植物bHLH转录因子的研究进展
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摘要
bHLH转录因子是植物中最大的转录因子家族之一。为深入了解植物bHLH转录因子的功能和结构特点,本研究阐述了bHLH转录因子的结构特点;总结了植物b HLH转录因子的功能,包括bHLH转录因子调控信号转导,bHLH转录因子调控植物体发育,bHLH转录因子参与植物的抗逆。但下一步仍需对该基因家族的功能进行系统的分类及分析,特别加强对其他非模式生物中bHLH转录因子的研究。
The bHLH transcription factors are one of the largest families of transcription factors in plants. In order to understand the functions and structure features of plant bHLH transcription factors, we clarify the structure features of bHLH transcription factors, and summarize the functions of plant bHLH transcription factors, including bHLH transcription factors regulating signal transduction, bHLH transcription factors regulating plant development, and bHLH transcription factors taking part in plant stress resistance. But the next step is still to systematically classify and analyze the functions of the gene family, in particular, the study of bHLH transcription factors in other non-model organisms should be strengthened.
The bHLH transcription factors are one of the largest families of transcription factors in plants. In order to understand the functions and structure features of plant bHLH transcription factors, we clarify the structure features of bHLH transcription factors, and summarize the functions of plant bHLH transcription factors, including bHLH transcription factors regulating signal transduction, bHLH transcription factors regulating plant development, and bHLH transcription factors taking part in plant stress resistance. But the next step is still to systematically classify and analyze the functions of the gene family, in particular, the study of bHLH transcription factors in other non-model organisms should be strengthened.
引文
[1]Heim M A,Jakoby M,Werber M,et al.The Basic Helix-LoopHelix Transcription Factor Family in Plants:A Genome-Wide Study of Protein Structure and Functional Diversity[J].Molecular Biology&Evolution,2003,20(5):735-747.
[2]Li X,Duan X,Jiang H,et al.Genome-Wide Analysis of Basic/Helix-Loop-Helix Transcription Factor Family in Rice and Arabidopsis[J].Plant Physiology,2006,141(4):1167-1184.
[3]Chang W H,Lai A G.Genome-wide analyses of the bHLHsuperfamily in crustaceans:reappraisal of higher-order groupings and evidence for lineage-specific duplications[J].Royal Society Open Science,2018,5(3):172433.
[4]Fitch A W M.A Natural Classification of the Basic Helix-LoopHelix Class of Transcription Factors[J].Proceedings of the National Academy of Sciences of the United States of America,1997,94(10):5172-5176.
[5]Ledent V,Vervoort M.The Basic Helix-Loop-Helix Protein Family:Comparative Genomics and Phylogenetic Analysis[J].Genome Research,2001,11(5):754-770.
[6]Ledent,Valérie,Paquet O,Vervoort M.Phylogenetic analysis of the human basic helix-loop-helix proteins[J].Genome Biology,2002,3(6):1-18.
[7]Fisher A,Caudy M.The function of hairy-related bHLH repressor proteins in cell fate decisions.[J].Bioessays,2015,20(4):298-306.
[8]Buck M J,Atchley W R.Phylogenetic Analysis of Plant Basic Helix-Loop-Helix Proteins[J].Journal of Molecular Evolution,2003,56(6):742-750.
[9]Ramsay N A,Glover B J.MYB-bHLH-WD40 protein complex and the evolution of cellular diversity[J].Trends in Plant Science,2005,10(2):0-70.
[10]Simon L,Bousquet J,Lévesque R C,et al.Origin and diversification of endomycorrhizal fungi and coincidence with vascular land plants[J].Nature,1993,363(6424):67-69.
[11]Li X,Duan X,Jiang H,et al.Genome-Wide Analysis of Basic/Helix-Loop-Helix Transcription Factor Family in Rice and Arabidopsis[J].Plant Physiology,2006,141(4):1167-1184.
[12]Toledoortiz G,Huq E,Quail P H.The Arabidopsis Basic/HelixLoop-Helix Transcription Factor Family[J].Plant Cell,2003,15(8):1749-1770.
[13]Qi T,Huang H,Wu D,et al.Arabidopsis DELLA and JAZ Proteins Bind the WD-Repeat/bHLH/MYB Complex to Modulate Gibberellin and Jasmonate Signaling Synergy[J].The Plant Cell,2014,26(3):1118-1133.
[14]Nakata M,Mitsuda N,Herde M,et al.A bHLH-Type Transcription Factor,ABA-INDUCIBLE BHLH-TYPE TRANSCRIPTIONFACTOR/JA-ASSOCIATED MYC2-LIKE1,Acts as a Repressor to Negatively Regulate Jasmonate Signaling in Arabidopsis[J].The Plant Cell,2013,25(5):1641-1656.
[15]Fan M,Bai M Y,Kim J G,et al.The b HLH Transcription Factor HBI1 Mediates the Trade-Off between Growth and PathogenAssociated Molecular Pattern-Triggered Immunity in Arabidopsis[J].The Plant Cell,2014,26(2):828-841.
[16]Liu W,Tai H,Li S,et al.b HLH122 is important for drought and osmotic stress resistance in Arabidopsis and in the repression of ABA catabolism[J].New Phytologist,2014,201(4):1192-1204.
[17]周波,李玉花.植物的光敏色素与光信号转导[J].植物生理学通讯,2006(01):134-140.
[18]Kumar S V,Lucyshyn D,Jaeger K E,et al.Transcription factor PIF4 controls the thermosensory activation of flowering[J].NATURE,2012,484(7393):242-245.
[19]Khanna R,Huq E,Kikis E A,et al.A Novel Molecular Recognition Motif Necessary for Targeting Photoactivated Phytochrome Signaling to Specific Basic Helix-Loop-Helix Transcription Factors[J].Plant Cell,2004,16(11):3033-3044.
[20]Friedrichsen D M,Nemhauser J,Muramitsu T,et al.Three redundant brassinosteroid early response genes encode putative bHLH transcription factors required for normal growth[J].Genetics,2002,162(3):1445.
[21]Li X,Duan X,Jiang H,et al.Genome-Wide Analysis of Basic/Helix-Loop-Helix Transcription Factor Family in Rice and Arabidopsis[J].Plant Physiology,2006,141(4):1167-1184.
[22]刘厚生.棉花bHLH蛋白基因的克隆、表达及生物信息学分析[D].重庆:西南大学,2014.
[23]Morita Y,Saitoh M,Hoshino A,et al.Isolation of c DNAs for R2R3-MYB,bHLH and WDR Transcriptional Regulators and Identification of c and ca Mutations Conferring White Flowers in the Japanese Morning Glory[J].Plant&Cell Physiology,2006,47(4):457-470.
[24]Groszmann M,Bylstra Y,Lampugnani E R,et al.Regulation of tissue-specific expression of SPATULA,a bHLH gene involved in carpel development,seedling germination,and lateral organ growth in Arabidopsis[J].Journal of Experimental Botany,2010,61(5):1495-1508.
[25]Groszmann M,Paicu T,Smyth D R.Functional domains of SPATULA,a bHLH transcription factor involved in carpel and fruit development in Arabidopsis[J].Plant Journal,2010,55(1):40-52.
[26]Gremski K,Ditta G,Yanofsky M F.The HECATE genes regulate female reproductive tract developmentin Arabidopsis thaliana[J].Development,2007,134(20):3593-3601.
[27]Sanders P M,Bui A Q,Weterings K,et al.Anther developmental defects in Arabidopsis thaliana male-sterile mutants[J].Sexual Plant Reproduction,1999,11(6):297-322.
[28]Zhang W,Sun Y,Timofejeva L,et al.Regulation of Arabidopsis tapetum development and function by DYSFUNCTIONALTAPETUM1(DYT1)encoding a putative b HLH transcription factor[J].Development,2006,133(16):3085-3095.
[29]An X H,Tian Y,Chen K Q,et al.The apple WD40 protein MdTTG1 interacts with b HLH but not MYB proteins to regulate anthocyanin accumulation[J].Journal of Plant Physiology,2012,169(7):710-717.
[30]Larkin J C,Oppenheimer D G,Pollock S,et al.Arabidopsis GLABROUS1 Gene Requires Downstream Sequences for Function[J].Plant Cell,1993,5(12):1739-1748.
[31]Nesi N,Debeaujon I,Jond C,et al.The TT8 gene encodes a basic helix-loop-helix domain protein required for expression of DFR and BAN genes in Arabidopsis siliques[J].Plant Cell,2000,12(10):1863-1878.
[32]Payne C T,Zhang F,Lloyd A M.GL3 encodes a bHLH protein that regulates trichome development in arabidopsis through interaction with GL1 and TTG1[J].Genetics,2000,156(3):1349-1362.
[33]Smolen G A,Pawlowski L,Wilensky S E,et al.Dominant alleles of the basic helix-loop-helix transcription factor ATR2 activate stressresponsive genes in Arabidopsis[J].Genetics,2002,161(3):1235-1246.
[34]Ikeda M,Fujiwara S,Mitsuda N,et al.A triantagonistic basic helixloop-helix system regulates cell elongation in Arabidopsis[J].Plant Cell,2012,24(11):4483-4497.
[35]Zhang L Y,Bai M Y,Wu J,et al.Antagonistic HLH/bHLHtranscription factors mediate brassinosteroid regulation of cell elongation and plant development in rice and Arabidopsis[J].Plant Cell,2009,21(12):3767-3780.
[36]Yao N,Jing L,Zheng H,et al.Research progress of jasmonateresponsive transcription factors in regulating plant secondary metabolism[J].China journal of Chinese materia medica,2018,43(5):897.
[37]Jin-Ying G,Felippes F F,Chang-Jun L,et al.Negative regulation of anthocyanin biosynthesis in Arabidopsis by a miR156-targeted SPLtranscription factor[J].Plant Cell,2011,23(4):1512-1522.
[38]Sorensen,Anna-Marie,Kr?ber,Sandra,Unte U S,et al.The Arabidopsis ABORTED MICROSPORES(AMS)gene encodes a MYC class transcription factor[J].Plant Journal for Cell&Molecular Biology,2010,33(2):413-423.
[39]Thorstensen T,Grini P E,Mercy I S,et al.The Arabidopsis SET-domain protein ASHR3 is involved in stamen development and interacts with the bHLH transcription factor ABORTEDMICROSPORES(AMS)[J].Plant Molecular Biology,2008,66(1-2):47-59.
[40]Carreteropaulet L,Galstyan A,Roigvillanova I,et al.Genome-wide classification and evolutionary analysis of the bHLH family of transcription factors in Arabidopsis,poplar,rice,moss,and algae[J].Plant Physiology,2010,153(3):1398-1412.
[41]Seo J S,Joo J,Kim M J,et al.OsbHLH148,a basic helix-loophelix protein,interacts with OsJAZ proteins in a jasmonate signaling pathway leading to drought tolerance in rice[J].Plant Journal for Cell&Molecular Biology,2011,65(6):907-921.
[42]Kiribuchi K,Sugimori M,Takeda M,et al.RERJ1,a jasmonic acidresponsive gene from rice,encodes a basic helix-loop-helix protein[J].Biochemical&Biophysical Research Communications,2004,325(3):857-863.
[43]Kiribuchi K,Jikumaru Y,Kaku H,et al.Involvement of the basic helix-loop-helix transcription factor RERJ1 in wounding and drought stress responses in rice plants[J].Bioscience Biotechnology&Biochemistry,2005,69(5):1042-1044.
[44]Chen Z H,Nimmo G A,Jenkins G I,et al.b HLH32 modulates several biochemical and morphological processes that respond to Pi starvation in Arabidopsis[J].Biochemical Journal,2007,405(1):191-198.
[45]Ogo Y,Itai R N,Nakanishi H,et al.Isolation and characterization of IRO2,a novel iron-regulated b HLH transcription factor in graminaceous plants[J].Journal of Experimental Botany,2006,57(11):2867-2878.
[46]Wu J,Wang C,Zheng L,et al.Ethylene is involved in the regulation of iron homeostasis by regulating the expression of ironacquisition-related genes in Oryza sativa[J].Journal of Experimental Botany,2011,62(2):667-674.
[47]Chen C L,Cui Y,Cui M,et al.A FIT-binding protein is involved in modulating iron and zinc homeostasis in Arabidopsis[J].Plant Cell&Environment,2018,41(7):1698.
[48]Li L,Gao W,Peng Q,et al.Two soybean b HLH factors regulate response to iron deficiency[J].Journal of Integrative Plant Biology,2018.
[49]Zhao Q,Xiang X,Liu D,et al.Tobacco transcription factor NtbHLH123 confers tolerance to cold stress by regulating the NtCBF pathway and reactive oxygen species homeostasis[J].Frontiers in Plant Science,2018,9:381.
[50]Yao P,Sun Z,Li C,et al.Overexpression of Fagopyrum tataricum FtbHLH2 enhances tolerance to cold stress in transgenic Arabidopsis[J].Plant Physiol Biochem,2018,125:85-94.
[51]Kim J,Kim H Y.Functional analysis of a calcium-binding transcription factor involved in plant salt stress signaling[J].Febs Letters,2006,580(22):0-5256.
[52]于月华,王莉萍,高文伟,等.小麦盐胁迫相关基因的克隆与表达分析[J].西北植物学报,2012,32(06):1073-1078.
[53]Liu W,Tai H,Li S,et al.bHLH122 is important for drought and osmotic stress resistance in Arabidopsis and in the repression of ABA catabolism.[J].New Phytologist,2014,201(4):1192-1204.
[54]Jiang Y,Deyholos M K.Comprehensive transcriptional profiling of NaCl-stressed Arabidopsis roots reveals novel classes of responsive genes[J].BMC Plant Biology,2006,6(1):25-30.
[55]王昕嘉,李昆志.植物bHLH转录因子参与非生物胁迫信号通路研究进展[J].生命科学,2015,27(02):208-216.
[56]张菁,邹淑雪,张茂林,等.miR-15a靶基因的预测及生物信息学分析[J].生物技术通讯,2010,21(02):179-183.
[57]王勇,陈克平,姚勤.bHLH转录因子家族研究进展[J].遗传,2008,30(7):821-830.
[2]Li X,Duan X,Jiang H,et al.Genome-Wide Analysis of Basic/Helix-Loop-Helix Transcription Factor Family in Rice and Arabidopsis[J].Plant Physiology,2006,141(4):1167-1184.
[3]Chang W H,Lai A G.Genome-wide analyses of the bHLHsuperfamily in crustaceans:reappraisal of higher-order groupings and evidence for lineage-specific duplications[J].Royal Society Open Science,2018,5(3):172433.
[4]Fitch A W M.A Natural Classification of the Basic Helix-LoopHelix Class of Transcription Factors[J].Proceedings of the National Academy of Sciences of the United States of America,1997,94(10):5172-5176.
[5]Ledent V,Vervoort M.The Basic Helix-Loop-Helix Protein Family:Comparative Genomics and Phylogenetic Analysis[J].Genome Research,2001,11(5):754-770.
[6]Ledent,Valérie,Paquet O,Vervoort M.Phylogenetic analysis of the human basic helix-loop-helix proteins[J].Genome Biology,2002,3(6):1-18.
[7]Fisher A,Caudy M.The function of hairy-related bHLH repressor proteins in cell fate decisions.[J].Bioessays,2015,20(4):298-306.
[8]Buck M J,Atchley W R.Phylogenetic Analysis of Plant Basic Helix-Loop-Helix Proteins[J].Journal of Molecular Evolution,2003,56(6):742-750.
[9]Ramsay N A,Glover B J.MYB-bHLH-WD40 protein complex and the evolution of cellular diversity[J].Trends in Plant Science,2005,10(2):0-70.
[10]Simon L,Bousquet J,Lévesque R C,et al.Origin and diversification of endomycorrhizal fungi and coincidence with vascular land plants[J].Nature,1993,363(6424):67-69.
[11]Li X,Duan X,Jiang H,et al.Genome-Wide Analysis of Basic/Helix-Loop-Helix Transcription Factor Family in Rice and Arabidopsis[J].Plant Physiology,2006,141(4):1167-1184.
[12]Toledoortiz G,Huq E,Quail P H.The Arabidopsis Basic/HelixLoop-Helix Transcription Factor Family[J].Plant Cell,2003,15(8):1749-1770.
[13]Qi T,Huang H,Wu D,et al.Arabidopsis DELLA and JAZ Proteins Bind the WD-Repeat/bHLH/MYB Complex to Modulate Gibberellin and Jasmonate Signaling Synergy[J].The Plant Cell,2014,26(3):1118-1133.
[14]Nakata M,Mitsuda N,Herde M,et al.A bHLH-Type Transcription Factor,ABA-INDUCIBLE BHLH-TYPE TRANSCRIPTIONFACTOR/JA-ASSOCIATED MYC2-LIKE1,Acts as a Repressor to Negatively Regulate Jasmonate Signaling in Arabidopsis[J].The Plant Cell,2013,25(5):1641-1656.
[15]Fan M,Bai M Y,Kim J G,et al.The b HLH Transcription Factor HBI1 Mediates the Trade-Off between Growth and PathogenAssociated Molecular Pattern-Triggered Immunity in Arabidopsis[J].The Plant Cell,2014,26(2):828-841.
[16]Liu W,Tai H,Li S,et al.b HLH122 is important for drought and osmotic stress resistance in Arabidopsis and in the repression of ABA catabolism[J].New Phytologist,2014,201(4):1192-1204.
[17]周波,李玉花.植物的光敏色素与光信号转导[J].植物生理学通讯,2006(01):134-140.
[18]Kumar S V,Lucyshyn D,Jaeger K E,et al.Transcription factor PIF4 controls the thermosensory activation of flowering[J].NATURE,2012,484(7393):242-245.
[19]Khanna R,Huq E,Kikis E A,et al.A Novel Molecular Recognition Motif Necessary for Targeting Photoactivated Phytochrome Signaling to Specific Basic Helix-Loop-Helix Transcription Factors[J].Plant Cell,2004,16(11):3033-3044.
[20]Friedrichsen D M,Nemhauser J,Muramitsu T,et al.Three redundant brassinosteroid early response genes encode putative bHLH transcription factors required for normal growth[J].Genetics,2002,162(3):1445.
[21]Li X,Duan X,Jiang H,et al.Genome-Wide Analysis of Basic/Helix-Loop-Helix Transcription Factor Family in Rice and Arabidopsis[J].Plant Physiology,2006,141(4):1167-1184.
[22]刘厚生.棉花bHLH蛋白基因的克隆、表达及生物信息学分析[D].重庆:西南大学,2014.
[23]Morita Y,Saitoh M,Hoshino A,et al.Isolation of c DNAs for R2R3-MYB,bHLH and WDR Transcriptional Regulators and Identification of c and ca Mutations Conferring White Flowers in the Japanese Morning Glory[J].Plant&Cell Physiology,2006,47(4):457-470.
[24]Groszmann M,Bylstra Y,Lampugnani E R,et al.Regulation of tissue-specific expression of SPATULA,a bHLH gene involved in carpel development,seedling germination,and lateral organ growth in Arabidopsis[J].Journal of Experimental Botany,2010,61(5):1495-1508.
[25]Groszmann M,Paicu T,Smyth D R.Functional domains of SPATULA,a bHLH transcription factor involved in carpel and fruit development in Arabidopsis[J].Plant Journal,2010,55(1):40-52.
[26]Gremski K,Ditta G,Yanofsky M F.The HECATE genes regulate female reproductive tract developmentin Arabidopsis thaliana[J].Development,2007,134(20):3593-3601.
[27]Sanders P M,Bui A Q,Weterings K,et al.Anther developmental defects in Arabidopsis thaliana male-sterile mutants[J].Sexual Plant Reproduction,1999,11(6):297-322.
[28]Zhang W,Sun Y,Timofejeva L,et al.Regulation of Arabidopsis tapetum development and function by DYSFUNCTIONALTAPETUM1(DYT1)encoding a putative b HLH transcription factor[J].Development,2006,133(16):3085-3095.
[29]An X H,Tian Y,Chen K Q,et al.The apple WD40 protein MdTTG1 interacts with b HLH but not MYB proteins to regulate anthocyanin accumulation[J].Journal of Plant Physiology,2012,169(7):710-717.
[30]Larkin J C,Oppenheimer D G,Pollock S,et al.Arabidopsis GLABROUS1 Gene Requires Downstream Sequences for Function[J].Plant Cell,1993,5(12):1739-1748.
[31]Nesi N,Debeaujon I,Jond C,et al.The TT8 gene encodes a basic helix-loop-helix domain protein required for expression of DFR and BAN genes in Arabidopsis siliques[J].Plant Cell,2000,12(10):1863-1878.
[32]Payne C T,Zhang F,Lloyd A M.GL3 encodes a bHLH protein that regulates trichome development in arabidopsis through interaction with GL1 and TTG1[J].Genetics,2000,156(3):1349-1362.
[33]Smolen G A,Pawlowski L,Wilensky S E,et al.Dominant alleles of the basic helix-loop-helix transcription factor ATR2 activate stressresponsive genes in Arabidopsis[J].Genetics,2002,161(3):1235-1246.
[34]Ikeda M,Fujiwara S,Mitsuda N,et al.A triantagonistic basic helixloop-helix system regulates cell elongation in Arabidopsis[J].Plant Cell,2012,24(11):4483-4497.
[35]Zhang L Y,Bai M Y,Wu J,et al.Antagonistic HLH/bHLHtranscription factors mediate brassinosteroid regulation of cell elongation and plant development in rice and Arabidopsis[J].Plant Cell,2009,21(12):3767-3780.
[36]Yao N,Jing L,Zheng H,et al.Research progress of jasmonateresponsive transcription factors in regulating plant secondary metabolism[J].China journal of Chinese materia medica,2018,43(5):897.
[37]Jin-Ying G,Felippes F F,Chang-Jun L,et al.Negative regulation of anthocyanin biosynthesis in Arabidopsis by a miR156-targeted SPLtranscription factor[J].Plant Cell,2011,23(4):1512-1522.
[38]Sorensen,Anna-Marie,Kr?ber,Sandra,Unte U S,et al.The Arabidopsis ABORTED MICROSPORES(AMS)gene encodes a MYC class transcription factor[J].Plant Journal for Cell&Molecular Biology,2010,33(2):413-423.
[39]Thorstensen T,Grini P E,Mercy I S,et al.The Arabidopsis SET-domain protein ASHR3 is involved in stamen development and interacts with the bHLH transcription factor ABORTEDMICROSPORES(AMS)[J].Plant Molecular Biology,2008,66(1-2):47-59.
[40]Carreteropaulet L,Galstyan A,Roigvillanova I,et al.Genome-wide classification and evolutionary analysis of the bHLH family of transcription factors in Arabidopsis,poplar,rice,moss,and algae[J].Plant Physiology,2010,153(3):1398-1412.
[41]Seo J S,Joo J,Kim M J,et al.OsbHLH148,a basic helix-loophelix protein,interacts with OsJAZ proteins in a jasmonate signaling pathway leading to drought tolerance in rice[J].Plant Journal for Cell&Molecular Biology,2011,65(6):907-921.
[42]Kiribuchi K,Sugimori M,Takeda M,et al.RERJ1,a jasmonic acidresponsive gene from rice,encodes a basic helix-loop-helix protein[J].Biochemical&Biophysical Research Communications,2004,325(3):857-863.
[43]Kiribuchi K,Jikumaru Y,Kaku H,et al.Involvement of the basic helix-loop-helix transcription factor RERJ1 in wounding and drought stress responses in rice plants[J].Bioscience Biotechnology&Biochemistry,2005,69(5):1042-1044.
[44]Chen Z H,Nimmo G A,Jenkins G I,et al.b HLH32 modulates several biochemical and morphological processes that respond to Pi starvation in Arabidopsis[J].Biochemical Journal,2007,405(1):191-198.
[45]Ogo Y,Itai R N,Nakanishi H,et al.Isolation and characterization of IRO2,a novel iron-regulated b HLH transcription factor in graminaceous plants[J].Journal of Experimental Botany,2006,57(11):2867-2878.
[46]Wu J,Wang C,Zheng L,et al.Ethylene is involved in the regulation of iron homeostasis by regulating the expression of ironacquisition-related genes in Oryza sativa[J].Journal of Experimental Botany,2011,62(2):667-674.
[47]Chen C L,Cui Y,Cui M,et al.A FIT-binding protein is involved in modulating iron and zinc homeostasis in Arabidopsis[J].Plant Cell&Environment,2018,41(7):1698.
[48]Li L,Gao W,Peng Q,et al.Two soybean b HLH factors regulate response to iron deficiency[J].Journal of Integrative Plant Biology,2018.
[49]Zhao Q,Xiang X,Liu D,et al.Tobacco transcription factor NtbHLH123 confers tolerance to cold stress by regulating the NtCBF pathway and reactive oxygen species homeostasis[J].Frontiers in Plant Science,2018,9:381.
[50]Yao P,Sun Z,Li C,et al.Overexpression of Fagopyrum tataricum FtbHLH2 enhances tolerance to cold stress in transgenic Arabidopsis[J].Plant Physiol Biochem,2018,125:85-94.
[51]Kim J,Kim H Y.Functional analysis of a calcium-binding transcription factor involved in plant salt stress signaling[J].Febs Letters,2006,580(22):0-5256.
[52]于月华,王莉萍,高文伟,等.小麦盐胁迫相关基因的克隆与表达分析[J].西北植物学报,2012,32(06):1073-1078.
[53]Liu W,Tai H,Li S,et al.bHLH122 is important for drought and osmotic stress resistance in Arabidopsis and in the repression of ABA catabolism.[J].New Phytologist,2014,201(4):1192-1204.
[54]Jiang Y,Deyholos M K.Comprehensive transcriptional profiling of NaCl-stressed Arabidopsis roots reveals novel classes of responsive genes[J].BMC Plant Biology,2006,6(1):25-30.
[55]王昕嘉,李昆志.植物bHLH转录因子参与非生物胁迫信号通路研究进展[J].生命科学,2015,27(02):208-216.
[56]张菁,邹淑雪,张茂林,等.miR-15a靶基因的预测及生物信息学分析[J].生物技术通讯,2010,21(02):179-183.
[57]王勇,陈克平,姚勤.bHLH转录因子家族研究进展[J].遗传,2008,30(7):821-830.