acid synthetase synthesis in differentiai 3T3-Ll preadipocytes.The Journal of Biological Chemistry,1980,255(25):4745-4750.
[30]BANERJEE S S,FEINBERG M W,WATANABE M,GRAY S,HASPEL R L,DENKINGER D J,KAWAHARA R,HAUNER H,JAIN M K.The Kruppel-like factor KLF2 inhibits peroxisome proliferator-activated receptor-gamma expression and adipogenesis.Journal of Biological Chemistry,2003,278(4):2581-2584.
[31]LEE H,KANG R,KIM Y S,CHUANG S I,YOON Y.Platycodin Dinhibits adipogenesis of 3T3-L 1 cells by modulating kruppel-like factor 2 and peroxisome proliferator-activated receptorγ.Phytotherapy Research,2010,24(S2):S161-S167.
[32]SUE N,JACK B H A,EATON S A,PEARSON R C M,FUNNELLA P W,TURNER J,CZOLIJ R,DENYER G,BAO S S,MOLERO-NAVAJAS J C,PERKINS A,FUJIWARA Y,ORKIN S H,BELL-ANDERSON K,CROSSLEY M.Targeted disruption ofthe basic Kriippel-like factor gene(Klf3)reveals a role in adipogenesis.Molecular and Cellular Biology,2008,28(12):3967-3978.
[33]EISENSTEIN A,CARROLL S H,JOHNSTON-COX H,FARB M,GOKCE N,RAVID K.An adenosine receptor-Krüppel-like factor 4protein axis inhibits adipogenesis.Journal of Biological Chemistry,2014,289(30):21071-21081.
[34]ZHANG Z W,WANG H X,SUN Y N,LI H,WANG N.Klf7modulates the differentiation and proliferation of chicken preadipocyte.Acta Biochimical et Biophysica Sinica(Shanghai),2013,45(4):280-288.
[35]LI D,YEA S,LI S,CHEN Z,NORLA G,BANCK M,LABORDA J,TAN S,FRIEDMAN J M,FRIEDMAN S L,WALSH M J.Kruppellike factor-6 promotes preadipocyte differentiation through histone deacetylase 3-dependent repression of DLK1.Journal of Biological Chemistry,2005,280(29):26941-26952.
[36]MATSUBARA Y,AOKI M,ENDO T,SATO K.Characterization of the expression profiles of adipogenesis-related factors,ZNF423,KLFs and FGF10,during preadipocyte differentiation and abdominal adipose tissue development in chickens.Comparative Biochemistry and Physiology B-Biochemistry&Molecular Biology,2013,165(3):189-195.
[37]AHMED M,GAFFEN S L.IL-17 inhibits adipogenesis in part via C/EBPα,PPARγand Krüppel-like factors.Cytokine,2013,61(3):898-905.
[38]OISHI Y,MANABE I,TOBE K,SHINDO T,FUJIU K,NISHIMURAG,MAEMURA K,YAMAUCHI T,KUBOTA N,SUZUKI R,KITAMURA T,AKIRA S,KADOWAKI T,NAGAI R.Kruppel-like transcription factor KLF5 is a key regulator of adipocyte differentiation.Cell Metabolism,2005,1(1):27-39.
[39]CERVANTES-CAMACHO C,BELTRáN-LANGARICA A,OCHOA-URIBE A K,MARSCH-MORENO M,AYALA-SUMUANO JT,VELEZ-DELVALLE C,KURI-HARCUCH W.The transient expression of Klf4 and Klf5 during adipogenesis depends on GSK3βactivity.Adipocyte,2015,4(4):248-255.
[40]LEE H,KIM H J,LEE Y J,LEE M Y,CHOI H,LEE H,KIM J W.Krüppel-like factor KLF8 plays a critical role in adipocyte differentiation.PLoS ONE,2012,7(12):e52474.
[41]PEI H,YAO Y,YANG Y,LIAO K,WU J R.Krüppel-like factor KLF9regulates PPARγtransactivation at the middle stage of adipogenesis.Cell Death and Differentition,2011,18(2):315-327.
[42]ILSLEY M D,GILLINDER K R,MAGOR G W,HUANG S,BAILEY T L,CROSSLEY M,PERKINS A C.Krüppel-like factors compete for promoters and enhancers to fine-tune transcription.Nucleic Acids Research,2017,45(11):6572-6588.
[43]FUNNELL A P,MAK K S,TWINE N A,PELKA G J,NORTON L J,RADZIEWIC T,POWER M,WIKINS M R,BELL-ANDERSON K S,FRASER S T,PERKINS A C,TAM P P,PEARSON R C M,CROSSLEY M.Generation of mice deficient in both KLF3/BKLFand KLF8 reveals a genetic interaction and a role for these factors in embryonic globin gene silencing.Molecular and Cellular Biology,2013,33(15):2976-2987.