水稻金属硫蛋白基因(MT2bL)启动子的克隆与表达分析
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摘要
金属硫蛋白(metallothionein,MT)富含半胱氨酸,而且能够结合多种金属离子,调控细胞内金属代谢的平衡。本研究根据水稻在减数分裂期、开花期和开花后5 d的基因芯片数据结果,从水稻品种黎榆B(O.sativa L.ssp.japonica C.V.Liyu B)基因组中克隆得到了一个植物MT-2类基因Metallothionein2b-Like(Os MT2b L)的启动子序列,序列全长2 063 bp。利用植物启动子区域顺式作用元件数据库(PLACE)分析表明该启动子序列含有5个CURE(copper-response element)元件(GTAC)以及多个其它顺式作用元件。构建了多个启动子融合表达载体(p Ca MV35S::GUS,p MT2b L::GUS,p Ubi1::GUS,p CYC::GUS和p ACT1::GUS),并通过根癌农杆菌介导转化水稻愈伤组织获得了转基因植株。GUS组织染色结果分析表明,Os MT2b L基因启动子在水稻幼苗期的胚根、胚芽和胚芽鞘中具有高水平的GUS表达活性,特别是在胚根根尖和胚芽顶端的GUS表达活性较高;在减数分裂期的植株叶片、颖壳和开花10 d后的未成熟种子中GUS表达活性也较高;GUS蛋白荧光定量分析结果表明,Os MT2b L基因启动子在水稻叶片中驱动GUS基因表达的活性比Ca MV35S基因启动子高出3倍以上。
Metallothionein(MT) is rich in cysteine, and can combine a variety of metalions to regulate the balance of intracellular metal metabolism. In this study, we cloned Oryza sativa Metallothionein2 b-Like(Os MT2 b L)promoter sequence with the length of 2 063 bp based on microarray results at meiotic, flowering and 5 days after pollination(DAP) stages from Liyu B(O. sativa L. ssp. japonica C.V.). We used plant promoter region cis-element database PLACE to analyze and found that these regulatory cis-elements include 5 copper-response elements(CURE) and other cis-elements. We constructed a number of promoter fusion expression vectors(p Ca MV35 S::GUS, pMT2bL::GUS, p Ubi1::GUS, p CYC::GUS and p ACT1::GUS) and obtained transgenic plants by Agrobacterium tumefaciens-mediated transformation metho d. The results of GUS histochemical assays showed that Os MT2bL gene promoter had high level of GUS expression in radicle, embryo and coleoptile at rice seedling stage, especially in the tip of radicle and the top of embryo. The level of GUS expression was also high in the leaf, panicles, and immature seeds of 10 days after pollination at the meiosis stage. The results of GUS protein fluorescence quantitative analysis showed that the expression level of GUS gene drove by Os MT2bL gene promoter was three times higher than Ca MV35 S gene promoter in rice leaves.
Metallothionein(MT) is rich in cysteine, and can combine a variety of metalions to regulate the balance of intracellular metal metabolism. In this study, we cloned Oryza sativa Metallothionein2 b-Like(Os MT2 b L)promoter sequence with the length of 2 063 bp based on microarray results at meiotic, flowering and 5 days after pollination(DAP) stages from Liyu B(O. sativa L. ssp. japonica C.V.). We used plant promoter region cis-element database PLACE to analyze and found that these regulatory cis-elements include 5 copper-response elements(CURE) and other cis-elements. We constructed a number of promoter fusion expression vectors(p Ca MV35 S::GUS, pMT2bL::GUS, p Ubi1::GUS, p CYC::GUS and p ACT1::GUS) and obtained transgenic plants by Agrobacterium tumefaciens-mediated transformation metho d. The results of GUS histochemical assays showed that Os MT2bL gene promoter had high level of GUS expression in radicle, embryo and coleoptile at rice seedling stage, especially in the tip of radicle and the top of embryo. The level of GUS expression was also high in the leaf, panicles, and immature seeds of 10 days after pollination at the meiosis stage. The results of GUS protein fluorescence quantitative analysis showed that the expression level of GUS gene drove by Os MT2bL gene promoter was three times higher than Ca MV35 S gene promoter in rice leaves.
引文
Abdullah S.N.A.,Cheah S.C.,and Murphy D.J.,2002,Isolation and characterisation of two divergent type 3 metallothioneins from oil palm,Elaeis guineensis,Plant Physiology and Biochemistry,40(3):255-263
Akashi K.,Nishimura N.,Ishida Y.,and Yokota A.,2004,Potent hydroxyl radical-scavenging activity of drought-induced type-2 metallothionein in wild watermelon,Biochem.Biophys.Res.Commun.,323(1):72-78
Butt A.,Mousley C.,Morris K.,Beynon J.,Can C.,Holub E.,Greenberg J.T.,and Buchanan-Wollaston V.,1998,Differential expression of a senescence enhanced metallothionein gene in Arabidopsis in response to isolates of Peronospora parasitica and Pseudomonas syringae,Plant J.,16(2):209-221
Chakravarthy S.,Tuori R.P.,D'Ascenzo M.D.,Fobert P.R.,Després C.,and Martin G.B.,2003,The tomato transcription factor Pti4 regulates defense-related gene expression via GCC box and non-GCC box cis-elements,The Plant Cell,15(12):3033-3050
Chatthai M.,Osusky M.,Osuska L.,Yevtushenko D.,and Misra S.,2004,Functional analysis of a Douglas-fir metallothionein-like gene promoter:transient assays in zygotic and somatic embryos and stable transformation in transgenic tobacco,Planta,220(1):118-128
Chen C.,and Chen Z.,2002,Potentiation of developmentally regulated plant defense response by At WRKY18,a pathogeninduced Arabidopsis transcription factor,Plant Physiol.,129(2):706-716
De Framond A.J.,1991,A metallothionein-like gene from maize(Zea mays)cloning and characterization,FEBS Lett.,290(1):103-106
Ellerstr觟m M.,St覽lberg K.,Ezcurra I.,and Rask L.,1996,Functional dissection of a napin gene promoter:identification of promoter elements required for embryo and endosperm-specific transcription,Plant Mol.Bio.,32(6):1019-1027
Evans I.M.,Gatehouse L.N.,Gatehouse J.A.,Robinson N.J.,and Croy R.R.,1990,A gene from pea(Pisum sativum L.)with homology to metallothionein genes,FEBS Lett.,262(1):29-32
Fehlberg V.,Vieweg M.F.,Dohmann E.M.N.,Hohnjec N.,Pühler A.,Perlick A.M.,and Küster H.,2005,The promoter of the leghaemoglobin gene Vf Lb29:functional analysis and identification of modules necessary for its activation in the infected cells of root nodules and in the arbuscule-containing cells of mycorrhizal roots,J.Exp.Bot.,56(413):799-806
Filichkin S.A.,Leonard J.M.,Monteros A.,Liu P.P.,and Nonogaki H.,2004,A novel endo-β-mannanase gene in tomato Le MAN5 is associated with anther and pollen development,Plant Physiol.,134(3):1080-1087
Fiume E.,Christou P.,GianìS.,and Breviario D.,2004,Introns are key regulatory elements of rice tubulin expression,Planta,218(5):693-703
Fordham-Skelton A.P.,Lilley C.,Urwin P.E.,and Robinson N.J.,1997,GUS expression in Arabidopsis directed by 5'-regions of the pea metallothionein-like gene Ps MTA,Plant Mol.Biol.,34(4):659-668
Fukuzawa H.,Yu L.H.,Umeda-Hara C.,Tagawa M.,and Uchimiya H.,2004,The rice metallothionein gene promoter does not direct foreign gene expression in seed endosperm,Plant Cell Rep.,23(4):231-235
Gallagher S.R.,ed.,1992,GUS protocols:using the GUS gene as a reporter of gene expression,Academic Press,New York,USA,pp.221
Goda H.,Sawa S.,Asami T.,Fujioka S.,Shimada Y.,and Yoshida S.,2004,Comprehensive comparison of auxin-regulated and brassinosteroid-regulated genes in Arabidopsis,Plant Physiol.,134(4):1555-1573
Guo W.J.,Bundithya W.,and Goldsbrough P.B.,2003,Characterization of the Arabidopsis metallothionein gene family:tissue-specific expression and induction during senescence and in response to copper,The New Phytologist,159(2):369-381
Hartmann U.,Sagasser M.,Mehrtens F.,Stracke R.,and Weisshaar B.,2005,Differential combinatorial interactions of cis-acting elements recognized by R2R3-MYB,BZIP,and BHLH factors control light-responsive and tissue-specific activation of phenylpropanoid biosynthesis genes,Plant Mol.Biol.,57(2):155-171
Hattori T.,Terada T.,and Hamasuna S.,1995,Regulation of the Osem gene by abscisic acid and the transcriptional activator VP1:analysis of cis-acting promoter elements required for regulation by abscisic acid and VP1,Plant J.,7(6):913-925
Hiei Y.,Ohta S.,Komari T.,and Kumashiro T.,1994,Efficient transformation of rice(Oryza sativa L.)mediated by Agrobacterium and sequence analysis of the boundaries of the T-DNA,Plant J.,6(2):271-282
Hwang Y.S.,Karrer E.E.,Thomas B.R.,Chen L.,and Rodriguez R.L.,1998,Three cis-elements required for riceα-amylase Amy3D expression during sugar starvation,Plant Mol.Biol.,36(3):331-341
Izawa T.,Foster R.,Nakajima M.,Shimamoto K.,and Chua N.H.,1994,The rice b ZIP transcriptional activator RITA-1 is highly expressed during seed development,Plant Cell,6(9):1277-1287
Jefferson R.A.,1987,Assaying chimeric genes in plants:the GUS gene fusion system,Plant Molecular Biology Reporter,5(4):387-405
Karin M.,Haslinger A.,Holtgreve H.,Richards R.I.,Krauter P.,Westphal H.M.,and Beato M.,1984,Characterization of DNA sequences through which cadmium and glucocorticoid hormones induce human metallothionein-IIA gene,Nature,308(5959):513-519
Kasuga M.,Liu Q.,Miura S.,Yamaguchi-Shinozaki K.,and Shinozaki K.,1999,Improving plant drought,salt,and freezing tolerance by gene transfer of a single stress-inducible transcription factor,Nat.Biotechnol.,17(3):287-291
Klemsdal S.S.,Hughes W.,L觟nneborg A.,Aalen R.B.,and Olsen O.A.,1991,Primary structure of a novel barley gene differentially expressed in immature aleurone layers,Mol.Gen.Genet,228(1):9-16
Kropat J.,Tottey S.,Birkenbihl R.P.,Depège N.,Huijser P.,and Merchant S.,2005,A regulator of nutritional copper signaling in Chlamydomonas is an SBP domain protein that recognizes the GTAC core of copper response element,Proc.Natl.Acad.Sci.USA,102(51):18730-18735
Lee D.S.,Chen L.J.,Li C.Y.,Liu Y.,Tan X.L.,Lu B.R.,Li J.,Gan S.X.,Kang S.G.,Suh H.S.,and Zhu Y.,2013,The Bsister MADS gene FST determines ovule patterning and development of the zygotic embryo and endosperm,PLo SOne,8(3):e58748
Liu J.,Hara C.,Umeda M.,Zhao Y.,Okita T.W.,and Uchimiya H.,1995,Analysis of randomly isolated c DNAs from developing endosperm of rice(Oryza sativa L.):evaluation of expressed sequence tags,and expression levels of m RNAs,Plant Mol.Biol.,29(4):685-689
Liu W.X.,Liu H.L.,Chai Z.J.,Xu X.P.,Song Y.R.,and Qu L.Q.,2010,Evaluation of seed storage-protein gene 5'-untranslated regions in enhancing gene expression in transgenic rice seed,Theor.Appl.Genet.,121(7):1267-1274
Luo H.,Song F.,Goodman R.M.,and Zheng Z.,2005,Up-regu lation of Os BIHD1,a rice gene encoding BELL homeodomain transcriptional factor,in disease resistance responses,Plant Biol.,7(5):459-468
Lv S.,Gu H.,Yuan X.,Wang X.,Wu A.M.,Qu L.,and Liu J.Y.,2007,The GUS reporter-aided analysis of the promoter activities of a rice metallothionein gene reveals different regulatory regions responsible for tissue-specific and inducible expression in transgenic Arabidopsis,Transgenic Res.,16(2):177-191
Murray M.G.,and Thompson W.F.,1980,Rapid isolation of high molecular weight plant DNA,Nucleic Acids Research,8(19):4321-4326
Nishiuchi T.,Hamada T.,Kodama H.,and Iba K.,1997,Wounding changes the spatial expression pattern of the Arabidopsis plastid omega-3 fatty acid desaturase gene(FAD7)through different signal transduction pathways,The Plant Cell,9(10):1701-1712
Park H.C.,Kim M.L.,Kang Y.H.,Jeon J.M.,Yoo J.H.,Kim M.C.,Park C.Y.,Jeong J.C.,Moon B.C.,Lee J.H.,Yoon H.W.,Lee S.H.,Chung W.S.,Lim C.O.,Lee S.Y.,Hong J.C.,and Cho M.J.,2004,Pathogen-and Na Cl-induced expression of the SCa M-4 promoter is mediated in part by a GT-1 box that interacts with a GT-1-like transcription factor,Plant Physiol.,135(4):2150-2161
Robinson N.B.,Chi E.Y.,and Robertson H.T.,1983,Ventilation-perfusion relationships after hemorrhage and resuscitation:an inert gas analysis,J.Appl.Physiol.Respir.Environ.Exerc.Physiol.,54(4):1131-1140
Rubio-Somoza I.,Martinez M.,Abraham Z.,Diaz I.,and Carbonero P.,2006,Ternary complex formation between Hv M-YBS3 and other factors involved in transcriptional control in barley seeds,Plant J.,47(2):269-281
Seguin C.,and Hamer D.H.,1987,Regulation in vitro of metallothionein gene binding factors,Science,235(4794):1383-1387
Shirsat A.,Wilford N.,Croy R.,and Boulter D.,1989,Sequences responsible for the tissue specific promoter activity of a pea legumin gene in tobacco,Mol.Gen.Genet.,215(2):326-331
Solano R.,Nieto C.,Avila J.,Ca觡as L.,Diaz I.,and Paz-Ares J.,1995,Dual DNA binding specificity of a petal epidermis-spe cific MYB transcription factor(MYB.Ph3)from Petunia hybrida,EMBO J.,14(8):1773-1784
Sutoh K.,and Yamauchi D.,2003,Two cis-acting elements necessary and sufficient for gibberellin-upregulated proteinase expression in rice seeds,Plant J.,34(5):635-645
Wenkel S.,Turck F.,Singer K.,Gissot L.,Gourrierec L.J.,Samach A.,and Coupland G.,2006,CONSTANS and the CCAATbox binding complex share a functionally important domain and interact to regulate flowering of Arabidopsis,Plant Cell,18(11):2971-2984
Whitelaw C.A.,Le Huquet J.A.,Thurman D.A.,and Tomsett A.B.,1997,The isolation and characterization of typeⅡmetallothionein-like genes from tomato(Lycopersicon esculentum L.),Plant Mol.Biol.,33(3):503-511
Wong H.L.,Sakamoto T.,Kawasaki T.,Umemura K.,and Shimamoto K.,2004,Down-regulation of metallothionein,a reactive oxygen scavenger,by the small GTPase Os Rac1 in rice,Plant Physiol.,135(3):1447-1456
Wu C.S.,Chen D.Y.,Chang C.F.,Li M.J.,Hung K.Y.,Chen L.J.,and Chen P.W.,2014,The promoter and the 5'-untranslated region of rice metallothionein Os MT2b gene are capable of directing high-level gene expression in germinated rice embryos,Plant Cell Rep.,33(5):793-806
Yu L.H.,Liu J.Y.,Umeda M.,Uchimiya H.,and Zhao N.M.,2000,Cloning and sequence characteristics of the genomic gene of a rice metallothionein,Chinese Science Bulletin,45(2):153-156
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Yuan J.,Chen D.,Ren Y.J.,Zhang X.L.,and Zhao J.,2008,Characteristic and expression analysis of a metallothionein gene,Os MT2b,down-regulated by cytokinin suggests functions in root development and seed embryo germination of rice,Plant Physiol.,146(4):1637-1650
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Zhao B.,2007,Expression and functional analysis of Bj MT2,a metallothionein type 2 from Brassica juncea,in Tobacco plant,Thesis for M.S.,Northeast Forestry University,Supervisor:An Z.G.,pp.1-49(赵博,2007,植物金属硫蛋白2型在烟草中的表达特性,硕士学位论文,东北林业大学,导师:安志刚,pp.1-49)
Zhou G.K.,Xu Y.F.,and Liu J.Y.,2005,Characterization of a rice class II metallothionein gene:tissue expression patterns and induction in response to abiotic factors,J.Plant Physiol.,162(6):686-696
Akashi K.,Nishimura N.,Ishida Y.,and Yokota A.,2004,Potent hydroxyl radical-scavenging activity of drought-induced type-2 metallothionein in wild watermelon,Biochem.Biophys.Res.Commun.,323(1):72-78
Butt A.,Mousley C.,Morris K.,Beynon J.,Can C.,Holub E.,Greenberg J.T.,and Buchanan-Wollaston V.,1998,Differential expression of a senescence enhanced metallothionein gene in Arabidopsis in response to isolates of Peronospora parasitica and Pseudomonas syringae,Plant J.,16(2):209-221
Chakravarthy S.,Tuori R.P.,D'Ascenzo M.D.,Fobert P.R.,Després C.,and Martin G.B.,2003,The tomato transcription factor Pti4 regulates defense-related gene expression via GCC box and non-GCC box cis-elements,The Plant Cell,15(12):3033-3050
Chatthai M.,Osusky M.,Osuska L.,Yevtushenko D.,and Misra S.,2004,Functional analysis of a Douglas-fir metallothionein-like gene promoter:transient assays in zygotic and somatic embryos and stable transformation in transgenic tobacco,Planta,220(1):118-128
Chen C.,and Chen Z.,2002,Potentiation of developmentally regulated plant defense response by At WRKY18,a pathogeninduced Arabidopsis transcription factor,Plant Physiol.,129(2):706-716
De Framond A.J.,1991,A metallothionein-like gene from maize(Zea mays)cloning and characterization,FEBS Lett.,290(1):103-106
Ellerstr觟m M.,St覽lberg K.,Ezcurra I.,and Rask L.,1996,Functional dissection of a napin gene promoter:identification of promoter elements required for embryo and endosperm-specific transcription,Plant Mol.Bio.,32(6):1019-1027
Evans I.M.,Gatehouse L.N.,Gatehouse J.A.,Robinson N.J.,and Croy R.R.,1990,A gene from pea(Pisum sativum L.)with homology to metallothionein genes,FEBS Lett.,262(1):29-32
Fehlberg V.,Vieweg M.F.,Dohmann E.M.N.,Hohnjec N.,Pühler A.,Perlick A.M.,and Küster H.,2005,The promoter of the leghaemoglobin gene Vf Lb29:functional analysis and identification of modules necessary for its activation in the infected cells of root nodules and in the arbuscule-containing cells of mycorrhizal roots,J.Exp.Bot.,56(413):799-806
Filichkin S.A.,Leonard J.M.,Monteros A.,Liu P.P.,and Nonogaki H.,2004,A novel endo-β-mannanase gene in tomato Le MAN5 is associated with anther and pollen development,Plant Physiol.,134(3):1080-1087
Fiume E.,Christou P.,GianìS.,and Breviario D.,2004,Introns are key regulatory elements of rice tubulin expression,Planta,218(5):693-703
Fordham-Skelton A.P.,Lilley C.,Urwin P.E.,and Robinson N.J.,1997,GUS expression in Arabidopsis directed by 5'-regions of the pea metallothionein-like gene Ps MTA,Plant Mol.Biol.,34(4):659-668
Fukuzawa H.,Yu L.H.,Umeda-Hara C.,Tagawa M.,and Uchimiya H.,2004,The rice metallothionein gene promoter does not direct foreign gene expression in seed endosperm,Plant Cell Rep.,23(4):231-235
Gallagher S.R.,ed.,1992,GUS protocols:using the GUS gene as a reporter of gene expression,Academic Press,New York,USA,pp.221
Goda H.,Sawa S.,Asami T.,Fujioka S.,Shimada Y.,and Yoshida S.,2004,Comprehensive comparison of auxin-regulated and brassinosteroid-regulated genes in Arabidopsis,Plant Physiol.,134(4):1555-1573
Guo W.J.,Bundithya W.,and Goldsbrough P.B.,2003,Characterization of the Arabidopsis metallothionein gene family:tissue-specific expression and induction during senescence and in response to copper,The New Phytologist,159(2):369-381
Hartmann U.,Sagasser M.,Mehrtens F.,Stracke R.,and Weisshaar B.,2005,Differential combinatorial interactions of cis-acting elements recognized by R2R3-MYB,BZIP,and BHLH factors control light-responsive and tissue-specific activation of phenylpropanoid biosynthesis genes,Plant Mol.Biol.,57(2):155-171
Hattori T.,Terada T.,and Hamasuna S.,1995,Regulation of the Osem gene by abscisic acid and the transcriptional activator VP1:analysis of cis-acting promoter elements required for regulation by abscisic acid and VP1,Plant J.,7(6):913-925
Hiei Y.,Ohta S.,Komari T.,and Kumashiro T.,1994,Efficient transformation of rice(Oryza sativa L.)mediated by Agrobacterium and sequence analysis of the boundaries of the T-DNA,Plant J.,6(2):271-282
Hwang Y.S.,Karrer E.E.,Thomas B.R.,Chen L.,and Rodriguez R.L.,1998,Three cis-elements required for riceα-amylase Amy3D expression during sugar starvation,Plant Mol.Biol.,36(3):331-341
Izawa T.,Foster R.,Nakajima M.,Shimamoto K.,and Chua N.H.,1994,The rice b ZIP transcriptional activator RITA-1 is highly expressed during seed development,Plant Cell,6(9):1277-1287
Jefferson R.A.,1987,Assaying chimeric genes in plants:the GUS gene fusion system,Plant Molecular Biology Reporter,5(4):387-405
Karin M.,Haslinger A.,Holtgreve H.,Richards R.I.,Krauter P.,Westphal H.M.,and Beato M.,1984,Characterization of DNA sequences through which cadmium and glucocorticoid hormones induce human metallothionein-IIA gene,Nature,308(5959):513-519
Kasuga M.,Liu Q.,Miura S.,Yamaguchi-Shinozaki K.,and Shinozaki K.,1999,Improving plant drought,salt,and freezing tolerance by gene transfer of a single stress-inducible transcription factor,Nat.Biotechnol.,17(3):287-291
Klemsdal S.S.,Hughes W.,L觟nneborg A.,Aalen R.B.,and Olsen O.A.,1991,Primary structure of a novel barley gene differentially expressed in immature aleurone layers,Mol.Gen.Genet,228(1):9-16
Kropat J.,Tottey S.,Birkenbihl R.P.,Depège N.,Huijser P.,and Merchant S.,2005,A regulator of nutritional copper signaling in Chlamydomonas is an SBP domain protein that recognizes the GTAC core of copper response element,Proc.Natl.Acad.Sci.USA,102(51):18730-18735
Lee D.S.,Chen L.J.,Li C.Y.,Liu Y.,Tan X.L.,Lu B.R.,Li J.,Gan S.X.,Kang S.G.,Suh H.S.,and Zhu Y.,2013,The Bsister MADS gene FST determines ovule patterning and development of the zygotic embryo and endosperm,PLo SOne,8(3):e58748
Liu J.,Hara C.,Umeda M.,Zhao Y.,Okita T.W.,and Uchimiya H.,1995,Analysis of randomly isolated c DNAs from developing endosperm of rice(Oryza sativa L.):evaluation of expressed sequence tags,and expression levels of m RNAs,Plant Mol.Biol.,29(4):685-689
Liu W.X.,Liu H.L.,Chai Z.J.,Xu X.P.,Song Y.R.,and Qu L.Q.,2010,Evaluation of seed storage-protein gene 5'-untranslated regions in enhancing gene expression in transgenic rice seed,Theor.Appl.Genet.,121(7):1267-1274
Luo H.,Song F.,Goodman R.M.,and Zheng Z.,2005,Up-regu lation of Os BIHD1,a rice gene encoding BELL homeodomain transcriptional factor,in disease resistance responses,Plant Biol.,7(5):459-468
Lv S.,Gu H.,Yuan X.,Wang X.,Wu A.M.,Qu L.,and Liu J.Y.,2007,The GUS reporter-aided analysis of the promoter activities of a rice metallothionein gene reveals different regulatory regions responsible for tissue-specific and inducible expression in transgenic Arabidopsis,Transgenic Res.,16(2):177-191
Murray M.G.,and Thompson W.F.,1980,Rapid isolation of high molecular weight plant DNA,Nucleic Acids Research,8(19):4321-4326
Nishiuchi T.,Hamada T.,Kodama H.,and Iba K.,1997,Wounding changes the spatial expression pattern of the Arabidopsis plastid omega-3 fatty acid desaturase gene(FAD7)through different signal transduction pathways,The Plant Cell,9(10):1701-1712
Park H.C.,Kim M.L.,Kang Y.H.,Jeon J.M.,Yoo J.H.,Kim M.C.,Park C.Y.,Jeong J.C.,Moon B.C.,Lee J.H.,Yoon H.W.,Lee S.H.,Chung W.S.,Lim C.O.,Lee S.Y.,Hong J.C.,and Cho M.J.,2004,Pathogen-and Na Cl-induced expression of the SCa M-4 promoter is mediated in part by a GT-1 box that interacts with a GT-1-like transcription factor,Plant Physiol.,135(4):2150-2161
Robinson N.B.,Chi E.Y.,and Robertson H.T.,1983,Ventilation-perfusion relationships after hemorrhage and resuscitation:an inert gas analysis,J.Appl.Physiol.Respir.Environ.Exerc.Physiol.,54(4):1131-1140
Rubio-Somoza I.,Martinez M.,Abraham Z.,Diaz I.,and Carbonero P.,2006,Ternary complex formation between Hv M-YBS3 and other factors involved in transcriptional control in barley seeds,Plant J.,47(2):269-281
Seguin C.,and Hamer D.H.,1987,Regulation in vitro of metallothionein gene binding factors,Science,235(4794):1383-1387
Shirsat A.,Wilford N.,Croy R.,and Boulter D.,1989,Sequences responsible for the tissue specific promoter activity of a pea legumin gene in tobacco,Mol.Gen.Genet.,215(2):326-331
Solano R.,Nieto C.,Avila J.,Ca觡as L.,Diaz I.,and Paz-Ares J.,1995,Dual DNA binding specificity of a petal epidermis-spe cific MYB transcription factor(MYB.Ph3)from Petunia hybrida,EMBO J.,14(8):1773-1784
Sutoh K.,and Yamauchi D.,2003,Two cis-acting elements necessary and sufficient for gibberellin-upregulated proteinase expression in rice seeds,Plant J.,34(5):635-645
Wenkel S.,Turck F.,Singer K.,Gissot L.,Gourrierec L.J.,Samach A.,and Coupland G.,2006,CONSTANS and the CCAATbox binding complex share a functionally important domain and interact to regulate flowering of Arabidopsis,Plant Cell,18(11):2971-2984
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