CMO启动子盐诱导功能元件鉴定及转录因子分离
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摘要
甜菜碱是植物体内的一种无毒渗透保护剂,它的积累使代谢中的许多重要酶类在渗透胁迫下能保持活性,保证植物正常生长。胆碱单加氧酶(CMO)是指导合成甜菜碱的关键酶。CMO基因的表达受其上游启动子及转录因子的调控,转录因子与启动子上的顺式作用元件相互作用,促进下游基因的表达。本文通过对CMO启动子盐诱导元件及与其结合的转录因子进行研究,分析CMO基因转录调控的机制。
本论文通过凝胶迁移分析实验找到了CMO启动子区段中盐诱导功能元件,以盐诱导功能元件为诱饵,通过酵母单杂交实验分离得到了可能与之结合的转录因子,主要结果如下:
1、用Genomatix软件对已知的辽宁碱蓬CMO启动子序列(-267～+128 bp)进行功能元件分析,发现在该序列上存在11个与非生物胁迫相关的顺式作用元件。从中选定了4个与盐诱导表达相关的元件:W-Box、Salt/drought responsive element、GT-1 cis-element和MYB元件用于凝胶迁移分析实验。
2、以选定的4个元件为核心序列设计探针,与辽宁碱蓬叶片核蛋白结合,经凝胶阻滞分析,发现有2条探针具有阻滞信号,初步认为Salt/drought responsive elements和MYB元件是CMO启动子中盐诱导相关的功能元件。
3、从获得的2个盐诱导功能元件中选取Salt/drought responsive element作为诱饵,将该元件的三串联体构建到酵母菌株中获得诱饵酵母菌株,同时构建辽宁碱蓬叶片cDNA文库,经酵母单杂交实验,筛选出阳性克隆,经测序及序列比对分析,获得了1个与Salt/drought responsive element相互作用的转录因子,该转录因子可能是PAPA-1- like family protein / zinc finger (HIT) family protein。
Betaine is a non-toxic penetration of protective agents in plants. Its accumulation makes many important enzymes of metabolism keep activity under osmotic stress, and protect the normal growth of plants. Choline monooxygenase (CMO) is the key enzyme for betaine synthesis. CMO gene expression is regulated by upstream promoter and transcription factors. The interaction between transcription factors and promoter cis-acting elements promote the expression of genes downstream. In this paper, the salt-related cis-acting elements in CMO promoter and their binding transcription factors were studied, and transcriptional regulation mechanism of CMO gene was analyzed.
This thesis finds salt-related cis-acting elements in CMO promoter by electrophoretic mobility shift assay(EMSA), and transcription factors binding to the cis-acting elements were isolated by yeast one-hybrid experiments. The main results are as follows:
1. CMO promoter sequence (-267～+128 bp) were analyzed using Genomatix Online software. 11 abiotic stress-related cis-acting elements were found in this sequence. 4 cis-acting elements:MYB element, W-box, GT-box and Salt/drought responsive element, were selected for EMSA.
2. 4 probes were designed using the selected cis-acting elements as the core sequence. The interaction between probes and nuclear proteins from Suaeda liaotungensis leaves was detected by EMSA, Blocking signals were found in 2 probes. It is concluded that MYB element and Salt/drought responsive element may be the salt-induced cis-acting elements in CMO promoter.
3. Using the Salt/drought responsive element as bait, a transcription factor was isolated from the cDNA library of Suaeda liaotungensis by yeast one-hybrid experiments, It is a PAPA-1-like family protein / zinc finger (HIT) family protein.
本论文通过凝胶迁移分析实验找到了CMO启动子区段中盐诱导功能元件,以盐诱导功能元件为诱饵,通过酵母单杂交实验分离得到了可能与之结合的转录因子,主要结果如下:
1、用Genomatix软件对已知的辽宁碱蓬CMO启动子序列(-267～+128 bp)进行功能元件分析,发现在该序列上存在11个与非生物胁迫相关的顺式作用元件。从中选定了4个与盐诱导表达相关的元件:W-Box、Salt/drought responsive element、GT-1 cis-element和MYB元件用于凝胶迁移分析实验。
2、以选定的4个元件为核心序列设计探针,与辽宁碱蓬叶片核蛋白结合,经凝胶阻滞分析,发现有2条探针具有阻滞信号,初步认为Salt/drought responsive elements和MYB元件是CMO启动子中盐诱导相关的功能元件。
3、从获得的2个盐诱导功能元件中选取Salt/drought responsive element作为诱饵,将该元件的三串联体构建到酵母菌株中获得诱饵酵母菌株,同时构建辽宁碱蓬叶片cDNA文库,经酵母单杂交实验,筛选出阳性克隆,经测序及序列比对分析,获得了1个与Salt/drought responsive element相互作用的转录因子,该转录因子可能是PAPA-1- like family protein / zinc finger (HIT) family protein。
Betaine is a non-toxic penetration of protective agents in plants. Its accumulation makes many important enzymes of metabolism keep activity under osmotic stress, and protect the normal growth of plants. Choline monooxygenase (CMO) is the key enzyme for betaine synthesis. CMO gene expression is regulated by upstream promoter and transcription factors. The interaction between transcription factors and promoter cis-acting elements promote the expression of genes downstream. In this paper, the salt-related cis-acting elements in CMO promoter and their binding transcription factors were studied, and transcriptional regulation mechanism of CMO gene was analyzed.
This thesis finds salt-related cis-acting elements in CMO promoter by electrophoretic mobility shift assay(EMSA), and transcription factors binding to the cis-acting elements were isolated by yeast one-hybrid experiments. The main results are as follows:
1. CMO promoter sequence (-267～+128 bp) were analyzed using Genomatix Online software. 11 abiotic stress-related cis-acting elements were found in this sequence. 4 cis-acting elements:MYB element, W-box, GT-box and Salt/drought responsive element, were selected for EMSA.
2. 4 probes were designed using the selected cis-acting elements as the core sequence. The interaction between probes and nuclear proteins from Suaeda liaotungensis leaves was detected by EMSA, Blocking signals were found in 2 probes. It is concluded that MYB element and Salt/drought responsive element may be the salt-induced cis-acting elements in CMO promoter.
3. Using the Salt/drought responsive element as bait, a transcription factor was isolated from the cDNA library of Suaeda liaotungensis by yeast one-hybrid experiments, It is a PAPA-1-like family protein / zinc finger (HIT) family protein.
引文
[1] Naomi S S , Ichiro M. Constitutive Promoters Available for transgene expression instead of CaMV35S RNA promoter:A rabi dopsis promoters of tryptophan synthase protein subunitand phytochrome B. Plant Biotechnology, 2002, 19 (1): 19-26.
[2] Seagull R W. The plant cytoskeleton.Crit Rev Plant Sci,1989, 8 : 131-167.
[3]张树珍,杨本鹏,刘飞虎,花特异表达启动子PchsA的克隆及其序列分析。农业生物技术学报,2002。10(2): 116-119。
[4] Narusaka Y, Nakashima K, Shinwari Z K, Sakuma Y, Furihata T ,Abe H , Narusaka M ,Shinozaki K ,Yamaguchi Shinozaki K. Interaction between two cis-acting elements ,ABRE and DRE ,in ABA dependent expression of A rabidopsis rd29A gene inresponse to dehydration and high salinity stresses. Plant J,2003, 34(2) : 137-148.
[5] Brown , A P , Dunn M A , Goddard N J , Hughes M A. Identification of a novel low-temperature-response element in the promoter of the barley ( Hordeum v ulgare L. ) gene blt101. 1. Planta,2001,213 (5) : 770-780.
[6] Xue GP. The DNA-binding activity of an AP2 transcriptional activator HvCBF2 involved in regulation of low-temperature responsive genes in barley is modulated by temperature. Plant J, 2003, 33:373-383.
[7] Stracke R, Werber M, Weisshaar B.The R2R3, MYB gene family in Arabidopsis thaliana. Curr Opin Plant Biol. 2001, (4): 447-456.
[8] Yamagata H, Yonesu K, Hirata A and Aizono Y.TGTCACA motif is a novel cis-regulatory enhancer element involved in fruit-specific expression of the cucumisin gene.J Biol Chem 2002, 277:11582-11590.
[9]杨英军,周鹏。番木瓜proteinase omega基因启动子的克隆及功能初步研究。云南植物研究, 2005,27 (5): 545-551。
[10] YIN Y, BEACHY R N.The regulatory regions of the rice tun-gro bacilliform virus promoter and interacting nuclear factors in rice (Oryza sativa L). Plant Journal, 1995, 7 (6): 969 - 980.
[11] Yamagata H, Yonesu K, Hirata A and Aizono Y.TGTCACA motif is a novel cis-regulatory enhancer element involved in fruit-specific expression of the cucumisin gene. J Biol Chem 2002, 277:11582-11590.
[12] Li JJ,Herskowitz1.IsolationofORC6,a component of the yeast origin recognition complex by a one-hybrid system. Science 1993;262:1870-1873.
[13] Delaney S K, Orford S J, Martin Harr ISM, et al. The fiber specificity of the cotton FSltp4 gene promoter is regulated by an AT rich p romoter region and the AT-Hook transcrip tion factor GhAT1. Plant Cell Physiology, 2007, 48 (10): 1426 - 1437.
[14] Rouster J.,Leah R.,Mundy J.,and Cameron-Mills V.Identification of a methyl jasmonate-responsive region in the promoter of a lipoxygenase 1 gene expressed in barley grain.Plant J 1997,11:513-523.
[15]张海文,谢丙炎,卢向阳,等.拟南芥防卫基因PDF1.2启动子中GCC盒是应答茉莉素反应必要的顺式作用元件。科学通报,2004,49(23): 2444-2448。
[16]吴纯清,张兴国,梁国鲁等。拟南芥AtRd29A启动子在烟草中的表达特性。西南农业学报, 2008,21(4):1045–1047。
[17] Rai M, He CK, Wu R.Comparative functional analysis of three abiotic stress-inducible promoters in transgenic rice.Transgenic Res, 2009, 18(5): 787–799.
[18]刘峻,朱国璋,解芳等。ACTA B IOCH IM ICA et BIOPHYSICA SIN ICAVo l. 29,1997,No.6:597-604。
[19] Maz n G, Lucena J M, Campoy S. LexA-binding sequences in Gram-positive and cyanobacteria are closely related [J].Mol Gen Genomics, 2004, 271: 40-49.
[20] Braunstein M, Rose AB, Holmes SG, Allis CD, Broach JR.Transcriptional silencing in yeast is associated with reduced nucleosome acetylation. Genes Dev, 1993, 7(4):592 ?6 04.
[21]张莉萍。运用染色质免疫沉淀技术分析基因表达转录调控方法的建立及初步应用。中华检验医学杂志,2006,29(5):449-451。
[22] Schnekenburger M, Talaska G, Puga A. Chromium cross-links histone deacetylase 1-DNA methyltransferase1 complexes to chromatin, inhibiting histone-remodeling marks critical for transcriptional activation. Mol Cell Biol,2007,27(20): 7089 ?7 101.
[23] Huang HS, Matevossian A, Whittle C, Kim SY,Schumacher A, Baker SP, Akbarian S. Prefrontal dysfunction in schizophrenia involves mixed-lineage leukemia 1-regulated histone methylation at GABAergic gene promoters. J Neurosci, 2007, 27(42): 112-154.
[24] Narendra Tuteja.Abscisic Acid and Abiotic Stress Signaling . Plant Signaling & Behavior ,2007,2(3):135-138.
[25] Yamaguchi-Shinozaki K, Shinozaki K. A novel cis-Acting Element in an Arabidopsis gene is involved in responsiveness to drought, low-temperature, or high-salt stress. Plant Cell, 1994,6(2):251-264.
[26] Baker S S, Wilhelm K S, Thomashow M F. The 5′region of Arabidopsis thaliana cor15a has cis - acting elements that confer cold - drought and ABA regulated gene expression. Plant Mol Biol , 1994,24 : 701-713.
[27] Stockinger EJ, Gilmour SJ, Thomashow MF. Arabidopsis thaliana CBF1 encodes an AP2 domain-containing transcriptional activator that binds to the C-repeat/DRE, a cis-acting DNA regulatory element that stimulates transcription in response to low temperature and water deficit. Proc Natl Acad Sci USA, 1997, 94(3): 1035-1040.
[28] Sakuma Y, Liu Q, Dubouzet J G, et al. DNA-binding specificity of the ERF /AP-domain of Arabidopsis DREBs, transcription factors involved in dehydration and cold-inducible gene expression. Biochem Biophys Res Commun, 2002,290 (3): 998-1009.
[29] Xu zs, Ni zy, Li zy, et al. Isolation and functional characterization of HvDREB1-a gene encoding a dehydration-responsive element binding protein in Hordeum vulgare. J Plant Res, 2009, 122(1):121-130.
[30] Matsukura S, Mizoi J, Yoshida T, et al. Comprehensive analysis of rice DREB2-type genes that encode transcription factors involved in the expression of abiotic stress-responsive genes. Mol Genet Genomics, 2010, 283(2):185-196.
[31] Iwasaki T,Yamaguchi-Shinozaki K, Shinoozaki K.Identifiati-on of a cis-regulatory region of a gene in Arabidopsis thaliana whose induction by dehydration is mediated by abscisic acid and requires protein synthesis. Mol Gen Genet, 1995, 247(4): 391-398.
[32] Yamaguchi-Shinozaki K, Urao T, Shinozaki K. Regulation of genes that are induced by drought stress in rabidopsis thaliana. J Plant Res,1995, 108: 127-136.
[33] Abe H, Yamaguchi-Shinozakik, Urao T, et al. Role of Arabidopsis MYC and MYB Homologs in Drought-and Abscisic Acid-Regulated Gene Expression. Plant Cell, 1997, 9:1859-1868
[34] J Paz-Ares, D Ghosal, U Wienand, et al. The regulatory cl loeus of Zea mays encodes a Protein with homology to Myb Proto-oneogene products and with struetural similarities to transcriptional activators. EMBOJ, 1987, 6(12): 3553-3558.
[35] Urao T, Yamaguchi-Shinozaki K, Urao S, et al. An Arabidopsis myb homolog is induced by dehydration stress and its gene product binds to the conserved MYB recognition sequence, Plant Cell, 1993, 5(11): 1529-1539.
[36] Manu Agarwal, Yujin Hao, Avnish Kapoor, et al. A R2R3 Type MYB Transcription Factor Is Involved in the Cold Regulation of CBF Genes and in Acquired Freezing Tolerance. JOURNAL OF BIOLOGICAL CHEMISTRY, 2006, 281(49): 37636-37645.
[37] Yong Liao, Hong-Feng Zou, Hui-Wen Wang, et al. Soybean GmMYB76, GmMYB92, and GmMYB177 genes confer stress tolerance in transgenic Arabidopsis plants. Cell Research, 2008, 18:1047-1060.
[38] Meshi T, Iwabuchi M. Plant transcription factors. Plant Cell Physiol,1995, 36:1405-1420.
[39] Onishi M, Tachi H, Kojima T, et al. Molecular cloning and characterization of a novel salt-inducible gene encoding an acidic isoform of PR-5 protein in soybean (Glycine max [L.] Merr. )Plant Physiol Biochem,2006,44(10):574-580.
[40] Manu Agarwal, Yujin Hao, Avnish Kapoor, et al. A R2R3 Type MYB Transcription Factor Is Involved in the Cold Regulation of CBF Genes and in Acquired Freezing Tolerance. JOURNAL OF BIOLOGICAL CHEMISTRY, 2006, 281(49): 37636-37645.
[41] Narendra Tuteja. Abscisic Acid and Abiotic Stress Signaling . Plant Signaling & Behavior, 2007, 2(3): 135-138.
[42] Marcotte Jr. W.R., Russel S.H., Quatrano R.S. Abscisic acid-response sequences from the Em gene of wheat.Plant Cell, 1989, 1(10): 969-976.
[43] Narusaka Y, Nakashima K, Shinwari Z K, et al. Interaction between two cis-acting elements, ABRE and DRE, in ABA-dependent expression of Arabidopsis rd29A gene in response to dehydration and high-salinity stresses.Plant J, 2003,34:137-148.
[44] Manavella PA, Dezar CA, Ariel FD, et al. Two ABREs, two redundant root-specific and one W-box cis-acting elements are functional in the sunflower HAHB4 promoter. Plant Physiol Biochem,2008,46(10):860-870.
[45] Zahur M, Maqbool A, Ifran M, et al. Isolation and functional analysis of cotton universal stress protein promoter in response to phytohormones and abiotic stresses. Mol Biol (Mosk),43(4):628-635.
[46] Liao Y, Zou HF, Wei W, et al. Soybean GmbZIP44, GmbZIP62 and GmbZIP78 genes function as negative regulator of ABA signaling and confer salt and freezing tolerance in transgenic Arabidopsis. Planta,2008,228(2):225-240.
[47] Xiao fen Jin, AiSheng Xiong, Ri He Peng, et al. OsAREB1, an ABRE-binding protein responding to ABA and glucose, has multiple functions in Arabidopsis. BMB reports, 2009, 34-39.
[48] Brown A P C, DunnM A, Godddard N J, et al.Identification of a novel low-temperature-response element in the promoter of the barley (Hordeum vulgare L) gene blt101. 1. Planta, 2001, 213: 770-780.
[49] Dunn M A, White1 A J, Vural S, et al. Identification of promoter elements in a low-temperature-responsive gene ( blt4.9 ) from barley ( Hordeum vulgare L. ) . Plant Molecular Biology, 1998,38: 551-564.
[50] Yoram Eyal, Yael Meller, Simcha Lev-Yadun, et al. A basic-type PR-1 promoter directs ethylene responsiveness, vascular and abscission zone-specific expression. The Plant Journal, 1993, 4(2):225-234.
[51] Ohme-Takagi M, Shinshi H. Ethylene-inducible DNA binding proteins that interact with an ethylene-responsive element. Plant Cell, 1995, 7: 173-182.
[52] Lijun Wu, Zhijin Zhang, Haiwen Zhang, et al. Transcriptional Modulation of Ethylene Response Factor Protein JERF3 in the Oxidative Stress Response Enhances Tolerance of Tobacco Seedlings to Salt, Drought and Freezing. Plant Physiology, 2008,148:1953-1963.
[53] Sylvia de Pater, Valentina Greco, Khanh Pham, et al. Members of a new family of DNA-binding proteins bind to a conserved cis-element in the promoters ofα-Amy2 genes Nucleic Acids Research, 1996,24(23): 4624-4631.
[54] Manavella PA, Dezar CA, Ariel FD, et al. Two ABREs, two redundant root-specific and one W-box cis-acting elements are functional in the sunflower HAHB4 promoter. Plant Physiol Biochem, 2008,46(10):860-870.
[55] Stuart Meier, RenéBastian1, Lara Donaldson, et al. Co-expression and promoter content analyses assign a role in biotic and abiotic stress responses to plant natriuretic peptides. BMC Plant Biology, 2008,8:24.
[56] Wang Z, Zhu Y, Wang L, et al. A WRKY transcription factor participates in dehydration tolerance in Boea hygrometrica by binding to the W-box elements of the galactinol synthase (BhGolS1) promoter. Planta,2009,230(6):1155-1166.
[57] Ishiguro S., Nakamura K. Characterization of a cDNA encoding a novel DNA-binding protein, SPF1, that recognizes SP8 sequences in the 50 upstream regions of genes coding for sporamin and beta-amylase from sweet potato. Mol. Gen. Genet., 1994,244 (6): 563-571.
[58] Giuliano G, Pichersky E, Malik VS. An evolutionarily con served protein binding sequence upstream of a plant light-regulated gene. Biochemistry, 1988,85: 7089-7093.
[59] Shen Q.Ho TH. Functional dissection of an abscisic acid (ABA) inducible gene reveals two independent ABA responsive complexes each containing a G-box and a novel cis-acting element. The Plant Cell, 1995,7:295-307.
[60] Shen Q.Zhang P.Ho TH. Modular nature of abscisic acid (ABA) response complexes: composite promoter unitsthat are necessary and sufficient for ABA induction of gene expression in barley. The Plant Cell, 1996,8:1107-1119.
[61] Zou M, Guan Y, Ren H, et al. A bZIP transcription factor, OsABI5, is involved in rice fertility and stress tolerance.Plant Mol Biol,2008,66(6):675-683.
[62] Ko DK, Lee MO, Hahn JS, et al. Submergence-inducible and circadian rhythmic basic helix-loop-helix protein gene in Nicotiana tabacum.J Plant Physiol, 2009,166 (10):1090-1100.
[63] Aida M, Ishida T, Fukaki H, et al. Genes involved in organ separation in Arabidopsis:analysis of the cup-shaped cotyledon mutant. Plant Cell, 1997, 9(6): 841-857.
[64] Tran L, Nakashima K, Sakuma Y, et al. Isolation and functional analysis of Arabidopsis stress-inducible NAC transcription factors that bind to a drought responsive cis-element in the EARLY RESPONSIVE TO DEHYDRATION STRESS promoter.Plant Cell, 2004, 16:2481-2498.
[65] Olsen A N, Ernst H A, Leggio L L, et al. NAC transcription factors:structurally distinct,functionally diverse .Trends in Plant Sci, 2005,10(2):79-87.
[66] Hongling Jiang, Hongmei Li, Qingyun Bu, et al. The RHA2a-interacting proteins ANAC019 and ANAC055 may play a dual role in regulating ABA response and jasmonate response. Plant Signaling&Behavior, 2009, 4(5):464-466.
[67] Seo PJ, Kim MJ, Song JS, et al. Proteolytic processing of an Arabidopsis membrane-bound NAC transcription factor is triggered by cold-induced changes in membrane fluidity. Biochem J,2010,427(3):359-367.
[68] Green PJ, Kay SA, Chua NH. Sequence-specific interactions of a pea nuclear factor with light-responsive elements upstream of the rbcS-3A gene . EMBO J, 1987, 6(9):25439-2549.
[69] Hyeong Cheol Park, Man Lyang Kim, Yun Hwan Kang, et al. Pathogen- and NaCl-Induced Expression of the SCaM-4 Promoter Is Mediated in Part by a GT-1 Box That Interacts with a GT-1-Like Transcription Factor. Plant Physiology, 2004, 135:2150-2161.
[70] Park HC, Kim ML, Kang YH, et al. Pathogen- and NaCl-induced expression of the SCaM-4 promoter is mediated in part by a GT-1 box that interacts with a GT-1-Like transcription factor. Plant Physiol, 2004, 135: 2150-2161.
[71] Rathinasabapathi B,Burnet K,Russel B L et al. Choline monoxygenase, an unusual iron-sulfur enzyme catalyzing the first step of glycine betaine synthesis in plants:prosthetic group characterization and cDNA cloning.Proc Nat1 Acad Sci.USA, 1997, 94: 3454–3458.
[72] Russell B L, Rathinasabapathi B, Hanson A D. Osmotic stress induces expression of choline monooxygenase in sugar beet and amaranth. Plant Physiol.1998,116(2): 859-865.
[73] Meng YL, Wang YM, Zhang B, Nii N. Isolation of a choline monooxygenase cDNA clone from Amaranthus tricolor and its expressions under stress conditions. Cell Res. 2001 Sep;11 (3): 187-193.
[74] LI Qiu-Li, YIN Hui, LI Dan, et al. Isolation and Characterization of CMO Gene Promoter from halophyte Suaeda liaotungensis K. Journal of Genetics and Genomics, 2007,34(4):355-361.
[75] Kenta Shirasawa, Tomoko Takabe, Tetsuko Takabe et al.Accumulation of Glycinebetaine in Rice Plants that Overexpress Choline Monooxygenase from Spinach and Evaluation of their Tolerance to Abiotic Stress Annals of Botany,2006,98: 565–571.
[76]尹辉。辽宁碱蓬CMO基因启动子功能分析。硕士论文,2007。
[77] Nazmul H B, Akira H, Nana Y et al. Regulation of betaine synthesis by precursor supply and choline monoxygenase expression in Amaranthus tricolor. Journal of Experimental Botany, 2007, 58:4203–4212.
[78] Yamaguchi-Shinozaki K,Shinozaki K.A novel cis-acting element in all Arabidopsis gene is involved in responsiveness to drought,low-temperature,or high-salt stress.Plant Cell,1994,6:251-264.
[79] Liu Q, Kasuga M, Sakuma Y et al.Two transcription factors, DREB1 and DREB2, with an EREBP/AP2 DNA binding domain separate two cellular signal transduction pathways in drought and low temperatu reresponsive gene expression, respectively, in Arabidopsis. Plant Cell, 1998, 10: 1391-1406.
[80] Lu CA, Ho TH, Ho SL et al. Three novel MYB proteins with one DNA binding repeat mediate sugar and hormone regulation ofα-amylase gene expression. Plant Cell, 2002,14 (8):1963-1980.
[81] Shen YG, Zhang WK, He SJ et al.An EREBP/AP2-type protein in Triticum aestivum was a DRE-binding transcription factor induced by cold, dehydration and ABA stress. Theor Appl Genet, 2003, 106: 923-930.
[82] EP, AC, JW et al.The sequence of rice chromosomes 11 and 12, rich in disease resistance genes and recent gene duplications.BMC Biol.2005 3:20.
[2] Seagull R W. The plant cytoskeleton.Crit Rev Plant Sci,1989, 8 : 131-167.
[3]张树珍,杨本鹏,刘飞虎,花特异表达启动子PchsA的克隆及其序列分析。农业生物技术学报,2002。10(2): 116-119。
[4] Narusaka Y, Nakashima K, Shinwari Z K, Sakuma Y, Furihata T ,Abe H , Narusaka M ,Shinozaki K ,Yamaguchi Shinozaki K. Interaction between two cis-acting elements ,ABRE and DRE ,in ABA dependent expression of A rabidopsis rd29A gene inresponse to dehydration and high salinity stresses. Plant J,2003, 34(2) : 137-148.
[5] Brown , A P , Dunn M A , Goddard N J , Hughes M A. Identification of a novel low-temperature-response element in the promoter of the barley ( Hordeum v ulgare L. ) gene blt101. 1. Planta,2001,213 (5) : 770-780.
[6] Xue GP. The DNA-binding activity of an AP2 transcriptional activator HvCBF2 involved in regulation of low-temperature responsive genes in barley is modulated by temperature. Plant J, 2003, 33:373-383.
[7] Stracke R, Werber M, Weisshaar B.The R2R3, MYB gene family in Arabidopsis thaliana. Curr Opin Plant Biol. 2001, (4): 447-456.
[8] Yamagata H, Yonesu K, Hirata A and Aizono Y.TGTCACA motif is a novel cis-regulatory enhancer element involved in fruit-specific expression of the cucumisin gene.J Biol Chem 2002, 277:11582-11590.
[9]杨英军,周鹏。番木瓜proteinase omega基因启动子的克隆及功能初步研究。云南植物研究, 2005,27 (5): 545-551。
[10] YIN Y, BEACHY R N.The regulatory regions of the rice tun-gro bacilliform virus promoter and interacting nuclear factors in rice (Oryza sativa L). Plant Journal, 1995, 7 (6): 969 - 980.
[11] Yamagata H, Yonesu K, Hirata A and Aizono Y.TGTCACA motif is a novel cis-regulatory enhancer element involved in fruit-specific expression of the cucumisin gene. J Biol Chem 2002, 277:11582-11590.
[12] Li JJ,Herskowitz1.IsolationofORC6,a component of the yeast origin recognition complex by a one-hybrid system. Science 1993;262:1870-1873.
[13] Delaney S K, Orford S J, Martin Harr ISM, et al. The fiber specificity of the cotton FSltp4 gene promoter is regulated by an AT rich p romoter region and the AT-Hook transcrip tion factor GhAT1. Plant Cell Physiology, 2007, 48 (10): 1426 - 1437.
[14] Rouster J.,Leah R.,Mundy J.,and Cameron-Mills V.Identification of a methyl jasmonate-responsive region in the promoter of a lipoxygenase 1 gene expressed in barley grain.Plant J 1997,11:513-523.
[15]张海文,谢丙炎,卢向阳,等.拟南芥防卫基因PDF1.2启动子中GCC盒是应答茉莉素反应必要的顺式作用元件。科学通报,2004,49(23): 2444-2448。
[16]吴纯清,张兴国,梁国鲁等。拟南芥AtRd29A启动子在烟草中的表达特性。西南农业学报, 2008,21(4):1045–1047。
[17] Rai M, He CK, Wu R.Comparative functional analysis of three abiotic stress-inducible promoters in transgenic rice.Transgenic Res, 2009, 18(5): 787–799.
[18]刘峻,朱国璋,解芳等。ACTA B IOCH IM ICA et BIOPHYSICA SIN ICAVo l. 29,1997,No.6:597-604。
[19] Maz n G, Lucena J M, Campoy S. LexA-binding sequences in Gram-positive and cyanobacteria are closely related [J].Mol Gen Genomics, 2004, 271: 40-49.
[20] Braunstein M, Rose AB, Holmes SG, Allis CD, Broach JR.Transcriptional silencing in yeast is associated with reduced nucleosome acetylation. Genes Dev, 1993, 7(4):592 ?6 04.
[21]张莉萍。运用染色质免疫沉淀技术分析基因表达转录调控方法的建立及初步应用。中华检验医学杂志,2006,29(5):449-451。
[22] Schnekenburger M, Talaska G, Puga A. Chromium cross-links histone deacetylase 1-DNA methyltransferase1 complexes to chromatin, inhibiting histone-remodeling marks critical for transcriptional activation. Mol Cell Biol,2007,27(20): 7089 ?7 101.
[23] Huang HS, Matevossian A, Whittle C, Kim SY,Schumacher A, Baker SP, Akbarian S. Prefrontal dysfunction in schizophrenia involves mixed-lineage leukemia 1-regulated histone methylation at GABAergic gene promoters. J Neurosci, 2007, 27(42): 112-154.
[24] Narendra Tuteja.Abscisic Acid and Abiotic Stress Signaling . Plant Signaling & Behavior ,2007,2(3):135-138.
[25] Yamaguchi-Shinozaki K, Shinozaki K. A novel cis-Acting Element in an Arabidopsis gene is involved in responsiveness to drought, low-temperature, or high-salt stress. Plant Cell, 1994,6(2):251-264.
[26] Baker S S, Wilhelm K S, Thomashow M F. The 5′region of Arabidopsis thaliana cor15a has cis - acting elements that confer cold - drought and ABA regulated gene expression. Plant Mol Biol , 1994,24 : 701-713.
[27] Stockinger EJ, Gilmour SJ, Thomashow MF. Arabidopsis thaliana CBF1 encodes an AP2 domain-containing transcriptional activator that binds to the C-repeat/DRE, a cis-acting DNA regulatory element that stimulates transcription in response to low temperature and water deficit. Proc Natl Acad Sci USA, 1997, 94(3): 1035-1040.
[28] Sakuma Y, Liu Q, Dubouzet J G, et al. DNA-binding specificity of the ERF /AP-domain of Arabidopsis DREBs, transcription factors involved in dehydration and cold-inducible gene expression. Biochem Biophys Res Commun, 2002,290 (3): 998-1009.
[29] Xu zs, Ni zy, Li zy, et al. Isolation and functional characterization of HvDREB1-a gene encoding a dehydration-responsive element binding protein in Hordeum vulgare. J Plant Res, 2009, 122(1):121-130.
[30] Matsukura S, Mizoi J, Yoshida T, et al. Comprehensive analysis of rice DREB2-type genes that encode transcription factors involved in the expression of abiotic stress-responsive genes. Mol Genet Genomics, 2010, 283(2):185-196.
[31] Iwasaki T,Yamaguchi-Shinozaki K, Shinoozaki K.Identifiati-on of a cis-regulatory region of a gene in Arabidopsis thaliana whose induction by dehydration is mediated by abscisic acid and requires protein synthesis. Mol Gen Genet, 1995, 247(4): 391-398.
[32] Yamaguchi-Shinozaki K, Urao T, Shinozaki K. Regulation of genes that are induced by drought stress in rabidopsis thaliana. J Plant Res,1995, 108: 127-136.
[33] Abe H, Yamaguchi-Shinozakik, Urao T, et al. Role of Arabidopsis MYC and MYB Homologs in Drought-and Abscisic Acid-Regulated Gene Expression. Plant Cell, 1997, 9:1859-1868
[34] J Paz-Ares, D Ghosal, U Wienand, et al. The regulatory cl loeus of Zea mays encodes a Protein with homology to Myb Proto-oneogene products and with struetural similarities to transcriptional activators. EMBOJ, 1987, 6(12): 3553-3558.
[35] Urao T, Yamaguchi-Shinozaki K, Urao S, et al. An Arabidopsis myb homolog is induced by dehydration stress and its gene product binds to the conserved MYB recognition sequence, Plant Cell, 1993, 5(11): 1529-1539.
[36] Manu Agarwal, Yujin Hao, Avnish Kapoor, et al. A R2R3 Type MYB Transcription Factor Is Involved in the Cold Regulation of CBF Genes and in Acquired Freezing Tolerance. JOURNAL OF BIOLOGICAL CHEMISTRY, 2006, 281(49): 37636-37645.
[37] Yong Liao, Hong-Feng Zou, Hui-Wen Wang, et al. Soybean GmMYB76, GmMYB92, and GmMYB177 genes confer stress tolerance in transgenic Arabidopsis plants. Cell Research, 2008, 18:1047-1060.
[38] Meshi T, Iwabuchi M. Plant transcription factors. Plant Cell Physiol,1995, 36:1405-1420.
[39] Onishi M, Tachi H, Kojima T, et al. Molecular cloning and characterization of a novel salt-inducible gene encoding an acidic isoform of PR-5 protein in soybean (Glycine max [L.] Merr. )Plant Physiol Biochem,2006,44(10):574-580.
[40] Manu Agarwal, Yujin Hao, Avnish Kapoor, et al. A R2R3 Type MYB Transcription Factor Is Involved in the Cold Regulation of CBF Genes and in Acquired Freezing Tolerance. JOURNAL OF BIOLOGICAL CHEMISTRY, 2006, 281(49): 37636-37645.
[41] Narendra Tuteja. Abscisic Acid and Abiotic Stress Signaling . Plant Signaling & Behavior, 2007, 2(3): 135-138.
[42] Marcotte Jr. W.R., Russel S.H., Quatrano R.S. Abscisic acid-response sequences from the Em gene of wheat.Plant Cell, 1989, 1(10): 969-976.
[43] Narusaka Y, Nakashima K, Shinwari Z K, et al. Interaction between two cis-acting elements, ABRE and DRE, in ABA-dependent expression of Arabidopsis rd29A gene in response to dehydration and high-salinity stresses.Plant J, 2003,34:137-148.
[44] Manavella PA, Dezar CA, Ariel FD, et al. Two ABREs, two redundant root-specific and one W-box cis-acting elements are functional in the sunflower HAHB4 promoter. Plant Physiol Biochem,2008,46(10):860-870.
[45] Zahur M, Maqbool A, Ifran M, et al. Isolation and functional analysis of cotton universal stress protein promoter in response to phytohormones and abiotic stresses. Mol Biol (Mosk),43(4):628-635.
[46] Liao Y, Zou HF, Wei W, et al. Soybean GmbZIP44, GmbZIP62 and GmbZIP78 genes function as negative regulator of ABA signaling and confer salt and freezing tolerance in transgenic Arabidopsis. Planta,2008,228(2):225-240.
[47] Xiao fen Jin, AiSheng Xiong, Ri He Peng, et al. OsAREB1, an ABRE-binding protein responding to ABA and glucose, has multiple functions in Arabidopsis. BMB reports, 2009, 34-39.
[48] Brown A P C, DunnM A, Godddard N J, et al.Identification of a novel low-temperature-response element in the promoter of the barley (Hordeum vulgare L) gene blt101. 1. Planta, 2001, 213: 770-780.
[49] Dunn M A, White1 A J, Vural S, et al. Identification of promoter elements in a low-temperature-responsive gene ( blt4.9 ) from barley ( Hordeum vulgare L. ) . Plant Molecular Biology, 1998,38: 551-564.
[50] Yoram Eyal, Yael Meller, Simcha Lev-Yadun, et al. A basic-type PR-1 promoter directs ethylene responsiveness, vascular and abscission zone-specific expression. The Plant Journal, 1993, 4(2):225-234.
[51] Ohme-Takagi M, Shinshi H. Ethylene-inducible DNA binding proteins that interact with an ethylene-responsive element. Plant Cell, 1995, 7: 173-182.
[52] Lijun Wu, Zhijin Zhang, Haiwen Zhang, et al. Transcriptional Modulation of Ethylene Response Factor Protein JERF3 in the Oxidative Stress Response Enhances Tolerance of Tobacco Seedlings to Salt, Drought and Freezing. Plant Physiology, 2008,148:1953-1963.
[53] Sylvia de Pater, Valentina Greco, Khanh Pham, et al. Members of a new family of DNA-binding proteins bind to a conserved cis-element in the promoters ofα-Amy2 genes Nucleic Acids Research, 1996,24(23): 4624-4631.
[54] Manavella PA, Dezar CA, Ariel FD, et al. Two ABREs, two redundant root-specific and one W-box cis-acting elements are functional in the sunflower HAHB4 promoter. Plant Physiol Biochem, 2008,46(10):860-870.
[55] Stuart Meier, RenéBastian1, Lara Donaldson, et al. Co-expression and promoter content analyses assign a role in biotic and abiotic stress responses to plant natriuretic peptides. BMC Plant Biology, 2008,8:24.
[56] Wang Z, Zhu Y, Wang L, et al. A WRKY transcription factor participates in dehydration tolerance in Boea hygrometrica by binding to the W-box elements of the galactinol synthase (BhGolS1) promoter. Planta,2009,230(6):1155-1166.
[57] Ishiguro S., Nakamura K. Characterization of a cDNA encoding a novel DNA-binding protein, SPF1, that recognizes SP8 sequences in the 50 upstream regions of genes coding for sporamin and beta-amylase from sweet potato. Mol. Gen. Genet., 1994,244 (6): 563-571.
[58] Giuliano G, Pichersky E, Malik VS. An evolutionarily con served protein binding sequence upstream of a plant light-regulated gene. Biochemistry, 1988,85: 7089-7093.
[59] Shen Q.Ho TH. Functional dissection of an abscisic acid (ABA) inducible gene reveals two independent ABA responsive complexes each containing a G-box and a novel cis-acting element. The Plant Cell, 1995,7:295-307.
[60] Shen Q.Zhang P.Ho TH. Modular nature of abscisic acid (ABA) response complexes: composite promoter unitsthat are necessary and sufficient for ABA induction of gene expression in barley. The Plant Cell, 1996,8:1107-1119.
[61] Zou M, Guan Y, Ren H, et al. A bZIP transcription factor, OsABI5, is involved in rice fertility and stress tolerance.Plant Mol Biol,2008,66(6):675-683.
[62] Ko DK, Lee MO, Hahn JS, et al. Submergence-inducible and circadian rhythmic basic helix-loop-helix protein gene in Nicotiana tabacum.J Plant Physiol, 2009,166 (10):1090-1100.
[63] Aida M, Ishida T, Fukaki H, et al. Genes involved in organ separation in Arabidopsis:analysis of the cup-shaped cotyledon mutant. Plant Cell, 1997, 9(6): 841-857.
[64] Tran L, Nakashima K, Sakuma Y, et al. Isolation and functional analysis of Arabidopsis stress-inducible NAC transcription factors that bind to a drought responsive cis-element in the EARLY RESPONSIVE TO DEHYDRATION STRESS promoter.Plant Cell, 2004, 16:2481-2498.
[65] Olsen A N, Ernst H A, Leggio L L, et al. NAC transcription factors:structurally distinct,functionally diverse .Trends in Plant Sci, 2005,10(2):79-87.
[66] Hongling Jiang, Hongmei Li, Qingyun Bu, et al. The RHA2a-interacting proteins ANAC019 and ANAC055 may play a dual role in regulating ABA response and jasmonate response. Plant Signaling&Behavior, 2009, 4(5):464-466.
[67] Seo PJ, Kim MJ, Song JS, et al. Proteolytic processing of an Arabidopsis membrane-bound NAC transcription factor is triggered by cold-induced changes in membrane fluidity. Biochem J,2010,427(3):359-367.
[68] Green PJ, Kay SA, Chua NH. Sequence-specific interactions of a pea nuclear factor with light-responsive elements upstream of the rbcS-3A gene . EMBO J, 1987, 6(9):25439-2549.
[69] Hyeong Cheol Park, Man Lyang Kim, Yun Hwan Kang, et al. Pathogen- and NaCl-Induced Expression of the SCaM-4 Promoter Is Mediated in Part by a GT-1 Box That Interacts with a GT-1-Like Transcription Factor. Plant Physiology, 2004, 135:2150-2161.
[70] Park HC, Kim ML, Kang YH, et al. Pathogen- and NaCl-induced expression of the SCaM-4 promoter is mediated in part by a GT-1 box that interacts with a GT-1-Like transcription factor. Plant Physiol, 2004, 135: 2150-2161.
[71] Rathinasabapathi B,Burnet K,Russel B L et al. Choline monoxygenase, an unusual iron-sulfur enzyme catalyzing the first step of glycine betaine synthesis in plants:prosthetic group characterization and cDNA cloning.Proc Nat1 Acad Sci.USA, 1997, 94: 3454–3458.
[72] Russell B L, Rathinasabapathi B, Hanson A D. Osmotic stress induces expression of choline monooxygenase in sugar beet and amaranth. Plant Physiol.1998,116(2): 859-865.
[73] Meng YL, Wang YM, Zhang B, Nii N. Isolation of a choline monooxygenase cDNA clone from Amaranthus tricolor and its expressions under stress conditions. Cell Res. 2001 Sep;11 (3): 187-193.
[74] LI Qiu-Li, YIN Hui, LI Dan, et al. Isolation and Characterization of CMO Gene Promoter from halophyte Suaeda liaotungensis K. Journal of Genetics and Genomics, 2007,34(4):355-361.
[75] Kenta Shirasawa, Tomoko Takabe, Tetsuko Takabe et al.Accumulation of Glycinebetaine in Rice Plants that Overexpress Choline Monooxygenase from Spinach and Evaluation of their Tolerance to Abiotic Stress Annals of Botany,2006,98: 565–571.
[76]尹辉。辽宁碱蓬CMO基因启动子功能分析。硕士论文,2007。
[77] Nazmul H B, Akira H, Nana Y et al. Regulation of betaine synthesis by precursor supply and choline monoxygenase expression in Amaranthus tricolor. Journal of Experimental Botany, 2007, 58:4203–4212.
[78] Yamaguchi-Shinozaki K,Shinozaki K.A novel cis-acting element in all Arabidopsis gene is involved in responsiveness to drought,low-temperature,or high-salt stress.Plant Cell,1994,6:251-264.
[79] Liu Q, Kasuga M, Sakuma Y et al.Two transcription factors, DREB1 and DREB2, with an EREBP/AP2 DNA binding domain separate two cellular signal transduction pathways in drought and low temperatu reresponsive gene expression, respectively, in Arabidopsis. Plant Cell, 1998, 10: 1391-1406.
[80] Lu CA, Ho TH, Ho SL et al. Three novel MYB proteins with one DNA binding repeat mediate sugar and hormone regulation ofα-amylase gene expression. Plant Cell, 2002,14 (8):1963-1980.
[81] Shen YG, Zhang WK, He SJ et al.An EREBP/AP2-type protein in Triticum aestivum was a DRE-binding transcription factor induced by cold, dehydration and ABA stress. Theor Appl Genet, 2003, 106: 923-930.
[82] EP, AC, JW et al.The sequence of rice chromosomes 11 and 12, rich in disease resistance genes and recent gene duplications.BMC Biol.2005 3:20.