参与植物非生物逆境响应的DREB/CBF转录因子研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2021-1219

摘要/Abstract

摘要：

DREB/CBF转录因子是植物AP2/ERF转录因子超家族的一个亚家族,具有AP2保守结构域,能特异性结合植物抗逆基因启动子区域DRE/CRT顺式作用元件。目前,越来越多研究表明,DREB/CBF转录因子参与调控植物响应干旱、盐、冷、热以及其他逆境胁迫的应答反应,是植物抗逆的关键调节因子。综述了近年来国内外DREB转录因子的研究进展,从DREB转录因子的结构、分类、家族及在非生物逆境胁迫中的功能等方面进行了总结,并结合当前研究的瓶颈,讨论了研究植物DREB/CBF转录因子的意义及存在的问题,并提出了今后的突破方向,以期为筛选优质抗逆植物资源,培育抗逆农林草新品种提供理论依据和指导。

关键词: 转录因子, DREB/CBF, 基因家族, 非生物胁迫

Abstract:

Dehydration responsive element binding（DREB）/CBF is a subfamily of the AP2/ERF（APETALA2/ethylene-responsive factor）superfamily in plant,having AP2 conserved domain and binding specifically to the dehydration response element/C-repeat（DRE/CRT）cis-elements（also known as the DNA sequence motif G/ACCGAC）found in the promoter region of stress resistance genes. Recent studies demonstrate that DREB transcription factors are involved in plant responses to drought,salt,cold,heat and other abiotic stresses,and is a key regulator in plant stress resistance. In this paper,recent progress in DREB is reviewed and several key aspects including structure,classification,family and function in abiotic stress are summarized. Combining the current bottlenecks of studying DREB transcription factors in plants,the prospects and issues in this field are also discussed. Finally,the breakthrough directions for future works are emphasized,aiming to provide a theoretical basis and guidance in screening high-quality plant resources and breeding novel varieties of crops,forests and grasses.

Key words: transcription factor, DREB/CBF, gene family, abiotic stress

刘坤, 李国婧, 杨杞. 参与植物非生物逆境响应的DREB/CBF转录因子研究进展[J]. 生物技术通报, 2022, 38(5): 201-214.

LIU Kun, LI Guo-jing, YANG Qi. Research Progress in DREB/CBF Transcription Factor Involved in Responses in Plant to Abiotic Stress[J]. Biotechnology Bulletin, 2022, 38(5): 201-214.

图/表 1

参考文献 110

[1]	苟艳丽, 张乐, 郭欢, 等. 植物AP2/ERF类转录因子研究进展[J]. 草业科学, 2020, 37(6):1150-1159.
	Gou YL, Zhang L, Guo H, et al. Research progress on the AP2/ERF transcription factor in plants[J]. Pratacultural Sci, 2020, 37(6):1150-1159.
[2]	李科友, 朱海兰. 植物非生物逆境胁迫DREB/CBF转录因子的研究进展[J]. 林业科学, 2011, 47(1):124-134.
	Li KY, Zhu HL. Research progress of DREB/CBF transcription factor in response to abiotic-stresses in plants[J]. Sci Silvae Sin, 2011, 47(1):124-134.
[3]	张麒, 陈静, 李俐, 等. 植物AP2/ERF转录因子家族的研究进展[J]. 生物技术通报, 2018, 34(8):1-7. doi: 10.13560/j.cnki.biotech.bull.1985.2017-1142
	Zhang Q, Chen J, Li L, et al. Research progress on plant AP2/ERF transcription factor family[J]. Biotechnol Bull, 2018, 34(8):1-7.
[4]	柏星轩, 闫雪, 要宇晨, 等. DREB/CBF转录因子在植物非生物胁迫中的作用及研究进展[J]. 生物学杂志, 2017, 34(4):88-93.
	Bai XX, Yan X, Yao YC, et al. The role and research progress of DREB/CBF transcription factors in plant abiotic stress[J]. J Biol, 2017, 34(4):88-93.
[5]	Li MY, Xu ZS, Huang Y, et al. Genome-wide analysis of AP2/ERF transcription factors in carrot(Daucus carota L.)reveals evolution and expression profiles under abiotic stress[J]. Mol Genet Genom, 2015, 290(6):2049-2061. doi: 10.1007/s00438-015-1061-3 URL
[6]	Licausi F, Ohme-Takagi M, Perata P. APETALA2/Ethylene responsive factor(AP2/ERF)transcription factors:mediators of stress responses and developmental programs[J]. New Phytol, 2013, 199(3):639-649. doi: 10.1111/nph.12291 URL
[7]	Allen MD, Yamasaki K, Ohme-Takagi M, et al. A novel mode of DNA recognition by a beta-sheet revealed by the solution structure of the GCC-box binding domain in complex with DNA[J]. EMBO J, 1998, 17(18):5484-5496. pmid: 9736626
[8]	Okamuro JK, Caster B, Villarroel R, et al. The AP2 domain of APETALA2 defines a large new family of DNA binding proteins in Arabidopsis[J]. PNAS, 1997, 94(13):7076-7081. pmid: 9192694
[9]	Sakuma Y, Liu Q, Dubouzet JG, et al. DNA-binding specificity of the ERF/AP2 domain of Arabidopsis DREBs, transcription factors involved in dehydration- and cold-inducible gene expression[J]. Biochem Biophys Res Commun, 2002, 290(3):998-1009. doi: 10.1006/bbrc.2001.6299 URL
[10]	Amalraj A, Luang S, Kumar MY, et al. Change of function of the wheat stress-responsive transcriptional repressor TaRAP2. 1L by repressor motif modification[J]. Plant Biotechnol J, 2016, 14(2):820-832. doi: 10.1111/pbi.12432 URL
[11]	Eini O, Yang N, Pyvovarenko T, et al. Complex regulation by Apetala2 domain-containing transcription factors revealed through analysis of the stress-responsive TdCor410b promoter from durum wheat[J]. PLoS One, 2013, 8(3):e58713. doi: 10.1371/journal.pone.0058713 URL
[12]	Xue GP. Characterisation of the DNA-binding profile of barley HvCBF₁ using an enzymatic method for rapid, quantitative and high-throughput analysis of the DNA-binding activity[J]. Nucleic Acids Res, 2002, 30(15):e77. doi: 10.1093/nar/gnf076 URL
[13]	Lata C, Prasad M. Role of DREBs in regulation of abiotic stress responses in plants[J]. J Exp Bot, 2011, 62(14):4731-4748. doi: 10.1093/jxb/err210 URL
[14]	Ohme-Takagi M, Shinshi H. Ethylene-inducible DNA binding proteins that interact with an ethylene-responsive element[J]. Plant Cell, 1995, 7(2):173-182. pmid: 7756828
[15]	Nakano T, Suzuki K, Fujimura T, et al. Genome-wide analysis of the ERF gene family in Arabidopsis and rice[J]. Plant Physiol, 2006, 140(2):411-432. doi: 10.1104/pp.105.073783 URL
[16]	Jaglo KR, Kleff S, Amundsen KL, et al. Components of the Arabidopsis C-repeat/dehydration-responsive element binding factor cold-response pathway are conserved in Brassica napus and other plant species[J]. Plant Physiol, 2001, 127(3):910-917. pmid: 11706173
[17]	Agarwal PK, Agarwal P, Reddy MK, et al. Role of DREB transcription factors in abiotic and biotic stress tolerance in plants[J]. Plant Cell Rep, 2006, 25(12):1263-1274. doi: 10.1007/s00299-006-0204-8 URL
[18]	Xiong Y, Fei SZ. Functional and phylogenetic analysis of a DREB/CBF-like gene in perennial ryegrass(Lolium perenne L.)[J]. Planta, 2006, 224(4):878-888. doi: 10.1007/s00425-006-0273-5 URL
[19]	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-temperature-responsive gene expression, respectively, in Arabidopsis[J]. Plant Cell, 1998, 10(8):1391-1406. pmid: 9707537
[20]	Gilmour SJ, Zarka DG, Stockinger EJ, et al. Low temperature regulation of the Arabidopsis CBF family of AP2 transcriptional activators as an early step in cold-induced COR gene expression[J]. Plant J, 1998, 16(4):433-442. pmid: 9881163
[21]	Fowler S, Thomashow MF. Arabidopsis transcriptome profiling indicates that multiple regulatory pathways are activated during cold acclimation in addition to the CBF cold response pathway[J]. Plant Cell, 2002, 14(8):1675-1690. pmid: 12172015
[22]	Haake V, Cook D, Riechmann JL, et al. Transcription factor CBF4 is a regulator of drought adaptation in Arabidopsis[J]. Plant Physiol, 2002, 130(2):639-648. doi: 10.1104/pp.006478 URL
[23]	Magome H, Yamaguchi S, Hanada A, et al. Dwarf and delayed-flowering 1, a novel Arabidopsis mutant deficient in gibberellin biosynthesis because of overexpression of a putative AP2 transcription factor[J]. Plant J, 2004, 37(5):720-729. doi: 10.1111/j.1365-313X.2003.01998.x URL
[24]	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[J]. Mol Genet Genom, 2010, 283(2):185-196. doi: 10.1007/s00438-009-0506-y URL
[25]	祝娟娟. 桑树DREB基因家族生物信息学分析及功能研究[D]. 重庆: 西南大学, 2013.
	Zhu JJ. Bioinformatics analysis and functional study of DREB genes from Morus spp[D]. Chongqing: Southwest University, 2013.
[26]	Wilson K, Long D, Swinburne J, et al. A Dissociation insertion causes a semidominant mutation that increases expression of TINY, an Arabidopsis gene related to APETALA2[J]. Plant Cell, 1996, 8(4):659-671. pmid: 8624440
[27]	Song X, Li Y, Hou X. Genome-wide analysis of the AP2/ERF transcription factor superfamily in Chinese cabbage(Brassica rapa ssp. pekinensis)[J]. BMC Genomics, 2013, 14:573. doi: 10.1186/1471-2164-14-573 URL
[28]	Liu Z, Kong L, Zhang M, et al. Genome-wide identification, phylogeny, evolution and expression patterns of AP2/ERF genes and cytokinin response factors in Brassica rapa ssp. pekinensis[J]. PLoS One, 2013, 8(12):e83444. doi: 10.1371/journal.pone.0083444 URL
[29]	邵文靖, 敖特根白音, 郎明林. AP2/ERF转录因子对植物非生物胁迫的应答机制研究进展[J]. 分子植物育种, 2020, 18(15):4981-4988.
	Shao WJ, Ao T, Lang ML. Research advances on the mechanism of AP2/ERF transcriptional factors in response to abiotic stresses in plants[J]. Mol Plant Breed, 2020, 18(15):4981-4988.
[30]	Charfeddine M, Saïdi MN, Charfeddine S, et al. Genome-wide analysis and expression profiling of the ERF transcription factor family in potato(Solanum tuberosum L.)[J]. Mol Biotechnol, 2015, 57(4):348-358. doi: 10.1007/s12033-014-9828-z pmid: 25491236
[31]	Sharma MK, Kumar R, Solanke AU, et al. Identification, phylogeny, and transcript profiling of ERF family genes during development and abiotic stress treatments in tomato[J]. Mol Genet Genomics, 2010, 284(6):455-475. doi: 10.1007/s00438-010-0580-1 pmid: 20922546
[32]	Sharoni AM, Nuruzzaman M, Satoh K, et al. Gene structures, classification and expression models of the AP2/EREBP transcription factor family in rice[J]. Plant Cell Physiol, 2011, 52(2):344-360. doi: 10.1093/pcp/pcq196 pmid: 21169347
[33]	Zhuang J, Deng DX, Yao QH, et al. Discovery, phylogeny and expression patterns of AP2-like genes in maize[J]. Plant Growth Regul, 2010, 62(1):51-58. doi: 10.1007/s10725-010-9484-7 URL
[34]	Zhuang J, Chen JM, Yao QH, et al. Discovery and expression profile analysis of AP2/ERF family genes from Triticum aestivum[J]. Mol Biol Rep, 2011, 38(2):745-753. doi: 10.1007/s11033-010-0162-7 pmid: 20407836
[35]	Zhuang J, Anyia A, Vidmar J, et al. Discovery and expression assessment of the AP2-like genes in Hordeum vulgare[J]. Acta Physiol Plant, 2011, 33(5):1639-1649. doi: 10.1007/s11738-010-0700-x URL
[36]	Huang Z, Zhong XJ, He J, et al. Identification and characterization of AP2/ERF transcription factors in moso bamboo(Phyllostachys edulis)[J]. Mol Biol:Mosk, 2016, 50(5):785-796.
[37]	Chen L, Han J, Deng X, et al. Expansion and stress responses of AP2/EREBP superfamily in Brachypodium distachyon[J]. Sci Rep, 2016, 6:21623. doi: 10.1038/srep21623 URL
[38]	Yan HW, Hong L, Zhou YQ, et al. A genome-wide analysis of the ERF gene family in Sorghum[J]. Genet Mol Res, 2013, 12(2):2038-2055. doi: 10.4238/2013.May.13.1 pmid: 23766026
[39]	Agarwal G, Garg V, Kudapa H, et al. Genome-wide dissection of AP2/ERF and HSP90 gene families in five legumes and expression profiles in chickpea and pigeonpea[J]. Plant Biotechnol J, 2016, 14(7):1563-1577. doi: 10.1111/pbi.12520 pmid: 26800652
[40]	Song X, Wang J, Ma X, et al. Origination, expansion, evolutionary trajectory, and expression bias of AP2/ERF superfamily in Brassica napus[J]. Front Plant Sci, 2016, 7:1186.
[41]	Thamilarasan SK, Park JI, Jung HJ, et al. Genome-wide analysis of the distribution of AP2/ERF transcription factors reveals duplication and CBFs genes elucidate their potential function in Brassica oleracea[J]. BMC Genomics, 2014, 15:422. doi: 10.1186/1471-2164-15-422 pmid: 24888752
[42]	Zhuang J, Cai B, Peng RH, et al. Genome-wide analysis of the AP2/ERF gene family in Populus trichocarpa[J]. Biochem Biophys Res Commun, 2008, 371(3):468-474. doi: 10.1016/j.bbrc.2008.04.087 URL
[43]	Zhang J, Shi SZ, Jiang Y, et al. Genome-wide investigation of the AP2/ERF superfamily and their expression under salt stress in Chinese willow(Salix matsudana)[J]. PeerJ, 2021, 9:e11076. doi: 10.7717/peerj.11076 URL
[44]	Xu W, Li F, Ling L, et al. Genome-wide survey and expression profiles of the AP2/ERF family in Castor bean(Ricinus communis L.)[J]. BMC Genomics, 2013, 14:785. doi: 10.1186/1471-2164-14-785 URL
[45]	Zhang H, Pan X, Liu S, et al. Genome-wide analysis of AP2/ERF transcription factors in pineapple reveals functional divergence during flowering induction mediated by ethylene and floral organ development[J]. Genomics, 2021, 113(2):474-489. doi: 10.1016/j.ygeno.2020.10.040 URL
[46]	Zhuang J, Peng RH, Cheng ZM, et al. Genome-wide analysis of the putative AP2/ERF family genes in Vitis vinifera[J]. Sci Hortic, 2009, 123(1):73-81. doi: 10.1016/j.scienta.2009.08.002 URL
[47]	Sun ZM, Zhou ML, Xiao XG, et al. Genome-wide analysis of AP2/ERF family genes from Lotus corniculatus shows LcERF054 enhances salt tolerance[J]. Funct Integr Genomics, 2014, 14(3):453-466. doi: 10.1007/s10142-014-0372-5 URL
[48]	Chinnusamy V, Ohta M, Kanrar S, et al. ICE1:a regulator of cold-induced transcriptome and freezing tolerance in Arabidopsis[J]. Genes Dev, 2003, 17(8):1043-1054. doi: 10.1101/gad.1077503 URL
[49]	Chinnusamy V, Zhu J, Zhu JK. Cold stress regulation of gene expression in plants[J]. Trends Plant Sci, 2007, 12(10):444-451. doi: 10.1016/j.tplants.2007.07.002 URL
[50]	Agarwal M, Hao Y, Kapoor A, et al. A R2R3 type MYB transcription factor is involved in the cold regulation of CBF genes and in acquired freezing tolerance[J]. J Biol Chem, 2006, 281(49):37636-37645. doi: 10.1074/jbc.M605895200 pmid: 17015446
[51]	Shi Y, Ding Y, Yang S. Molecular regulation of CBF signaling in cold acclimation[J]. Trends Plant Sci, 2018, 23(7):623-637. doi: 10.1016/j.tplants.2018.04.002 URL
[52]	Ding YL, Jia YX, Shi YT, et al. OST 1-mediated BTF 3L phosphorylation positively regulates CBF s during plant cold responses[J]. EMBO J, 2018, 37(8):e98228.
[53]	Liu Z, Jia Y, Ding Y, et al. Plasma membrane CRPK1-mediated phosphorylation of 14-3-3 proteins induces their nuclear import to fine-tune CBF signaling during cold response[J]. Mol Cell, 2017, 66(1):117-128.e5. doi: 10.1016/j.molcel.2017.02.016 URL
[54]	Wang X, Ding Y, Li Z, et al. PUB25 and PUB26 promote plant freezing tolerance by degrading the cold signaling negative regulator MYB15[J]. Dev Cell, 2019, 51(2):222-235.e5. doi: S1534-5807(19)30691-4 pmid: 31543444
[55]	Seki M, Narusaka M, Abe H, et al. Monitoring the expression pattern of 1300 Arabidopsis genes under drought and cold stresses by using a full-length cDNA microarray[J]. Plant Cell, 2001, 13(1):61-72. pmid: 11158529
[56]	刘坤, 王光霞, 王瑞刚, 等. 中间锦鸡儿CiDREB1C基因增强转基因拟南芥抵抗非生物胁迫的能力[J]. 农业生物技术学报, 2018, 26(10):1688-1697.
	Liu K, Wang GX, Wang RG, et al. CiDREB1C gene from Caragana intermedia enhances abiotic stress tolerance of transgenic Arabidopsis thaliana[J]. J Agric Biotechnol, 2018, 26(10):1688-1697.
[57]	Zhang ZY, Yang Q, Zhang CL, et al. A CkDREB1 gene isolated from Caragana korshinskii Kom. enhances Arabidopsis drought and cold tolerance[J]. Braz J Bot, 2019, 42(1):97-105. doi: 10.1007/s40415-018-0509-1 URL
[58]	Medina J, Bargues M, Terol J, et al. The Arabidopsis CBF gene family is composed of three genes encoding AP2 domain-containing proteins whose expression Is regulated by low temperature but not by abscisic acid or dehydration[J]. Plant Physiol, 1999, 119(2):463-470. pmid: 9952441
[59]	Zhao C, Zhang Z, Xie S, et al. Mutational evidence for the critical role of CBF transcription factors in cold acclimation in Arabidopsis[J]. Plant Physiol, 2016, 171(4):2744-2759. doi: 10.1104/pp.16.00533 URL
[60]	Jia Y, Ding Y, Shi Y, et al. The cbfs triple mutants reveal the essential functions of CBFs in cold acclimation and allow the definition of CBF regulons in Arabidopsis[J]. New Phytol, 2016, 212(2):345-353. doi: 10.1111/nph.14088 URL
[61]	Novillo F, Alonso JM, Ecker JR, et al. CBF2/DREB1C is a negative regulator of CBF1/DREB1B and CBF3/DREB1A expression and plays a central role in stress tolerance in Arabidopsis[J]. PNAS, 2004, 101(11):3985-3990. doi: 10.1073/pnas.0303029101 URL
[62]	Novillo F, Medina J, Salinas J. Arabidopsis CBF1 and CBF3 have a different function than CBF2 in cold acclimation and define different gene classes in the CBF regulon[J]. PNAS, 2007, 104(52):21002-21007. doi: 10.1073/pnas.0705639105 URL
[63]	Hua J. Defining roles of tandemly arrayed CBF genes in freezing tolerance with new genome editing tools[J]. New Phytol, 2016, 212(2):301-302. doi: 10.1111/nph.14183 URL
[64]	Sakuma Y, Maruyama K, Osakabe Y, et al. Functional analysis of an Arabidopsis transcription factor, DREB2A, involved in drought-responsive gene expression[J]. Plant Cell, 2006, 18(5):1292-1309. doi: 10.1105/tpc.105.035881 pmid: 16617101
[65]	Agarwal P, Agarwal PK, Nair S, et al. Stress-inducible DREB2A transcription factor from Pennisetum glaucum is a phosphoprotein and its phosphorylation negatively regulates its DNA-binding activity[J]. Mol Genet Genom, 2007, 277(2):189-198. doi: 10.1007/s00438-006-0183-z URL
[66]	Chen J, Xia X, Yin W. Expression profiling and functional characterization of a DREB2-type gene from Populus euphratica[J]. Biochem Biophys Res Commun, 2009, 378(3):483-487. doi: 10.1016/j.bbrc.2008.11.071 URL
[67]	Egawa C, Kobayashi F, Ishibashi M, et al. Differential regulation of transcript accumulation and alternative splicing of a DREB2 homolog under abiotic stress conditions in common wheat[J]. Genes Genet Syst, 2006, 81(2):77-91. doi: 10.1266/ggs.81.77 URL
[68]	Qin F, Kakimoto M, Sakuma Y, et al. Regulation and functional analysis of ZmDREB2A in response to drought and heat stresses in Zea mays L[J]. Plant J, 2007, 50(1):54-69. doi: 10.1111/j.1365-313X.2007.03034.x URL
[69]	秦红霞, 贾志平, 张海超, 等. 银新杨中与DRE元件结合的转录因子的克隆及鉴定分析[J]. 生物工程学报, 2005, 21(6):906-910
	Qin HX, Jia ZP, Zhang HC, et al. Isolation and Characterization of a DRE-binding Transcription Factor from Yinxin Poplar(Populus alba × P. alba var. pyramidalis)[J]. Chin J Biotechnol, 2005, 21(6):906-910
[70]	谢永丽, 王自章, 刘强, 等. 草坪草狗牙根中抗逆基因BeDREB的克隆及功能鉴定[J]. 中国生物化学与分子生物学报, 2005, 21(4):521-527.
	Xie YL, Wang ZZ, Liu Q, et al. Cloning and functional identification of stress-resistant BeDREB genes from Bermuda grass[J]. Chin J Biochem Mol Biol, 2005, 21(4):521-527.
[71]	Sun S, Yu JP, Chen F, et al. Tiny, a Dehydration-responsive element(dre)-binding protein-like transcription factor connecting the dre- and ethylene-responsive element-mediated signaling pathways in Arabidopsis[J]. J Biol Chem, 2008, 283(10):6261-6271. doi: 10.1074/jbc.M706800200 URL
[72]	Chen M, Wang QY, Cheng XG, et al. GmDREB2, a soybean DRE-binding transcription factor, conferred drought and high-salt tolerance in transgenic plants[J]. Biochem Biophys Res Commun, 2007, 353(2):299-305. doi: 10.1016/j.bbrc.2006.12.027 URL
[73]	Chen M, Xu Z, Xia L, et al. Cold-induced modulation and functional analyses of the DRE-binding transcription factor gene, GmDREB3, in soybean(Glycine max L.)[J]. J Exp Bot, 2009, 60(1):121-135. doi: 10.1093/jxb/ern269 URL
[74]	Figueroa-Yañez L, Pereira-Santana A, Arroyo-Herrera A, et al. RAP2. 4a is transported through the phloem to regulate cold and heat tolerance in Papaya tree(Carica papaya cv. maradol):implications for protection against abiotic stress[J]. PLoS One, 2016, 11(10):e0165030. doi: 10.1371/journal.pone.0165030 URL
[75]	Qin F, Sakuma Y, Tran LS, et al. Arabidopsis DREB2A-interacting proteins function as RING E3 ligases and negatively regulate plant drought stress-responsive gene expression[J]. Plant Cell, 2008, 20(6):1693-1707. doi: 10.1105/tpc.107.057380 URL
[76]	Rogers S, Wells R, Rechsteiner M. Amino acid sequences common to rapidly degraded proteins:the PEST hypothesis[J]. Science, 1986, 234(4774):364-368. doi: 10.1126/science.2876518 pmid: 2876518
[77]	Zhou ML, Ma JT, Pang JF, et al. Regulation of plant stress response by dehydration responsive element binding(DREB)transcription factors[J]. Afr. J. Biotechnol, 2010, 9(54):9255-9269.
[78]	Salmerón A, Janzen J, Soneji Y, et al. Direct phosphorylation of NF-kappaB1 p105 by the IkappaB kinase complex on serine 927 is essential for signal-induced p105 proteolysis[J]. J Biol Chem, 2001, 276(25):22215-22222. doi: 10.1074/jbc.M101754200 pmid: 11297557
[79]	Agarwal PK, Gupta K, Lopato S, et al. Dehydration responsive element binding transcription factors and their applications for the engineering of stress tolerance[J]. J Exp Bot, 2017, 68(9):2135-2148. doi: 10.1093/jxb/erx118 URL
[80]	Morimoto K, Ohama N, Kidokoro S, et al. BPM-CUL3 E3 ligase modulates thermotolerance by facilitating negative regulatory domain-mediated degradation of DREB2A in Arabidopsis[J]. PNAS, 2017, 114(40):E8528-E8536.
[81]	Kudo M, Kidokoro S, Yoshida T, et al. Double overexpression of DREB and PIF transcription factors improves drought stress tolerance and cell elongation in transgenic plants[J]. Plant Biotechnol J, 2017, 15(4):458-471. doi: 10.1111/pbi.12644 URL
[82]	Kidokoro S, Watanabe K, Ohori T, et al. Soybean DREB1/CBF-type transcription factors function in heat and drought as well as cold stress-responsive gene expression[J]. Plant J, 2015, 81(3):505-518. doi: 10.1111/tpj.12746 URL
[83]	Engels C, Fuganti-Pagliarini R, Marin SRR, et al. Introduction of the rd29A:AtDREB2A CA gene into soybean(Glycine max L. Merril)and its molecular characterization in leaves and roots during dehydration[J]. Genet Mol Biol, 2013, 36(4):556-565. doi: 10.1590/S1415-47572013000400015 URL
[84]	Mizoi J, Ohori T, Moriwaki T, et al. GmDREB2A;2, a canonical DEHYDRATION-RESPONSIVE ELEMENT-BINDING PROTEIN2-type transcription factor in soybean, is posttranslationally regulated and mediates dehydration-responsive element-dependent gene expression[J]. Plant Physiol, 2013, 161(1):346-361. doi: 10.1104/pp.112.204875 URL
[85]	Reis RR, da Cunha BA, Martins PK, et al. Induced over-expression of AtDREB2A CA improves drought tolerance in sugarcane[J]. Plant Sci, 2014, 221/222:59-68. doi: 10.1016/j.plantsci.2014.02.003 URL
[86]	Souza WR, Oliveira NG, Vinecky F, et al. Field evaluation of AtDR-EB 2A CA overexpressing sugarcane for drought tolerance[J]. J Agro Crop Sci, 2019, 205(6):545-553. doi: 10.1111/jac.12341 URL
[87]	Schmidt R, Mieulet D, Hubberten HM, et al. Salt-responsive ERF1 regulates reactive oxygen species-dependent signaling during the initial response to salt stress in rice[J]. Plant Cell, 2013, 25(6):2115-2131. doi: 10.1105/tpc.113.113068 URL
[88]	Dubouzet JG, Sakuma Y, Ito Y, et al. OsDREB genes in rice, Oryza sativa L., encode transcription activators that function in drought-, high-salt- and cold-responsive gene expression[J]. Plant J, 2003, 33(4):751-763. pmid: 12609047
[89]	Gutha LR, Reddy AR. Rice DREB1B promoter shows distinct stress-specific responses, and the overexpression of cDNA in tobacco confers improved abiotic and biotic stress tolerance[J]. Plant Mol Biol, 2008, 68(6):533-555. doi: 10.1007/s11103-008-9391-8 URL
[90]	Wei T, Deng K, Zhang Q, et al. Modulating AtDREB1C expression improves drought tolerance in Salvia miltiorrhiza[J]. Front Plant Sci, 2017, 8:52.
[91]	An D, Ma Q, Wang H, et al. Cassava C-repeat binding factor 1 gene responds to low temperature and enhances cold tolerance when overexpressed in Arabidopsis and cassava[J]. Plant Mol Biol, 2017, 94(1/2):109-124. doi: 10.1007/s11103-017-0596-6 URL
[92]	Bouaziz D, Pirrello J, Ben Amor H, et al. Ectopic expression of dehydration responsive element binding proteins(StDREB2)confers higher tolerance to salt stress in potato[J]. Plant Physiol Biochem, 2012, 60:98-108. doi: 10.1016/j.plaphy.2012.07.029 URL
[93]	Bouaziz D, Pirrello J, Charfeddine M, et al. Overexpression of StDREB1 transcription factor increases tolerance to salt in transgenic potato plants[J]. Mol Biotechnol, 2013, 54(3):803-817. doi: 10.1007/s12033-012-9628-2 pmid: 23250722
[94]	Huang B, Liu JY. Cloning and functional analysis of the novel gene GhDBP3 encoding a DRE-binding transcription factor from Gossypium hirsutum[J]. Biochim et Biophys Acta BBA Gene Struct Expr, 2006, 1759(6):263-269.
[95]	Liang YQ, Li XS, Yang RR, et al. BaDBL1, a unique DREB gene from desiccation tolerant moss Bryum argenteum, confers osmotic and salt stress tolerances in transgenic Arabidopsis[J]. Plant Sci, 2021, 313:111047. doi: 10.1016/j.plantsci.2021.111047 URL
[96]	Liang Y, Li X, Zhang D, et al. ScDREB8, a novel A-5 type of DREB gene in the desert moss Syntrichia caninervis, confers salt tolerance to Arabidopsis[J]. Plant Physiol Biochem, 2017, 120:242-251. doi: 10.1016/j.plaphy.2017.09.014 URL
[97]	Li XS, Liang YQ, Gao B, et al. ScDREB10, an A-5c type of DREB gene of the desert moss Syntrichia caninervis, confers osmotic and salt tolerances to Arabidopsis[J]. Genes, 2019, 10(2):146. doi: 10.3390/genes10020146 URL
[98]	Chen JR, Lü JJ, Wang TX, et al. Activation of a DRE-binding transcription factor from Medicago truncatula by deleting a Ser/Thr-rich region[J]. Vitro Cell Dev Biol Plant, 2009, 45(1):1-11.
[99]	Lim CJ, Hwang JE, Chen H, et al. Over-expression of the Arabidopsis DRE/CRT-binding transcription factor DREB2C enhances thermotolerance[J]. Biochem Biophys Res Commun, 2007, 362(2):431-436. doi: 10.1016/j.bbrc.2007.08.007 URL
[100]	Singh S, Chopperla R, Shingote P, et al. Overexpression of EcDREB2A transcription factor from finger millet in tobacco enhances tolerance to heat stress through ROS scavenging[J]. J Biotechnol, 2021, 336:10-24. doi: 10.1016/j.jbiotec.2021.06.013 URL
[101]	Mao L, Deng M, Jiang S, et al. Characterization of the DREBA4-type transcription factor(SlDREBA4), which contributes to heat tolerance in tomatoes[J]. Front Plant Sci, 2020, 11:554520. doi: 10.3389/fpls.2020.554520 URL
[102]	Du X, Li W, Sheng L, et al. Over-expression of Chrysanthemum CmDREB6 enhanced tolerance of Chrysanthemum to heat stress[J]. BMC Plant Biol, 2018, 18(1):178. doi: 10.1186/s12870-018-1400-8 URL
[103]	Zhang Z, Li W, Gao X, et al. DEAR4, a member of DREB/CBF family, positively regulates leaf senescence and response to multiple stressors in Arabidopsis thaliana[J]. Front Plant Sci, 2020, 11:367. doi: 10.3389/fpls.2020.00367 URL
[104]	Liu XQ, Liu CY, Guo Q, et al. Mulberry transcription factor MnDREB4A confers tolerance to multiple abiotic stresses in transgenic tobacco[J]. PLoS One, 2015, 10(12):e0145619. doi: 10.1371/journal.pone.0145619 URL
[105]	Sharabi-Schwager M, Lers A, Samach A, et al. Overexpression of the CBF2 transcriptional activator in Arabidopsis delays leaf senescence and extends plant longevity[J]. J Exp Bot, 2010, 61(1):261-273. doi: 10.1093/jxb/erp300 pmid: 19854800
[106]	Porat R, Sharabi-Schwager M, Samach A. Overexpression of the CBF₂ transcriptional activator enhances oxidative stress tolerance in Arabidopsis plants[J]. Int J Biol, 2011, 3(2):94.
[107]	Zwack PJ, Robinson BR, Risley MG, et al. Cytokinin response factor 6 negatively regulates leaf senescence and is induced in response to cytokinin and numerous abiotic stresses[J]. Plant Cell Physiol, 2013, 54(6):971-981. doi: 10.1093/pcp/pct049 pmid: 23539244
[108]	Ban Q, Liu G, Wang Y. A DREB gene from Limonium bicolor mediates molecular and physiological responses to copper stress in transgenic tobacco[J]. J Plant Physiol, 2011, 168(5):449-458. doi: 10.1016/j.jplph.2010.08.013 URL
[109]	Hong JP, Kim WT. Isolation and functional characterization of the Ca-DREBLP1 gene encoding a dehydration-responsive element binding-factor-like protein 1 in hot pepper(Capsicum annuum L. cv. Pukang)[J]. Planta, 2005, 220(6):875-888. doi: 10.1007/s00425-004-1412-5 URL
[110]	Du HW, Huang M, Zhang ZX, et al. Genome-wide analysis of the AP2/ERF gene family in maize waterlogging stress response[J]. Euphytica, 2014, 198(1):115-126. doi: 10.1007/s10681-014-1088-2 URL