森林草莓MYB家族鉴定及生物信息学分析
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摘要
为研究草莓MYB基因家族,采用生物信息学方法检索森林草莓(Fragaria vesca L.)基因组中MYB基因,并对其基因结构、蛋白保守结构域、染色体定位、基因进化及花药发育过程中表达水平进行分析。结果表明,草莓基因组中可能含有105个R2R3-MYB、4个MYB3R、1个MYB4R基因,它们不均匀分布在7条草莓染色体上,其中5号染色体上数量最多为26个,部分基因在染色体上形成基因簇。理化分析发现这些MYB蛋白长度和大小差异较大,但均具有保守的R2和R3型结构域,R重复单元均含有特征性的氨基酸,包括最保守的有序间隔出现的色氨酸(W)残基。通过系统进化关系和基因结构信息,草莓MYB基因家族被分成了34个亚组(A1～A34),同时利用拟南芥基因功能预测草莓中对应的直系同源基因的功能,结果表明草莓MYB基因家族参与调控黄酮物质合成、细胞发育、生物/非生物胁迫。花药发育过程中表达水平分析发现FvMYB2、FvMYB85、FvMYB74、FvMYB28在花药发育的5个时期中表达量逐渐升高,尤其是发育后期表达量较高,表明这些基因可能调控花药发育过程。本研究结果为草莓MYB基因家族的功能研究提供参考。
To get insight of MYB genes in strawberry,we used bioinformatics methods to screen the MYB genes from the Fragaria vesca L. genome database,and further analyzed gene structure,conserved motif,chromosomal location,phylogenetic analysis and expression in the period of the development of anther.Our results showed that three types of MYB transcription factors,including 105 R2 R3-MYB,4 MYB3 R,1 MYB4 R,were identified.These genes unevenly distributed on 7 chromosomes,e.g.,that chromosome Fvb5 contained the largest number of 26 MYB genes.Some of MYBs formed distinct tandem duplicate gene clusters.Although the protein length of strawberry MYB genes differed,all of MYBs contained conserved R2 R3 domain,with characteristic amino acids,especially orderly spaced conservative tryptophan residues(W).Phylogenetic and gene structure analysis showed that strawberry MYB genes were divided into 34 distinct groups(A1-A34).The function prediction of strawberry orthologous genes using Arabidopsis thaliana L. gene annotation showed strawberry MYB genes were most likely involved in ?avone synthesis,cell fate,biotic and abiotic stresses.Temporal or spatial expression trends of MYB genes during strawberry anther development indicated that FvMYB2,FvMYB85,FvMYB74,and FvMYB28 had increased expression during 5 period of the development of strawberry anther,which were mainly expressed in the later period of anther development.These results suggested that those MYB genes may play important role during strawberry anther development.Thus,this paper provided information for functional identi?cation of strawberry MYB genes.
To get insight of MYB genes in strawberry,we used bioinformatics methods to screen the MYB genes from the Fragaria vesca L. genome database,and further analyzed gene structure,conserved motif,chromosomal location,phylogenetic analysis and expression in the period of the development of anther.Our results showed that three types of MYB transcription factors,including 105 R2 R3-MYB,4 MYB3 R,1 MYB4 R,were identified.These genes unevenly distributed on 7 chromosomes,e.g.,that chromosome Fvb5 contained the largest number of 26 MYB genes.Some of MYBs formed distinct tandem duplicate gene clusters.Although the protein length of strawberry MYB genes differed,all of MYBs contained conserved R2 R3 domain,with characteristic amino acids,especially orderly spaced conservative tryptophan residues(W).Phylogenetic and gene structure analysis showed that strawberry MYB genes were divided into 34 distinct groups(A1-A34).The function prediction of strawberry orthologous genes using Arabidopsis thaliana L. gene annotation showed strawberry MYB genes were most likely involved in ?avone synthesis,cell fate,biotic and abiotic stresses.Temporal or spatial expression trends of MYB genes during strawberry anther development indicated that FvMYB2,FvMYB85,FvMYB74,and FvMYB28 had increased expression during 5 period of the development of strawberry anther,which were mainly expressed in the later period of anther development.These results suggested that those MYB genes may play important role during strawberry anther development.Thus,this paper provided information for functional identi?cation of strawberry MYB genes.
引文
[1] Latchman D S.Transcription factors:an overview-The International Journal of Biochemistry&Cell Biology, 1997,29(12):1305-1312
[2] Riechmann J L, Heard J, Martin G, Reuber L, Jiang C, Keddie J, Adam L, Pineda O, Ratcliffe O J, Samaha R R, Creelman R, Pilgrim M, Broun P, Zhang J Z, Ghandehari D, Sherman B K, Yu G-Arabidopsis transcription factors:Genome-wide comparative analysis among eukaryotes-Science, 2000,290(5499):2105-2110
[3] Dubos C, Stracke R, Grotewold E, Weisshaar B, Martin C,Lepiniec L-MYB transcription factors in Arabidopsis.Trends in Plant Science, 2010, 15(10):573-581
[4]Ogata K, Kaneiishii C, Sasaki M, Hatanaka H, Nagadoi A,Enari M,Nakamura H,Nishimura Y,Ishii S,Sarai A.The cavity in the hydrophobic core of Myb DNA binding domain is reserved for DNA recognition and trans-activation-Nature Structural&Molecular Biology, 1996,3(2):178-187
[5]魏麦玲,裴丽丽,刘佳明,闵东红,陈明,李连城,马有志,徐兆师,张小红·大豆MYB转录因子基因GmMYB174的克隆及分子特性分析·植物遗传资源学报,2015,16(1):94-99Wei M L, Pei L L, Liu J M, Min D H, Chen M, Li L C,Ma Y Z, Xu Z S, Zhang X H-Isolation and molecular characteristics analysis of soybean transcription factor gene GmMYB174.Journal of Plant Genetic Resources, 2015,16(1):94-99
[6]魏潇,曹珂,王力荣,朱更瑞,方伟超,陈昌文·红根甘肃桃根系MYB基因片段的克隆及序列分析·植物遗传资源学报,2011,12(3):455-459,463Wei X, Cao K, Wang L R, Zhu G R, Fang W C, Chen C W-Cloning and characterization of MYB genes from root system of Honggengansutao peach(Prunus kansuensis L-)-Journal of Plant Genetic Resources, 2011,12(3):455-459,463
[7]Katiyar A, Smita S, Lenka S K, Rajwanshi R, Chinnusamy V,Bansal K C-Genome-wide classification and expression analysis of MYB transcription factor families in rice and Arabidopsis.BMC Genomics, 2012, 13:544
[8] Chen Y H, Yang X Y, He K, Liu M H, Li J G, Gao Z F, Lin Z Q, Zhang Y F, Wang X X, Qiu X M, Shen Y P, Zhang L,Deng X H, Luo J C, Deng X W, Chen Z L, Gu H Y.The MYB transcription factor super family of Arabidopsis:expression analysis and phylogenetic comparison with the rice MYB family-Nature Structural&Molecular Biology, 2006, 60:107-124
[9] Dias A P, Braun E L, McMullen M D, Grotewold E-Recently duplicated maize R2R3 Myb genes provide evidence for distinct mechanisms of evolutionary divergence after duplication-Plant Physiology, 2003, 131(2):610-620
[10] Zimmermann I M, Heim M A, Weisshaar B, Uhrig J F.Comprehensive identification of Arabidopsis thaliana MYB transcription factors interacting with R/B-like BHLH proteins.The Plant Journal, 2004,40:22-34
[11] Medina-Puche L, Cumplido-Laso G, Amil-Ruiz F, Hoffmann T, Ring L, Rodri guez-Franco A, Caballero J L, Schwab W,Mun oz-Blanco J, Blanco-Portales R.MYB10 plays a major role in the regulation of flavonoid/phenylpropanoid metabolism during ripening of Fragaria x ananassa fruits-Journal of Experimental Botany, 2015,65:401-417
[12] Kadomura-Ishikawa Y, Miyawaki K, Takahashi A, Noji S.RNAi-mediated silencing and overexpression of the FaMYBl gene and its effect on anthocyanin accumulation in strawberry fruit-Biologia Plantarum, 2015,59(4):677-685
[13] Tropf S, Lanz T-Evidence that stilbene synthases have developed from chalcone synthases several times in the course of evolution-Journal of Molecular Evolution, 1994, 38:610-618
[14] Cominelli E, Galbiati M, Vavasseur A, Conti L, Sala T,Vuylsteke M, Leonhardt N, Dellaporta S L, Tonelli C-A guardcell-specific MYB transcription factor regulates stomatal movements and plant drought tolerance-Current Biology, 2005,15:1196-1200
[15] Seo P J, Xiang F, Qiao M, Park J Y, Lee Y N, Kim S G, Lee Y H, Park W J, Park C M-The MYB96 transcription factor mediates abscisic acid signaling during drought stress response in Arabidopsis-Plant Physiology, 2009, 151:275-289
[16] Seo P J and Park C M-MYB96-mediated abscisic acid signals induce pathogen resistance response by promoting salicylic acid biosynthesis in Arabidopsis-The New Physiology, 2010, 186:471-483
[17] Mandaokar A and Browse J.MYB108 acts together with MYB24 to regulate jasmonate-mediated stamen maturation in Arabidopsis-Plant Physiology, 2009, 149:851-862
[18] Zhang Z B, Zhu J, Gao J F, Wang C, Li H, Li H, Zhang H Q, Zhang S, Wang D M, Wang Q X, Huang H, Xia H J,Yang Z N-Transcription factor AtMYB103 is required for anther development by regulating tapetum development, callose dissolution and exine formation in Arabidopsis-The Plant Journal, 2007, 52:528-538
[19] Brownfield L,Hafidh S,Borg M,Sidorova A,Mori T,Twell D. A plant germline-specific integrator of sperm specification and cell cycle progression-PLoS Genetics, 2009, 5(3):e1000430
[20] Du H, Feng B R, Yang S S, Huang Y B, Tang Y X The R2R3-MYB transcription factor gene family in maize-PLoS One,2012,7(6):e37463
[21] Stracke R, Holtgra we D, Schneider J, Pucker B, So rensen T R,Weisshaar B-Genome-wide identification and characterisation of R2R3-MYB genes in sugar beet(Beta vulgaris).BMC Plant Biology, 2014, 14(1):249-266
[22] Matus J T, Aquea F, Arce-Johnson P-Analysis of the grape MYB R2R3 subfamily reveals expanded wine quality-related clades and conserved gene structure organization across Vitis and Arabidopsis genomes-BMC Plant Biology, 2008,8(1):83-98
[23]Zhou H, Peng Q, Zhao J, Owiti A, Ren F, Liao L, Wang L,Deng X, Jiang Q, Han Y.Multiple R2R3-MYB transcription factors involved in the regulation of anthocyanin accumulation in peach flower-Frontiers in Plant Science, 2016, 7(28):1557-1568
[24] Shulaev V, Sargent D J, Crowhurst R N, Mockler T C, Folkerts O, Delcher A L, Jaiswal P, Mockaitis K, Liston A, Mane S P,Bums P, Davis T M, Slovin J P, Bassil N, Hellens R P, Evans C, Harkins T, Kodira C, Desany B, Crasta O R, Jensen R V,Allan A C, Michael T P, Setubal J C, Celton J M, Rees D J G,Williams K P, Holt S H, Rojas J J R, Chatterjee M, Liu B,Silva H, Meisel L, Adato A, Filichkin S A, Troggio M, Viola R, Ashman T L, Wang H, Dharmawardhana P, Elser J, Raja R, Priest H D, Bryant D W, Fox S E, Givan S A, Wilhelm L J,Naithani S, Christoffels A, Salama D Y, Carter J, Girona E L,Zdepski A, Wang W Q, Kerstetter R A, Schwab W, Korban S S, Davik J, Monfort A, Denoyes-Rothan B, Arus P, Mittler R,Flinn B, Aharoni A, Bennetzen J L, Salzberg S L, Dickerman A W, Velasco R, Borodovsky M, Veilleux R E, Folta K M-The genome of woodland strawberry(Fragaria vesca).Nature Genetics, 2010,43:109-116
[25]Eddy S R Profile hidden Markov models.Bioinformatics,1998,14(9):755-763
[26]Thompson J D, Higgins D G, Gibson T J-CLUSTAL W:improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific alignment through sequence weighting, position-specific gap penalties and weight matrix choice-Nucleic Acids Research,1994,22(22):4673-4680
[27] Crooks G E, Hon G, Chandonia J M, Brenner S E.WebLogo:A sequence logo generator.Genome Research, 2004, 14:1188-1190
[28] Hu B,Jin J P,Guo A Y,Zhang H,Luo J,Gao G GSDS 2 0:an upgraded gene feature visualization server Bioinformatics,2015,31(8):1296-1297
[29]Tamura K,Peterson D,Peterson N,Stecher G,Nei M,Kumar S-MEGA5:Molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods.Molecular Biology and Evolution, 2011,28(10):2731-2739
[30] Darwish O, Slovin J P, Kang C, Hollender C A, Geretz A,Houston S, Liu Z, Alkharouf N W-SGR:an online genomic resource for the woodland strawberry-BMC Plant Biology,2013, 13(1):223
[31] Deng W K,Wang Y B,Liu Z X,Cheng H,Xue Y.Heml:A toolkit for illustrating heatmaps-PLoS One, 2014,9(11):e111988
[32] Stracke R,Ishihara H,Huep G,Barsch A,Mehrtens F,Niehaus K, Weisshaar B Differential regulation of closely related R2R3-MYB transcription factors controls flavonol accumulation in different parts of the Arabidopsis thaliana seedling-Plant Journal, 2007, 50:660-677
[33] Gonzalez A, Zhao M, Leavitt J M, Lloyd A M-Regulation of the anthocyanin biosynthetic pathway by the TTG1/bHLH/Myb transcriptional complex in Arabidopsis seedlings-The Plant Journal, 2008, 53:814-827
[34] Kang Y H, Kirik V, Hulskamp M, Nam K H, Hagely K, Lee M M, Schiefelbein J.The MYB23 gene provides a positive feedback loop for cell fate specification in the Arabidopsis root epidermis-Plant Cell, 2009, 21:1080-1094
[35] Cominelli E, Galbiati M, Vavasseur A, Conti L, Sala T,Vuylsteke M, Leonhardt N, Dellaporta S L, Tonelli C.A guardcell-specific MYB transcription factor regulates stomatal movements and plant drought tolerance-Current Biology, 2005,15:1196-1200
[36] Rosinski J A, Atchley W R.Molecular evolution of the Mybfamily of transcription factors:evidence for polyphyletic origin.Journal of Molecular Evolution, 1998,46:74-83
[37] Haga N, Kato K, Murase M, Araki S, Kubo M, Demura T,Suzuki K, Mu ller I, Voss U, Ju rgens G, Ito M.R1R2R3-Myb proteins positively regulate cytokinesis through activation of KNOLLE transcription in Arabidopsis thaliana-Development,2007, 134:1101-1110
[38]袁华招,赵密珍,吴伟民,于红梅,王庆莲·葡萄CHS和STS基因家族生物信息学鉴定和表达分析·植物遗传资源学报,2016, 17(4):756-765Yuan H Z, Zhao M Z, Wu W M, Yu H M, Wang Q L.Genomewide identification and expression analysis of CHS and STS gene families in grape-Journal of Plant Genetic Resources,2016, 17(4):756-765
[39] Mandaokar A, Browse J.MYB108 acts together with MYB24 to regulate jasmonate-mediated stamen maturation in Arabidopsis.Plant Physiology, 2009, 149:851-862
[2] Riechmann J L, Heard J, Martin G, Reuber L, Jiang C, Keddie J, Adam L, Pineda O, Ratcliffe O J, Samaha R R, Creelman R, Pilgrim M, Broun P, Zhang J Z, Ghandehari D, Sherman B K, Yu G-Arabidopsis transcription factors:Genome-wide comparative analysis among eukaryotes-Science, 2000,290(5499):2105-2110
[3] Dubos C, Stracke R, Grotewold E, Weisshaar B, Martin C,Lepiniec L-MYB transcription factors in Arabidopsis.Trends in Plant Science, 2010, 15(10):573-581
[4]Ogata K, Kaneiishii C, Sasaki M, Hatanaka H, Nagadoi A,Enari M,Nakamura H,Nishimura Y,Ishii S,Sarai A.The cavity in the hydrophobic core of Myb DNA binding domain is reserved for DNA recognition and trans-activation-Nature Structural&Molecular Biology, 1996,3(2):178-187
[5]魏麦玲,裴丽丽,刘佳明,闵东红,陈明,李连城,马有志,徐兆师,张小红·大豆MYB转录因子基因GmMYB174的克隆及分子特性分析·植物遗传资源学报,2015,16(1):94-99Wei M L, Pei L L, Liu J M, Min D H, Chen M, Li L C,Ma Y Z, Xu Z S, Zhang X H-Isolation and molecular characteristics analysis of soybean transcription factor gene GmMYB174.Journal of Plant Genetic Resources, 2015,16(1):94-99
[6]魏潇,曹珂,王力荣,朱更瑞,方伟超,陈昌文·红根甘肃桃根系MYB基因片段的克隆及序列分析·植物遗传资源学报,2011,12(3):455-459,463Wei X, Cao K, Wang L R, Zhu G R, Fang W C, Chen C W-Cloning and characterization of MYB genes from root system of Honggengansutao peach(Prunus kansuensis L-)-Journal of Plant Genetic Resources, 2011,12(3):455-459,463
[7]Katiyar A, Smita S, Lenka S K, Rajwanshi R, Chinnusamy V,Bansal K C-Genome-wide classification and expression analysis of MYB transcription factor families in rice and Arabidopsis.BMC Genomics, 2012, 13:544
[8] Chen Y H, Yang X Y, He K, Liu M H, Li J G, Gao Z F, Lin Z Q, Zhang Y F, Wang X X, Qiu X M, Shen Y P, Zhang L,Deng X H, Luo J C, Deng X W, Chen Z L, Gu H Y.The MYB transcription factor super family of Arabidopsis:expression analysis and phylogenetic comparison with the rice MYB family-Nature Structural&Molecular Biology, 2006, 60:107-124
[9] Dias A P, Braun E L, McMullen M D, Grotewold E-Recently duplicated maize R2R3 Myb genes provide evidence for distinct mechanisms of evolutionary divergence after duplication-Plant Physiology, 2003, 131(2):610-620
[10] Zimmermann I M, Heim M A, Weisshaar B, Uhrig J F.Comprehensive identification of Arabidopsis thaliana MYB transcription factors interacting with R/B-like BHLH proteins.The Plant Journal, 2004,40:22-34
[11] Medina-Puche L, Cumplido-Laso G, Amil-Ruiz F, Hoffmann T, Ring L, Rodri guez-Franco A, Caballero J L, Schwab W,Mun oz-Blanco J, Blanco-Portales R.MYB10 plays a major role in the regulation of flavonoid/phenylpropanoid metabolism during ripening of Fragaria x ananassa fruits-Journal of Experimental Botany, 2015,65:401-417
[12] Kadomura-Ishikawa Y, Miyawaki K, Takahashi A, Noji S.RNAi-mediated silencing and overexpression of the FaMYBl gene and its effect on anthocyanin accumulation in strawberry fruit-Biologia Plantarum, 2015,59(4):677-685
[13] Tropf S, Lanz T-Evidence that stilbene synthases have developed from chalcone synthases several times in the course of evolution-Journal of Molecular Evolution, 1994, 38:610-618
[14] Cominelli E, Galbiati M, Vavasseur A, Conti L, Sala T,Vuylsteke M, Leonhardt N, Dellaporta S L, Tonelli C-A guardcell-specific MYB transcription factor regulates stomatal movements and plant drought tolerance-Current Biology, 2005,15:1196-1200
[15] Seo P J, Xiang F, Qiao M, Park J Y, Lee Y N, Kim S G, Lee Y H, Park W J, Park C M-The MYB96 transcription factor mediates abscisic acid signaling during drought stress response in Arabidopsis-Plant Physiology, 2009, 151:275-289
[16] Seo P J and Park C M-MYB96-mediated abscisic acid signals induce pathogen resistance response by promoting salicylic acid biosynthesis in Arabidopsis-The New Physiology, 2010, 186:471-483
[17] Mandaokar A and Browse J.MYB108 acts together with MYB24 to regulate jasmonate-mediated stamen maturation in Arabidopsis-Plant Physiology, 2009, 149:851-862
[18] Zhang Z B, Zhu J, Gao J F, Wang C, Li H, Li H, Zhang H Q, Zhang S, Wang D M, Wang Q X, Huang H, Xia H J,Yang Z N-Transcription factor AtMYB103 is required for anther development by regulating tapetum development, callose dissolution and exine formation in Arabidopsis-The Plant Journal, 2007, 52:528-538
[19] Brownfield L,Hafidh S,Borg M,Sidorova A,Mori T,Twell D. A plant germline-specific integrator of sperm specification and cell cycle progression-PLoS Genetics, 2009, 5(3):e1000430
[20] Du H, Feng B R, Yang S S, Huang Y B, Tang Y X The R2R3-MYB transcription factor gene family in maize-PLoS One,2012,7(6):e37463
[21] Stracke R, Holtgra we D, Schneider J, Pucker B, So rensen T R,Weisshaar B-Genome-wide identification and characterisation of R2R3-MYB genes in sugar beet(Beta vulgaris).BMC Plant Biology, 2014, 14(1):249-266
[22] Matus J T, Aquea F, Arce-Johnson P-Analysis of the grape MYB R2R3 subfamily reveals expanded wine quality-related clades and conserved gene structure organization across Vitis and Arabidopsis genomes-BMC Plant Biology, 2008,8(1):83-98
[23]Zhou H, Peng Q, Zhao J, Owiti A, Ren F, Liao L, Wang L,Deng X, Jiang Q, Han Y.Multiple R2R3-MYB transcription factors involved in the regulation of anthocyanin accumulation in peach flower-Frontiers in Plant Science, 2016, 7(28):1557-1568
[24] Shulaev V, Sargent D J, Crowhurst R N, Mockler T C, Folkerts O, Delcher A L, Jaiswal P, Mockaitis K, Liston A, Mane S P,Bums P, Davis T M, Slovin J P, Bassil N, Hellens R P, Evans C, Harkins T, Kodira C, Desany B, Crasta O R, Jensen R V,Allan A C, Michael T P, Setubal J C, Celton J M, Rees D J G,Williams K P, Holt S H, Rojas J J R, Chatterjee M, Liu B,Silva H, Meisel L, Adato A, Filichkin S A, Troggio M, Viola R, Ashman T L, Wang H, Dharmawardhana P, Elser J, Raja R, Priest H D, Bryant D W, Fox S E, Givan S A, Wilhelm L J,Naithani S, Christoffels A, Salama D Y, Carter J, Girona E L,Zdepski A, Wang W Q, Kerstetter R A, Schwab W, Korban S S, Davik J, Monfort A, Denoyes-Rothan B, Arus P, Mittler R,Flinn B, Aharoni A, Bennetzen J L, Salzberg S L, Dickerman A W, Velasco R, Borodovsky M, Veilleux R E, Folta K M-The genome of woodland strawberry(Fragaria vesca).Nature Genetics, 2010,43:109-116
[25]Eddy S R Profile hidden Markov models.Bioinformatics,1998,14(9):755-763
[26]Thompson J D, Higgins D G, Gibson T J-CLUSTAL W:improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific alignment through sequence weighting, position-specific gap penalties and weight matrix choice-Nucleic Acids Research,1994,22(22):4673-4680
[27] Crooks G E, Hon G, Chandonia J M, Brenner S E.WebLogo:A sequence logo generator.Genome Research, 2004, 14:1188-1190
[28] Hu B,Jin J P,Guo A Y,Zhang H,Luo J,Gao G GSDS 2 0:an upgraded gene feature visualization server Bioinformatics,2015,31(8):1296-1297
[29]Tamura K,Peterson D,Peterson N,Stecher G,Nei M,Kumar S-MEGA5:Molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods.Molecular Biology and Evolution, 2011,28(10):2731-2739
[30] Darwish O, Slovin J P, Kang C, Hollender C A, Geretz A,Houston S, Liu Z, Alkharouf N W-SGR:an online genomic resource for the woodland strawberry-BMC Plant Biology,2013, 13(1):223
[31] Deng W K,Wang Y B,Liu Z X,Cheng H,Xue Y.Heml:A toolkit for illustrating heatmaps-PLoS One, 2014,9(11):e111988
[32] Stracke R,Ishihara H,Huep G,Barsch A,Mehrtens F,Niehaus K, Weisshaar B Differential regulation of closely related R2R3-MYB transcription factors controls flavonol accumulation in different parts of the Arabidopsis thaliana seedling-Plant Journal, 2007, 50:660-677
[33] Gonzalez A, Zhao M, Leavitt J M, Lloyd A M-Regulation of the anthocyanin biosynthetic pathway by the TTG1/bHLH/Myb transcriptional complex in Arabidopsis seedlings-The Plant Journal, 2008, 53:814-827
[34] Kang Y H, Kirik V, Hulskamp M, Nam K H, Hagely K, Lee M M, Schiefelbein J.The MYB23 gene provides a positive feedback loop for cell fate specification in the Arabidopsis root epidermis-Plant Cell, 2009, 21:1080-1094
[35] Cominelli E, Galbiati M, Vavasseur A, Conti L, Sala T,Vuylsteke M, Leonhardt N, Dellaporta S L, Tonelli C.A guardcell-specific MYB transcription factor regulates stomatal movements and plant drought tolerance-Current Biology, 2005,15:1196-1200
[36] Rosinski J A, Atchley W R.Molecular evolution of the Mybfamily of transcription factors:evidence for polyphyletic origin.Journal of Molecular Evolution, 1998,46:74-83
[37] Haga N, Kato K, Murase M, Araki S, Kubo M, Demura T,Suzuki K, Mu ller I, Voss U, Ju rgens G, Ito M.R1R2R3-Myb proteins positively regulate cytokinesis through activation of KNOLLE transcription in Arabidopsis thaliana-Development,2007, 134:1101-1110
[38]袁华招,赵密珍,吴伟民,于红梅,王庆莲·葡萄CHS和STS基因家族生物信息学鉴定和表达分析·植物遗传资源学报,2016, 17(4):756-765Yuan H Z, Zhao M Z, Wu W M, Yu H M, Wang Q L.Genomewide identification and expression analysis of CHS and STS gene families in grape-Journal of Plant Genetic Resources,2016, 17(4):756-765
[39] Mandaokar A, Browse J.MYB108 acts together with MYB24 to regulate jasmonate-mediated stamen maturation in Arabidopsis.Plant Physiology, 2009, 149:851-862