小麦糖基转移酶基因TaUGT73D1的克隆及表达分析
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摘要
为探讨TaUGT73D1基因在小麦花发育过程中雄蕊和雌蕊形成的作用,本研究利用小麦三雌蕊突变体TP和雄蕊同源转化型不育突变体HTS-1两个品种,从中克隆得到了TaUGT73D1(UDP-glycosyltransferase73D1)的3个同源基因TaUGT73D1-2A、TaUGT73D1-2B、TaUGT73D1-2D,通过基因碱基序列分析表明TaUGT73D1-2A、TaUGT73D1-2B、TaUGT73D1-2D基因序列全长分别为1 907 bp、1 912 bp、1 905 bp,开放阅读框分别为1 365 bp、1 467 bp、1 461 bp,分别编码454、488、486个氨基酸残基,通过与小麦基因组比对,发现这3个同源基因分别位于染色体2AL、2BL、2DL上。TaUGT73D1-2A、TaUGT73D1-2B、TaUGT73D1-2D与乌拉尔图小麦Tu UGT73D1的编码区相似度分别为97.68%、96.52%、99.05%,蛋白序列相似度分别为73.57%、95.08%、95.70%。系统进化分析表明,该类基因与单子叶植物的UGT基因聚在一起,表明其亲缘关系较近,而与双子叶植物相隔较远,表明其亲缘关系较远。利用Real-time PCR分析TTaUGT73D1-2A、TaUGT73D1-2B、TaUGT73D1-2D基因在不同花器官雌蕊(P)、雄蕊(S)及雌蕊化雄蕊(PS)中表达模式。结果表明,该类基因在雄蕊中表达量最高,在雌蕊及雌蕊化中几乎不表达。因此推测TaUGT73D1基因可能与小麦雄蕊同源转化为雌蕊性状有关。本研究为进一步探讨小麦TaUGT73D1基因的功能提供了理论依据。
To investigate the role of TaUGT73D1 gene in the formation of wheat stamens and pistils during the development of wheat flower, three homologous genes TaUGT73D1-2A, TaUGT73D1-2B and TaUGT73D1-2D were cloned of TaUGT73D1(UDP-glycosyltransferase 73D1) from wheat three pistils TP and pistillody mutant HTS-1 in this study. By nucleotide sequence analysis, the results showed that TaUGT73D1-2A, TaUGT73D1-2B,TaUGT73D1-2D were 1 907 bp, 1 912 bp, 1 905 bp in full-length, respectively. The ORF of TaUGT73D1-2A,TaUGT73D1-2B and TaUGT73D1-2D were 1 365 bp, 1 467 bp, 1461 bp, and coded for deduced protein of 454,488 and 486 amino acids, respectively. By comparing with the wheat genome, three TaUGT73D1 homologous genes(TaUGT73D1-2A, TaUGT73D1-2B, TaUGT73D1-2D) were isolated and found to be located on chromosomes2 AL, 2BL, and 2DL, respectively. The Coding region of TaUGT73D1-2A, TaUGT73D1-2B and TaUGT73D1-2D with Tu UGT73D1 are respectively 97.68%, 96.52% and 99.05% similarity. And the similarity of protein sequence was 73.57%, 95.08% and 95.70%, respectively. Phylogenetic analysis showed that the TaUGT73D1 genes was most closely related to monocots` UGT genes, indicating their close genetic relationship. However, it was far from the diocts, which indicated their distant relationship. Real-time PCR was used to studythe pattern of TaUGT73D1-2A,TaUGT73D1-2B and TaUGT73D1-2D expression in pistil(P), stamen(S) and pistillody stamen(PS). The results showed that TaUGT73D1-2A, TaUGT73D1-2B and TaUGT73D1-2D had a higher expression levels in Stamen, but very rare in Pistillody, Stamen and Pistil. So we speculated that TaUGT73D1 gene may be related to the homologous transformation of stamen into pistil in wheat. This study provided a theoretical basis for the further study of TaUGT73D1 gene function in wheat.
To investigate the role of TaUGT73D1 gene in the formation of wheat stamens and pistils during the development of wheat flower, three homologous genes TaUGT73D1-2A, TaUGT73D1-2B and TaUGT73D1-2D were cloned of TaUGT73D1(UDP-glycosyltransferase 73D1) from wheat three pistils TP and pistillody mutant HTS-1 in this study. By nucleotide sequence analysis, the results showed that TaUGT73D1-2A, TaUGT73D1-2B,TaUGT73D1-2D were 1 907 bp, 1 912 bp, 1 905 bp in full-length, respectively. The ORF of TaUGT73D1-2A,TaUGT73D1-2B and TaUGT73D1-2D were 1 365 bp, 1 467 bp, 1461 bp, and coded for deduced protein of 454,488 and 486 amino acids, respectively. By comparing with the wheat genome, three TaUGT73D1 homologous genes(TaUGT73D1-2A, TaUGT73D1-2B, TaUGT73D1-2D) were isolated and found to be located on chromosomes2 AL, 2BL, and 2DL, respectively. The Coding region of TaUGT73D1-2A, TaUGT73D1-2B and TaUGT73D1-2D with Tu UGT73D1 are respectively 97.68%, 96.52% and 99.05% similarity. And the similarity of protein sequence was 73.57%, 95.08% and 95.70%, respectively. Phylogenetic analysis showed that the TaUGT73D1 genes was most closely related to monocots` UGT genes, indicating their close genetic relationship. However, it was far from the diocts, which indicated their distant relationship. Real-time PCR was used to studythe pattern of TaUGT73D1-2A,TaUGT73D1-2B and TaUGT73D1-2D expression in pistil(P), stamen(S) and pistillody stamen(PS). The results showed that TaUGT73D1-2A, TaUGT73D1-2B and TaUGT73D1-2D had a higher expression levels in Stamen, but very rare in Pistillody, Stamen and Pistil. So we speculated that TaUGT73D1 gene may be related to the homologous transformation of stamen into pistil in wheat. This study provided a theoretical basis for the further study of TaUGT73D1 gene function in wheat.
引文
Airoldi C.A.,2010,Determination of sexual organ development Sexual Plant Reproduction,23(1):53-62
Alvarez-Buylla E.R.,Azpeitia E.,Barrio R.,Benítez M.,and Padilla-Longoria P.,2010,From ABC genes to regulatory networks,epigenetic landscapes and flower morphogenesis making biological sense of theoretical approaches,Seminar in Cell&Developmental Biology,21(1):108-117
Angenent G.C.,Franken J.,Busscher M.,Van Dijken A.,Van Went J.L.,Dons H.J.M.,and Van Tunen A.J.V.,1995,Anovel class of MADS box genes is involved in ovule devel opment in petunia,The Plant Cell,7(10):1569-1582
Bowman J.L.,Smyth D.R.,and Meyerowitz E.M.,1991,Geneti interactions among floral homeotic genes of Arabidopsis Development,112(1):1-20
Chang Y.Y.,Kao N.H.,Li J.Y.,Hsu W.H.,Liang Y.L.,Wu J.W.and Yang C.H.,2009,Characterization of the possible role for B class MADS box genes in regulation of perianth forma tion in orchid,Plant Physiology,152(2):837-853
Chen J.S.,Zhang L.H.,and Wu B.X.,1983,A preminary report on the discovery and breeding of the“Trigrain Wheat”,1983Zuowu Xuebao(Acta Agronomica Sinica),9(1):69-71(陈济世,张岭华,吴秉礼,1983,“三粒小麦”的发现及选育初报,作物学报,9(1):69-71)
Chen N.,Hu D.Q.,Pan L.J.,Chi X.Y.,Chen M.N.,Wang T.Wang M.,Yang Z.,and Yu S.L.,2014,Cloning of UDPglucosyltransferase gene from peanut(Arachis hypogaea L.and its expression analysis during abiotic stress,Zhongguo Youliao Zuowu Xuebao(Chinese Journal of Oil Crop Sci ences),36(3):308-315(陈娜,胡东青,潘丽娟,迟晓云,陈明娜,王通,王冕,杨珍,禹山林,2014,花生中UDP-葡萄糖基转移酶基因的克隆及在非生物胁迫下的表达研究中国油料作物学报,36(3):308-315)
Gao X.M.,Xia Y.M.,and Li Q.J.,2006,Isolation of two putativ homologues of PISTILLATA and AGAMOUS from Alpini oblongifolia(Zingiberaceae)and characterization of their ex pression,Plant Science,170(3):674-684
Honma T.,and Goto K.,2001,Complexes of MADS-box pro teins are sufficient to convert leaves into floral organs,Na ture,409(6819):525-529
Ji X.N.,He F.,Duan C.Q.,and Wang J.,2013,Recent progress in biochemical properties and functions of UDP-glycosyl trans ferase during plant secondary metabolism,Shipin Kexu(Food Science),34(9):316-323(姬向楠,何非,段长青,王军,2013,植物UDP-糖基转移酶生化特性和功能研究进展,食品科学,34(9):316-323)
Kanno A.,Nakada M.,Akita Y.,and Hirai M.,2007,Class Bgene expression and the modified ABC model in Nongras monocots,The Scientific World Journal,7(7):268-279
Kobayashi K.,Maekawa M.,Miyao A.,Hirochika H.,and Ky ozuka J.,2010,PANICLE PHYTOMER2(PAP2),encoding a SEPALLATA subfamily MADS-box protein,positively controls Spikelet meristem identity in rice,Plant&Cel Physiology,51(1):47-57
Li Y.X.,Zhang A.M.,and Huang T.C.,1996,Relationship be tween w heat cytoplasm ic m ale sterility and the content o endogenous horm ones in the anther,Nongye Shengwu Jishu Xuebao(Journal of Agricultural Biotechnology),4(4)307-313(李英贤,张爱民,黄铁城,1996,小麦细胞质雄性不育与花药组织内源激素的关系,农业生物技术学报,4(4):307-313)
Ling H.Q.,Zhao S.,Liu D.C.,Wang J.Y.,Sun H.,Zhang C.,Fan H.J.,Li D.,Dong L.L.,Tao Y.,Gao C.,Wu H.L.,Li Y.W.,Cu Y.,Guo X.S.,Zheng S.S.,Wang B.,Yu K.,Liang Q.S.,Yang W.L.,Lou X.Y.,Chen J.,Feng M.J.,Jian J.B.,Zhang X.F.Luo G.B.,Jiang,Y.,Liu J.J.,Wang Z.B.,Sha Y.H.,Zhang B.R.,Wu H.J.,Tang D.Z.,Shen Q.H.,Xue P.Y.,Zou S.H.Wang X.J.,Liu X.,Wang F.M.,Yang Y.P.,An X.L.,Dong Z.Y.,Zhang K.P.,Zhang X.Q.,Luo M.C.,Dvorak J.,Tong YP.,Wang J.,Yang H.M.,Li Z.S.,Wang D.W.,Zhang A.Mand Wang J.,2013,Draft genome of the wheat A-genom progenitor Triticum urartu,Nature,496(7443):87-90
Liska A.J.,and Cassman K.G.,2008,Towards standardization o Life-Cycle metrics for biofuels:greenhouse gas emission mitigation and Net Energy Yield,Journal of Biobased Mate rials&Bioenergy,2(3):187-203
Litt A.,and Kramer E.M.,2010,The ABC model and the diversi fication of floral organ identity,Seminars in Cell&Devel opmental Biology,21(1):129-137
Liu L.S.,Wang C.X.,and Feng L.,eds.,2002,Plant Molecula Genetics,Science Press,Beijing,China,pp.426-434(刘良式,王春新,冯亮,编著,2002,植物分子遗传学,科学出版社,中国,北京,pp.426-434)
Liu S.J.,Zhu Q.,Xing X.J.,Wang B.Q.,Yu F.X.,and Zhou H.2014,Advanced research on male sterility of plant,Zhong guo Nongxue Tongbao(Chinese Agricultural Science Bul letin),30(34):46-50(刘淑娟,朱祺,幸学俊,王碧琴,余发新,周华,2014,植物雄性不育影响因素研究进展,中国农学通报,30(34):46-50)
Liu Z.,Yan J.P.,Li D.K.,Luo Q.,Yan Q.,Liu Z.B.,Y L.M.Wang J.M.,Li X.F.,and Yang Y.,2015,UDP-glucosyltrans ferase71C5,a major glucosyltransferase,mediates abscisi acid homeostasis in Arabidopsis,Plant Physiol.,167(4)1659-1670
Livak K.J.,and Schmittgen T.D.,2001,Analysis of relative gen expression data using Real-time quantitative PCR and th2-ΔΔCT(Delta Delta C(T))method,Methods,25(4):402-408
Murai K.,Takumi S.,Koga H.,and Ogihara Y.,2002,Pistillody homeotic transformation of stamens into pistil-like struc tures,caused by nuclear-cytoplasm interaction in wheat,Th Plant Journal,29(2):169-181
Nam J.,Depamphilis C.W.,Hong MMa H.,and Nei M.,2003Antiquity and evolution of the MADS-box gene family con trolling flower development in plants,Molecular Biology&Evolution,20(9):1435-1447
Pelaz S.,Ditta G.S,Baumann E.,Wisman E.,and Yanofsky M.F.2000,B and C floral organ identity functions require Sepal lata EPALLATA mads MADS-box genes,Nature,405(6783)200-203
Pelaz S.,Tapia-López R.,Alvarez-Buylla E.R.,and Yanofsky MF.,2001,Conversion of leaves into petals in Arabidopsis Current Biology,11(3):182-184
Peng Z.S.,2003,A new mutation in wheat producing three pistil in a floret,Journal of Agronomy and Crop Science,189(4)270-272
Peng Z.S.,Yang J.,Wei S.H.,and Zeng J.H.,2004,Characteriza tion of the common wheat(Triticum aestivum L.)mutation line producing three pistils in a floret,Hereditas,141(1)15-18
Prokopyk D.O.,and Ternovska T.K.,2011,Homeotic genes and their role in development of morphological traits in wheat Cytology and Genetics,45(1):41-54
Wang Y.W.,Peng Z.S.,Yang Z.J.,Wei S.H.,Liao M.L.,Zhao H.Yang H.,Yang Y.F.,and Wang Q.H.,2014,Cloning and ex pression analysis of Ta AP1-3 genes of three pistils mutation in wheat,Huabei Nongxuebao(Acta Agriculturae Boreali-Sinica),29(6):28-35(王育伟,彭正松,杨在君,魏淑红,廖明莉赵欢,杨会,杨宇凤,王清海,2014,小麦三雌蕊突变体Ta AP1-3基因的克隆及表达分析,华北农学报,29(6):28-35)
Yamaguchi T.,and Hirano H.Y.,2006,Function and diversifica tion of MADS-box genes in rice,The Scientific World Jour nal,6:1923-1932
Yang Y.F.,Peng Z.S.,Yang Z.J.,Wei S.H.,Liao M.L.,Wang YW.,Wang Q.H.,and Yang H.,2015,Cloning and expression analysis of Class-B MADS-box genes WPI1 and WPI2 in wheat,Henongxue Bao(Journal of Nuclear Agricultural Sci ences),29(2):221-228(杨宇凤,彭正松,杨在君,魏淑红廖明莉,王育伟,王清海,杨会,2015,小麦B类MADS-bo基因WPI1和WPI2的克隆及表达分析,核农学报,29(2)221-228)
Zhang Y.,and Liu Q.L.,2003,Proceedings on molecular mecha nism of plant flower development,Zhiwuxue Tongbao(Chi nese Bulletin of Bontany),20(5):589-601(张云,刘青林2003,植物花发育的分子机理研究进展,植物学通报,20(5):589-601)
Zhang Y.,Sui Z.H.,Ding H.Y.,and Zhong J.,2011,Cloning and analysis of the UDP-Glucose pyrophosphoryiase(UGPase gene of Gracilaria leimaneiformis(Rhodophyta)and correla tion between gene expression and agar synthesis,Zhongguo
Haiyang Daxue Xuebao(Periodical of Ocean University o China(National Science Edtion)),41(S):80-86(张杨,隋正红,丁弘叶,仲洁,2011,龙须菜UDP-葡萄糖焦磷酸化酶基因的克隆及其表达与琼胶产量的关系,中国海洋大学学报(自然科学版),41(S):80-86)
Alvarez-Buylla E.R.,Azpeitia E.,Barrio R.,Benítez M.,and Padilla-Longoria P.,2010,From ABC genes to regulatory networks,epigenetic landscapes and flower morphogenesis making biological sense of theoretical approaches,Seminar in Cell&Developmental Biology,21(1):108-117
Angenent G.C.,Franken J.,Busscher M.,Van Dijken A.,Van Went J.L.,Dons H.J.M.,and Van Tunen A.J.V.,1995,Anovel class of MADS box genes is involved in ovule devel opment in petunia,The Plant Cell,7(10):1569-1582
Bowman J.L.,Smyth D.R.,and Meyerowitz E.M.,1991,Geneti interactions among floral homeotic genes of Arabidopsis Development,112(1):1-20
Chang Y.Y.,Kao N.H.,Li J.Y.,Hsu W.H.,Liang Y.L.,Wu J.W.and Yang C.H.,2009,Characterization of the possible role for B class MADS box genes in regulation of perianth forma tion in orchid,Plant Physiology,152(2):837-853
Chen J.S.,Zhang L.H.,and Wu B.X.,1983,A preminary report on the discovery and breeding of the“Trigrain Wheat”,1983Zuowu Xuebao(Acta Agronomica Sinica),9(1):69-71(陈济世,张岭华,吴秉礼,1983,“三粒小麦”的发现及选育初报,作物学报,9(1):69-71)
Chen N.,Hu D.Q.,Pan L.J.,Chi X.Y.,Chen M.N.,Wang T.Wang M.,Yang Z.,and Yu S.L.,2014,Cloning of UDPglucosyltransferase gene from peanut(Arachis hypogaea L.and its expression analysis during abiotic stress,Zhongguo Youliao Zuowu Xuebao(Chinese Journal of Oil Crop Sci ences),36(3):308-315(陈娜,胡东青,潘丽娟,迟晓云,陈明娜,王通,王冕,杨珍,禹山林,2014,花生中UDP-葡萄糖基转移酶基因的克隆及在非生物胁迫下的表达研究中国油料作物学报,36(3):308-315)
Gao X.M.,Xia Y.M.,and Li Q.J.,2006,Isolation of two putativ homologues of PISTILLATA and AGAMOUS from Alpini oblongifolia(Zingiberaceae)and characterization of their ex pression,Plant Science,170(3):674-684
Honma T.,and Goto K.,2001,Complexes of MADS-box pro teins are sufficient to convert leaves into floral organs,Na ture,409(6819):525-529
Ji X.N.,He F.,Duan C.Q.,and Wang J.,2013,Recent progress in biochemical properties and functions of UDP-glycosyl trans ferase during plant secondary metabolism,Shipin Kexu(Food Science),34(9):316-323(姬向楠,何非,段长青,王军,2013,植物UDP-糖基转移酶生化特性和功能研究进展,食品科学,34(9):316-323)
Kanno A.,Nakada M.,Akita Y.,and Hirai M.,2007,Class Bgene expression and the modified ABC model in Nongras monocots,The Scientific World Journal,7(7):268-279
Kobayashi K.,Maekawa M.,Miyao A.,Hirochika H.,and Ky ozuka J.,2010,PANICLE PHYTOMER2(PAP2),encoding a SEPALLATA subfamily MADS-box protein,positively controls Spikelet meristem identity in rice,Plant&Cel Physiology,51(1):47-57
Li Y.X.,Zhang A.M.,and Huang T.C.,1996,Relationship be tween w heat cytoplasm ic m ale sterility and the content o endogenous horm ones in the anther,Nongye Shengwu Jishu Xuebao(Journal of Agricultural Biotechnology),4(4)307-313(李英贤,张爱民,黄铁城,1996,小麦细胞质雄性不育与花药组织内源激素的关系,农业生物技术学报,4(4):307-313)
Ling H.Q.,Zhao S.,Liu D.C.,Wang J.Y.,Sun H.,Zhang C.,Fan H.J.,Li D.,Dong L.L.,Tao Y.,Gao C.,Wu H.L.,Li Y.W.,Cu Y.,Guo X.S.,Zheng S.S.,Wang B.,Yu K.,Liang Q.S.,Yang W.L.,Lou X.Y.,Chen J.,Feng M.J.,Jian J.B.,Zhang X.F.Luo G.B.,Jiang,Y.,Liu J.J.,Wang Z.B.,Sha Y.H.,Zhang B.R.,Wu H.J.,Tang D.Z.,Shen Q.H.,Xue P.Y.,Zou S.H.Wang X.J.,Liu X.,Wang F.M.,Yang Y.P.,An X.L.,Dong Z.Y.,Zhang K.P.,Zhang X.Q.,Luo M.C.,Dvorak J.,Tong YP.,Wang J.,Yang H.M.,Li Z.S.,Wang D.W.,Zhang A.Mand Wang J.,2013,Draft genome of the wheat A-genom progenitor Triticum urartu,Nature,496(7443):87-90
Liska A.J.,and Cassman K.G.,2008,Towards standardization o Life-Cycle metrics for biofuels:greenhouse gas emission mitigation and Net Energy Yield,Journal of Biobased Mate rials&Bioenergy,2(3):187-203
Litt A.,and Kramer E.M.,2010,The ABC model and the diversi fication of floral organ identity,Seminars in Cell&Devel opmental Biology,21(1):129-137
Liu L.S.,Wang C.X.,and Feng L.,eds.,2002,Plant Molecula Genetics,Science Press,Beijing,China,pp.426-434(刘良式,王春新,冯亮,编著,2002,植物分子遗传学,科学出版社,中国,北京,pp.426-434)
Liu S.J.,Zhu Q.,Xing X.J.,Wang B.Q.,Yu F.X.,and Zhou H.2014,Advanced research on male sterility of plant,Zhong guo Nongxue Tongbao(Chinese Agricultural Science Bul letin),30(34):46-50(刘淑娟,朱祺,幸学俊,王碧琴,余发新,周华,2014,植物雄性不育影响因素研究进展,中国农学通报,30(34):46-50)
Liu Z.,Yan J.P.,Li D.K.,Luo Q.,Yan Q.,Liu Z.B.,Y L.M.Wang J.M.,Li X.F.,and Yang Y.,2015,UDP-glucosyltrans ferase71C5,a major glucosyltransferase,mediates abscisi acid homeostasis in Arabidopsis,Plant Physiol.,167(4)1659-1670
Livak K.J.,and Schmittgen T.D.,2001,Analysis of relative gen expression data using Real-time quantitative PCR and th2-ΔΔCT(Delta Delta C(T))method,Methods,25(4):402-408
Murai K.,Takumi S.,Koga H.,and Ogihara Y.,2002,Pistillody homeotic transformation of stamens into pistil-like struc tures,caused by nuclear-cytoplasm interaction in wheat,Th Plant Journal,29(2):169-181
Nam J.,Depamphilis C.W.,Hong MMa H.,and Nei M.,2003Antiquity and evolution of the MADS-box gene family con trolling flower development in plants,Molecular Biology&Evolution,20(9):1435-1447
Pelaz S.,Ditta G.S,Baumann E.,Wisman E.,and Yanofsky M.F.2000,B and C floral organ identity functions require Sepal lata EPALLATA mads MADS-box genes,Nature,405(6783)200-203
Pelaz S.,Tapia-López R.,Alvarez-Buylla E.R.,and Yanofsky MF.,2001,Conversion of leaves into petals in Arabidopsis Current Biology,11(3):182-184
Peng Z.S.,2003,A new mutation in wheat producing three pistil in a floret,Journal of Agronomy and Crop Science,189(4)270-272
Peng Z.S.,Yang J.,Wei S.H.,and Zeng J.H.,2004,Characteriza tion of the common wheat(Triticum aestivum L.)mutation line producing three pistils in a floret,Hereditas,141(1)15-18
Prokopyk D.O.,and Ternovska T.K.,2011,Homeotic genes and their role in development of morphological traits in wheat Cytology and Genetics,45(1):41-54
Wang Y.W.,Peng Z.S.,Yang Z.J.,Wei S.H.,Liao M.L.,Zhao H.Yang H.,Yang Y.F.,and Wang Q.H.,2014,Cloning and ex pression analysis of Ta AP1-3 genes of three pistils mutation in wheat,Huabei Nongxuebao(Acta Agriculturae Boreali-Sinica),29(6):28-35(王育伟,彭正松,杨在君,魏淑红,廖明莉赵欢,杨会,杨宇凤,王清海,2014,小麦三雌蕊突变体Ta AP1-3基因的克隆及表达分析,华北农学报,29(6):28-35)
Yamaguchi T.,and Hirano H.Y.,2006,Function and diversifica tion of MADS-box genes in rice,The Scientific World Jour nal,6:1923-1932
Yang Y.F.,Peng Z.S.,Yang Z.J.,Wei S.H.,Liao M.L.,Wang YW.,Wang Q.H.,and Yang H.,2015,Cloning and expression analysis of Class-B MADS-box genes WPI1 and WPI2 in wheat,Henongxue Bao(Journal of Nuclear Agricultural Sci ences),29(2):221-228(杨宇凤,彭正松,杨在君,魏淑红廖明莉,王育伟,王清海,杨会,2015,小麦B类MADS-bo基因WPI1和WPI2的克隆及表达分析,核农学报,29(2)221-228)
Zhang Y.,and Liu Q.L.,2003,Proceedings on molecular mecha nism of plant flower development,Zhiwuxue Tongbao(Chi nese Bulletin of Bontany),20(5):589-601(张云,刘青林2003,植物花发育的分子机理研究进展,植物学通报,20(5):589-601)
Zhang Y.,Sui Z.H.,Ding H.Y.,and Zhong J.,2011,Cloning and analysis of the UDP-Glucose pyrophosphoryiase(UGPase gene of Gracilaria leimaneiformis(Rhodophyta)and correla tion between gene expression and agar synthesis,Zhongguo
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