板栗WRKY基因家族鉴定及其在干旱胁迫下的表达分析

doi:10.7668/hbnxb.20194000

摘要/Abstract

摘要：

WRKY转录因子在植物生长和防御反应的调控中发挥着重要作用。为了探究WRKY基因家族在板栗抗旱中的功能,对板栗WRKY基因家族成员进行了鉴定,对其编码蛋白的理化性质、系统发育、结构等进行分析,并通过转录组测序和qRT-PCR技术分析其在干旱胁迫下的表达特征。通过生物信息学技术在板栗中共鉴定到65个WRKY基因家族成员,将其分为Ⅰ、Ⅱ、Ⅲ 3组,其中Ⅱ组根据其结构和系统发育关系分为5个亚组。WRKY基因的结构和保守基序分析表明,外显子数目为1~7个,同一亚组的外显子数目和基序分布类似。WRKY基因保守结构域发生了一定的变异,包括WRKY七肽结构域的缺失,锌指结构的缺失,七肽结构域变异为WRKYGKK、WRKYGRK和WRKYGRK等。转录组测序数据表明,有4个WRKY基因家族成员在样品中没有表达,干旱胁迫诱导表达的WRKY基因家族成员表达量上调的有49个,而表达量下调的WRKY家族成员有11个。以上结果表明,板栗WRKY基因在对干旱胁迫的响应中发挥了重要作用。

关键词: 板栗, 基因家族, WRKY, 干旱胁迫

Abstract:

WRKY transcription factors play an important role in the regulation of plant growth and defence responses.In order to explore the function of WRKY gene family in chestnut under drought resistance,the chestnut WRKY gene family members were identified and the physical and chemical properties,phylogeny and structure of the encoded proteins were analyzed.Transcriptome sequencing and qRT-PCR were used to analyze the expression characteristics of WRKY gene family members under drought stress.The results showed that 65 WRKY gene family members were identified and divided into three groups:Ⅰ,Ⅱ and Ⅲ,among which group Ⅱ was divided into five subgroups according to its structure and phylogenetic relationship.Analysis of WRKY gene structure and conserved motif showed that the number of exons ranged from 1 to 7,and the number of exons and motif distribution in the same subgroup were similar.The conserved domain of WRKY gene had certain variation,including the deletion of WRKY heptapeptide domain,the deletion of zinc finger structure,and the heptapeptide domain was mutated into WRKYGKK,WRKYGRK and WRKYGRK.Transcriptome sequencing data showed that 4 WRKY family members were not expressed in the samples,and 49 WRKY family members were up-regulated under drought stress,while 11 WRKY family members were down-regulated under drought stress.The above results showed that,chestnut WRKY genes played an important role in response to drought stress.

Key words: Chestnut, Gene family, WRKY, Drought stress

赵爽, 葛朝红, 石鹤飞, 闵卓, 王广鹏, 李伟明. 板栗WRKY基因家族鉴定及其在干旱胁迫下的表达分析[J]. 华北农学报, 2024, 39(1): 72-82. doi: 10.7668/hbnxb.20194000.

ZHAO Shuang, GE Chaohong, SHI Hefei, MIN Zhuo, WANG Guangpeng, LI Weiming. Identification and Expression Analysis of the Chestnut WRKY Gene Family under Drought Stress[J]. Acta Agriculturae Boreali-Sinica, 2024, 39(1): 72-82. doi: 10.7668/hbnxb.20194000.

http://www.hbnxb.net/CN/Y2024/V39/I1/72

图/表 9

参考文献 55

[1]	Xu Z S, Chen M, Li L C, Ma Y Z. Functions of the ERF transcription factor family in plants[J]. Botany, 2008, 86(9):969-977.doi:10.1139/B08-041. URL
[2]	Singh K B, Foley R C, Oñate-Sánchez L. Transcription factors in plant defense and stress responses[J]. Current Opinion in Plant Biology, 2002, 5(5):430-436.doi:10.1016/s1369-5266(02)00289-3. pmid: 12183182
[3]	Sun C X, Palmqvist S, Olsson H, Borén M, Ahlandsberg S, Jansson C. A novel WRKY transcription factor,SUSIBA2,participates in sugar signaling in barley by binding to the sugar-responsive elements of the iso1 promoter[J]. The Plant Cell, 2003, 15(9):2076-2092.doi:10.1105/tpc.014597. URL
[4]	Johnson C S, Kolevski B, Smyth D R. TRANSPARENT TESTA GLABRA2,a trichome and seed coat development gene of Arabidopsis,encodes a WRKY transcription factor[J]. The Plant Cell, 2002, 14(6):1359-1375.doi:10.1105/tpc.001404. URL
[5]	Zhang Z L, Xie Z, Zou X L, Casaretto J, David Ho T H, Shen Q X J. A rice WRKY gene encodes a transcriptional repressor of the gibberellin signaling pathway in aleurone cells[J]. Plant Physiology, 2004, 134(4):1500-1513.doi:10.1104/pp.103.034967. URL
[6]	Lan J, Lin Q B, Zhou C L, Ren Y K, Liu X, Miao R, Jing R N, Mou C L, Nguyen T, Zhu X J, Wang Q, Zhang X, Guo X P, Liu S J, Jiang L, Wan J M. Small grain and semi-dwarf 3,a WRKY transcription factor,negatively regulates plant height and grain size by stabilizing SLR1 expression in rice[J]. Plant Molecular Biology, 2020, 104(4/5):429-450.doi:10.1007/s11103-020-01049-0.
[7]	Wu X, Tao M Z, Meng Y, Zhu X Y, Qian L W, Shah A, Wang W, Cao S Q. The role of WRKY47 gene in regulating selenium tolerance in Arabidopsis thaliana[J]. Plant Biotechnology Reports, 2020, 14(1):121-129.doi:10.1007/s11816-019-00560-1.
[8]	Stock J, Bräutigam A, Melzer M, Bienert G P, Bunk B, Nagel M, Overmann J, Keller E R J, Mock H P. The transcription factor WRKY22 is required during cryo-stress acclimation in Arabidopsis shoot tips[J]. Journal of Experimental Botany, 2020, 71(16):4993-5009.doi:10.1093/jxb/eraa224. URL
[9]	Eulgem T, Rushton P J, Robatzek S, Somssich I E. The WRKY superfamily of plant transcription factors[J]. Trends in Plant Science, 2000, 5(5):199-206.doi:10.1016/S1360-1385(00)01600-9. pmid: 10785665
[10]	Rishmawi L, Pesch M, Juengst C, Schauss A C, Schrader A, Hülskamp M. Non-cell-autonomous regulation of root hair patterning genes by WRKY75 in Arabidopsis[J]. Plant Physiology, 2014, 165(1):186-195.doi:10.1104/pp.113.233775. pmid: 24676857
[11]	Chen H, Lai Z B, Shi J W, Xiao Y, Chen Z X, Xu X P. Roles of Arabidopsis WRKY18,WRKY40 and WRKY60 transcription factors in plant responses to abscisic acid and abiotic stress[J]. BMC Plant Biology, 2010, 10:281.doi:10.1186/1471-2229-10-281. pmid: 21167067
[12]	Park C Y, Lee J H, Yoo J H, Moon B C, Choi M S, Kang Y H, Lee S M, Kim H S, Kang K Y, Chung W S, Lim C O, Cho M J. WRKY group IId transcription factors interact with calmodulin[J]. FEBS Letters, 2005, 579(6):1545-1550.doi:10.1016/j.febslet.2005.01.057. pmid: 15733871
[13]	Liu L S, White M J, MacRae T H. Transcription factors and their genes in higher plants functional domains,evolution and regulation[J]. European Journal of Biochemistry, 1999, 262(2):247-257.doi:10.1046/j.1432-1327.1999.00349.x. pmid: 10336605
[14]	Chen L G, Song Y, Li S J, Zhang L P, Zou C S, Yu D Q. The role of WRKY transcription factors in plant abiotic stresses[J]. Biochimica et Biophysica Acta, 2012, 1819(2):120-128.doi:10.1016/j.bbagrm.2011.09.002. pmid: 21964328
[15]	Ishiguro S, Nakamura K. Characterization of a cDNA encoding a novel DNA-binding protein,SPF1,that recognizes SP8 sequences in the 5' upstream regions of genes coding for sporamin and beta-amylase from sweet potato[J]. Molecular and General Genetics, 1994, 244(6):563-571.doi:10.1007/bf00282746. pmid: 7969025
[16]	Ülker B, Somssich I E. WRKY transcription factors:from DNA binding towards biological function[J]. Current Opinion in Plant Biology, 2004, 7(5):491-498.doi:10.1016/j.pbi.2004.07.012. URL
[17]	Ross C A, Liu Y, Shen Q X J. The WRKY gene family in rice(Oryza sativa)[J]. Journal of Integrative Plant Biology, 2007, 49(6):827-842.doi:10.1111/j.1744-7909.2007.00504.x. URL
[18]	Bencke-Malato M, Cabreira C, Wiebke-Strohm B, Bücker-Neto L, Mancini E, Osorio M B, Homrich M S, Turchetto-Zolet A C, De Carvalho M C, Stolf R, Weber R L, Westergaard G, Castagnaro A P, Abdelnoor R V, Marcelino-Guimarães F C, Margis-Pinheiro M, Bodanese-Zanettini M H. Genome-wide annotation of the soybean WRKY family and functional characterization of genes involved in response to Phakopsora pachyrhizi infection[J]. BMC Plant Biology, 2014, 14:236.doi:10.1186/s12870-014-0236-0. pmid: 25201117
[19]	He H S, Dong Q, Shao Y H, Jiang H Y, Zhu S W, Cheng B J, Xiang Y. Genome-wide survey and characterization of the WRKY gene family in Populus trichocarpa[J]. Plant Cell Reports, 2012, 31(7):1199-1217.doi:10.1007/s00299-012-1241-0. URL
[20]	Tripathi P, Rabara R C, Langum T J, Boken A K, Rushton D L, Boomsma D D, Rinerson C I, Rabara J, Reese R N, Chen X F, Rohila J S, Rushton P J. The WRKY transcription factor family in Brachypodium distachyon[J]. BMC Genomics, 2012, 13:270.doi:10.1186/1471-2164-13-270. pmid: 22726208
[21]	Ling J, Jiang W J, Zhang Y, Yu H J, Mao Z C, Gu X F, Huang S W, Xie B Y. Genome-wide analysis of WRKY gene family in Cucumis sativus[J]. BMC Genomics, 2011, 12:471.doi:10.1186/1471-2164-12-471. pmid: 21955985
[22]	Wei K F, Chen J, Chen Y F, Wu L J, Xie D X. Molecular phylogenetic and expression analysis of the complete WRKY transcription factor family in maize[J]. DNA Research, 2012, 19(2):153-164.doi:10.1093/dnares/dsr048. URL
[23]	Liu Z, Saiyinduleng, Chang Q Y, Cheng C W, Zheng Z M, Yu S. Identification of yellowhorn(Xanthoceras sorbifolium)WRKY transcription factor family and analysis of abiotic stress response model[J]. Journal of Forestry Research, 2021, 32(3):987-1004.doi:10.1007/s11676-020-01134-6.
[24]	Dong X S, Yang Y N, Zhang Z Y, Xiao Z W, Bai X H, Gao J, Hur Y K, Dong R S, Yang A, Hao S M, He F F. Genome-wide identification of WRKY genes and their response to cold stress in Coffea canephora[J]. Forests, 2019, 10(4):335.doi:10.3390/f10040335. URL
[25]	Li W, Wang H P, Yu D Q. Arabidopsis WRKY transcription factors WRKY12 and WRKY13 oppositely regulate flowering under short-day conditions[J]. Molecular Plant, 2016, 9(11):1492-1503.doi:10.1016/j.molp.2016.08.003. URL
[26]	Robatzek S, Somssich I E. Targets of AtWRKY6 regulation during plant senescence and pathogen defense[J]. Genes & Development, 2002, 16(9):1139-1149.doi:10.1101/gad.222702. URL
[27]	Yu F F, Huaxia Y F, Lu W J, Wu C G, Cao X C, Guo X Q. GhWRKY15,a member of the WRKY transcription factor family identified from cotton(Gossypium hirsutum L.),is involved in disease resistance and plant development[J]. BMC Plant Biology, 2012, 12:144.doi:10.1186/1471-2229-12-144.
[28]	Wei Y X, Liu W, Hu W, Yan Y, Shi H T. The chaperone MeHSP90 recruits MeWRKY20 and MeCatalase1 to regulate drought stress resistance in cassava[J]. New Phytologist, 2020, 226(2):476-491.doi:10.1111/nph.16346. pmid: 31782811
[29]	Zhang Y, Yu H J, Yang X Y, Li Q, Ling J, Wang H, Gu X F, Huang S W, Jiang W J. CsWRKY46,a WRKY transcription factor from cucumber,confers cold resistance in transgenic-plant by regulating a set of cold-stress responsive genes in an ABA-dependent manner[J]. Plant Physiology Biochemistry, 2016, 108:478-487.doi:10.1016/j.plaphy.2016.08.013. URL
[30]	Li R, Zhang J, Li J C, Zhou G X, Wang Q, Bian W B, Erb M, Lou Y G. Prioritizing plant defence over growth through WRKY regulation facilitates infestation by non-target herbivores[J]. ELife, 2015, 4:e04805.doi:10.7554/elife.04805. URL
[31]	Liu X Q, Bai X Q, Wang X J, Chu C C. OsWRKY71,a rice transcription factor,is involved in rice defense response[J]. Journal of Plant Physiology, 2007, 164(8):969-979.doi:10.1016/j.jplph.2006.07.006. URL
[32]	Tian S L, Shen G N, Xu L, Sun X L. Effects of different water-saving irrigation modes on chestnut growth and fruiting in drought hilly land[J]. Chinese Journal of Applied Ecology, 2012, 23(3):639-644.doi:10.2307/2404042.
[33]	郭燕, 张树航, 李颖, 张馨方, 王广鹏. 中国板栗238份品种(系)叶片形态、解剖结构及其抗旱性评价[J]. 园艺学报, 2020, 47(6):1033-1046.doi:10.16420/j.issn.0513-353x.2019-0771.
	Guo Y, Zhang S H, Li Y, Zhang X F, Wang G P. Studies on the leaf morphology,anatomical structure and drought resistance evaluation of 238 Chinese chestnut varieties(strains)[J]. Acta Horticulturae Sinica, 2020, 47(6):1033-1046.
[34]	Luo X, Bai X, Sun X L, Zhu D, Liu B H, Ji W, Cai H, Cao L, Wu J, Hu M R, Liu X, Tang L L, Zhu Y M. Expression of wild soybean WRKY20 in Arabidopsis enhances drought tolerance and regulates ABA signalling[J]. Journal of Experimental Botany, 2013, 64(8):2155-2169.doi:10.1093/jxb/ert073. URL
[35]	Yang Z, Chi X Y, Guo F F, Jin X Y, Luo H L, Hawar A, Chen Y X, Feng K K, Wang B, Qi J L, Yang Y H, Sun B. SbWRKY30 enhances the drought tolerance of plants and regulates a drought stress-responsive gene,SbRD19,in sorghum[J]. Journal of Plant Physiology, 2020, 246/247:153142.doi:10.1016/j.jplph.2020.153142. URL
[36]	Shahzad R, Ewas M, Harlina P W, Nishawy E, Ayaad M, Manan A, Maher M, Khames E. Multiple stress responsive WRKY transcription factor,StWRKY2,enhances drought and late blight resistance in transgenic potato[J]. International Journal of Agriculture and Biology, 2020, 24(1):154-164.doi:10.17957/IJAB/15.1419.
[37]	Xing Y, Liu Y, Zhang Q, Nie X H, Sun Y M, Zhang Z Y, Li H C, Fang K F, Wang G P, Huang H W, Bisseling T, Cao Q Q, Qin L. Hybrid de novo genome assembly of Chinese chestnut(Castanea mollissima)[J]. GigaScience, 2019, 8(9):giz112.doi:10.1093/gigascience/giz112. URL
[38]	Camacho C, Coulouris G, Avagyan V, Ma N, Papadopoulos J, Bealer K, Madden T L. BLAST+:architecture and applications[J]. BMC Bioinformatics, 2009, 10:421.doi:10.1186/1471-2105-10-421.
[39]	Hu B, Jin J P, Guo A Y, Zhang H, Luo J C, Gao G. GSDS 2.0:an upgraded gene feature visualization server[J]. Bioinformatics, 2015, 31(8):1296-1297.doi:10.1093/bioinformatics/btu817. URL
[40]	Bailey T L, Boden M, Buske F A, Frith M, Grant C E, Clementi L, Ren J Y, Li W W, Noble W S. MEME SUITE:tools for motif discovery and searching[J]. Nucleic Acids Research, 2009, 37(Web Server Issue):W202-W208.doi:10.1093/nar/gkp335. URL
[41]	Livak K J, Schmittgen T D. Analysis of relative gene expression data using real-time quantitative PCR and the 2^-ΔΔCt method[J]. Methods, 2001, 25(4):402-408.doi:10.1006/meth.2001.1262. pmid: 11846609
[42]	Qiu Y P, Jing S J, Fu J, Li L, Yu D Q. Cloning and analysis of expression profile of 13 WRKY genes in rice[J]. Chinese Science Bulletin, 2004, 49(20):2159-2168.doi:10.1007/BF03185783.
[43]	Seungmin S, Ho-Keun A, Young-Joo S, Sang R P, Jong H I. Rice transcription factor WRKY114 directly regulates the expression of OsPR1a and Chitinase to enhance resistance against Xanthomonas oryzae pv.oryzae[J]. Biochemical and Biophysical Research Communications, 2020, 533(4):1262-1268.doi:10.1016/j.bbrc.2020.09.141. pmid: 33051057
[44]	Wei W, Hu Y, Han Y T, Zhang K, Zhao F L, Feng J Y. The WRKY transcription factors in the diploid woodland strawberry Fragaria vesca:identification and expression analysis under biotic and abiotic stresses[J]. Plant Physiology & Biochemistry, 2016, 105:129-144.doi:10.1016/j.plaphy.2016.04.014.
[45]	Chen F, Hu Y, Vannozzi A, Wu K C, Cai H Y, Qin Y, Mullis A, Lin Z G, Zhang L S. The WRKY transcription factor family in model plants and crops[J]. Critical Reviews in Plant Sciences, 2018, 36(5/6):311-335.doi:10.1080/07352689.2018.1441103. URL
[46]	Qiu Y P, Yu D Q. Over-expression of the stress-induced OsWRKY45 enhances disease resistance and drought tolerance in Arabidopsis[J]. Environmental and Experimental Botany, 2009, 65(1):35-47.doi:10.1016/j.envexpbot.2008.07.002. URL
[47]	Jin J P, Tian F, Yang D C, Meng Y Q, Kong L, Luo J C, Gao G. PlantTFDB 4.0:toward a central hub for transcription factors and regulatory interactions in plants[J]. Nucleic Acids Research, 2017, 45(D1):D1040-D1045.doi:10.1093/nar/gkw982. URL
[48]	Zhao N N, He M J, Li L, Cui S L, Hou M Y, Wang L, Mu G J, Liu L F, Yang X L. Identification and expression analysis of WRKY gene family under drought stress in peanut(Arachis hypogaea L.)[J]. PLoS One, 2020, 15(4):e0231396.doi:10.1371/journal.pone.0231396. URL
[49]	Kobe B, Kajava A V. The leucine-rich repeat as a protein recognition motif[J]. Current Opinion in Structural Biology, 2001, 11(6):725-732.doi:10.1016/s0959-440x(01)00266-4. pmid: 11751054
[50]	Matsushima N, Enkhbayar P, Kamiya M, Osaki M, Kretsinger R. Leucine-Rich Repeats(LRRs):structure,function,evolution and interaction with ligands[J]. Drug Design Reviews, 2005, 2(4):305-322.doi:10.2174/1567269054087613. URL
[51]	Bella J, Hindle K L, McEwan P A, Lovell S C. The leucine-rich repeat structure[J]. Cellular and Molecular Life Sciences, 2008, 65(15):2307-2333.doi:10.1007/s00018-008-8019-0. pmid: 18408889
[52]	Jiang J J, Ma S H, Ye N H, Jiang M, Cao J S, Zhang J H. WRKY transcription factors in plant responses to stresses[J]. Journal of Integrative Plant Biology, 2017, 59(2):86-101.doi:10.1111/jipb.12513.
[53]	Zhao K X, Chu S S, Zhang X D, Wang L P, Rono J K, Yang Z M. AtWRKY21 negatively regulates tolerance to osmotic stress in Arabidopsis[J]. Environmental and Experimental Botany, 2020, 169:103920.doi:10.1016/j.envexpbot.2019.103920. URL
[54]	El-Esawi M A, Al-Ghamdi A A, Ali H M, Ahmad M. Overexpression of AtWRKY30 transcription factor enhances heat and drought stress tolerance in wheat(Triticum aestivum L.)[J]. Genes, 2019, 10(2):163.doi:10.3390/genes10020163. URL
[55]	Chen J N, Nolan T M, Ye H X, Zhang M C, Tong H N, Xin P Y, Chu J F, Chu C C, Li Z H, Yin Y H. Arabidopsis WRKY46,WRKY54,and WRKY70 transcription factors are involved in brassinosteroid-regulated plant growth and drought responses[J]. The Plant Cell, 2017, 29(6):1425-1439.doi:10.1105/tpc.17.00364.

基因名称 Gene name	上游引物(5'—3') Forward primer (5'—3')	下游引物(5'—3') Reverse primer(5'—3')
CmWRKY1	ACAGGGTTGGAAGGACCTCA	CTTGTTTTGTTGCCCAGCAGT
CmWRKY5	GGTCCCTGCGATTAGCTTGA	TTTACGTGCTGGACACCCTC
CmWRKY6	AGAAGAGGTGCCACGAACAC	TAACCCCGTGGGTATGGTGA
CmWRKY10	GAGACTGGTGCTTTGGTGGA	ATCCATCAACTGGCTTCGCA
CmWRKY11	TGGCAGTTGGTTGTCCTGTT	CTGCTGTTGTGGTTGATGCC
CmWRKY19	AGTCGCCTTCCATTTCAGCA	AACTGTTGGAAAGCCGGTGA
CmWRKY21	CTCTGACCTCCTTGCCTCAC	AATTTAGGCACACCCGACCC
CmWRKY23	ACGAATGGCGGAAATATGGC	CTGGTAAGGCATGGTTGTGC
CmWRKY26	GGACAACAGTCTCTCGCACA	TGCACTTGTTTGGTTGCTCG
CmWRKY27	TGCTTGTGTAGGAGAGCCCA	CTGTCCATCGGGGCAATGT
CmWRKY28	TTTTCGTTGGAGAAGGCGGT	AAGGCAGCATCACCACTCTG
CmWRKY41	AGAGTCCCTGCAATTAGCCC	CTTGGATGCGGGGAACCTTT
内参基因 Reference gene	ATTCACGAGACCACCTACA	TGCCACAACCTTAATCTTCAT

基因名称 Gene name	上游引物(5'—3') Forward primer (5'—3')	下游引物(5'—3') Reverse primer(5'—3')
CmWRKY1	ACAGGGTTGGAAGGACCTCA	CTTGTTTTGTTGCCCAGCAGT
CmWRKY5	GGTCCCTGCGATTAGCTTGA	TTTACGTGCTGGACACCCTC
CmWRKY6	AGAAGAGGTGCCACGAACAC	TAACCCCGTGGGTATGGTGA
CmWRKY10	GAGACTGGTGCTTTGGTGGA	ATCCATCAACTGGCTTCGCA
CmWRKY11	TGGCAGTTGGTTGTCCTGTT	CTGCTGTTGTGGTTGATGCC
CmWRKY19	AGTCGCCTTCCATTTCAGCA	AACTGTTGGAAAGCCGGTGA
CmWRKY21	CTCTGACCTCCTTGCCTCAC	AATTTAGGCACACCCGACCC
CmWRKY23	ACGAATGGCGGAAATATGGC	CTGGTAAGGCATGGTTGTGC
CmWRKY26	GGACAACAGTCTCTCGCACA	TGCACTTGTTTGGTTGCTCG
CmWRKY27	TGCTTGTGTAGGAGAGCCCA	CTGTCCATCGGGGCAATGT
CmWRKY28	TTTTCGTTGGAGAAGGCGGT	AAGGCAGCATCACCACTCTG
CmWRKY41	AGAGTCCCTGCAATTAGCCC	CTTGGATGCGGGGAACCTTT
内参基因 Reference gene	ATTCACGAGACCACCTACA	TGCCACAACCTTAATCTTCAT

基因名称 Gene name	基因号 Gene ID	序列 Scaffold			氨基酸长度/aa Amino acid length	分子质量/ku MW	等电点 pI	CDS长度/bp CDS length	七肽域 Heptapeptide	锌指结构 Zinc finger	组别 Group
		序列号 No.	位置/bp Position
		序列号 No.	From	To
CmWRKY1	CMHBY200296	ctg0002	3 819 280	3 820 865	358	40.032 24	5.42	1 074	WRKYGQK	C2HC	Ⅲ
CmWRKY2	CMHBY200841	ctg0006	3 189 450	3 192 831	178	20.104 56	9.45	534	WRKYGQK	C2H2	Ⅱc
CmWRKY3	CMHBY201642	ctg0015	551 307	552 000	187	21.099 93	9.74	561	WRKYGQK	C2H2	Ⅱc
CmWRKY4	CMHBY201644	ctg0015	563 993	565 021	297	33.317 43	5.69	891	WRKYGQK	C2H2	Ⅱe
CmWRKY5	CMHBY201907	ctg0017	3 084 906	3 086 292	337	37.165 88	9.61	1 011	WRKYGQK	C2H2	Ⅱd
CmWRKY6	CMHBY201976	ctg0018	2 011 234	2 013 043	319	34.602 16	9.48	957	WRKYGQK	C2H2	Ⅱd
CmWRKY7	CMHBY202049	ctg0019	672 049	678 460	511	56.155 34	8.12	1 533	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY8	CMHBY202109	ctg0019	2 421 959	2 425 973	560	60.871 58	6.02	1 680	None	C2H2	Ⅱb
CmWRKY9	CMHBY202144	ctg0020	466 917	469 565	562	61.015 00	8.27	1 686	WRKYGQK	C2H2	Ⅱb
CmWRKY10	CMHBY202286	ctg0022	658 043	660 044	311	34.494 63	8.63	933	WRKYGQK	C2H2	Ⅱa
CmWRKY11	CMHBY202726	ctg0028	1 607 424	1 609 926	631	68.497 65	6.41	1 893	WRKYGQK	C2H2	Ⅱb
CmWRKY12	CMHBY202917	ctg0031	1 368 429	1 370 657	415	45.247 92	6.25	1 245	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY13	CMHBY203010	ctg0033	78 218	80 958	607	65.585 85	5.93	1 821	WRKYGQK	C2H2	Ⅱb
CmWRKY14	CMHBY203172	ctg0036	1 788 738	1 791 911	125	14.509 46	9.84	375	WRKYGQK	C2H2	Ⅱc
CmWRKY15	CMHBY203286	ctg0038	575 897	577 764	323	36.224 96	5.90	969	WRKYGQK	C2HC	Ⅲ
CmWRKY16	CMHBY203524	ctg0043	723 095	724 650	348	38.529 42	6.18	1 044	WRKYGQK	C2H2	Ⅱc
CmWRKY17	CMHBY204080	ctg0054	757 019	759 176	517	57.331 39	5.19	1 551	WRKYGQK	C2H2	Ⅱb
CmWRKY18	CMHBY204310	ctg0059	705 525	716 193	532	58.049 02	5.02	1 596	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY19	CMHBY205107	ctg0077	1 155 849	1 158 176	317	35.610 91	7.65	951	WRKYGQK	C2H2	Ⅱa
CmWRKY20	CMHBY205108	ctg0077	1 195 492	1 196 805	266	29.940 01	8.95	798	WRKYGQK	C2H2	Ⅱa
CmWRKY21	CMHBY205461	ctg0084	100 470	103 546	601	66.277 11	6.71	1 803	WRKYGQK	C2H2/C2H2	Ⅰ
									/WRKYGQK
CmWRKY22	CMHBY206287	ctg0106	976 128	982 839	716	77.723 59	6.13	2 148	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY23	CMHBY206346	ctg0108	754 075	754 711	104	11.708 08	9.61	312	WRKYGQK	C2H2	Ⅰ
CmWRKY24	CMHBY206743	ctg0119	823 860	829 392	610	67.028 55	6.85	1 830	WRKYGQK	C2H2	Ⅱb
CmWRKY25	CMHBY207206	ctg0132	474 665	478 302	370	40.334 03	6.01	1 110	WRKYGQK	C2HC	Ⅲ
CmWRKY26	CMHBY207207	ctg0132	480 307	481 816	328	37.069 01	5.54	984	WRKYGQK	C2HC	Ⅲ
CmWRKY27	CMHBY207827	ctg0156	185 305	187 918	345	38.637 93	5.28	1 035	WRKYGQK	C2HC	Ⅲ
CmWRKY28	CMHBY208621	ctg0184	331 282	334 536	587	65.384 84	8.05	1 761	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY29	CMHBY208659	ctg0185	160 154	163 898	479	52.124 97	9.15	1 437	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY30	CMHBY208739	ctg0187	40 062	41 344	308	34.164 37	6.67	924	WRKYGQK	C2H2	Ⅱc
CmWRKY31	CMHBY208751	ctg0187	514 452	515 760	262	29.673 23	5.21	786	WRKYGQK	C2H2	Ⅱe
CmWRKY32	CMHBY209160	ctg0201	231 216	233 088	153	17.033 46	9.10	459	WRKYGQK	C2H2	Ⅱd
CmWRKY33	CMHBY209246	ctg0205	155 844	161 515	602	65.198 13	7.10	1 806	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY34	CMHBY209375	ctg0209	601 155	60 3187	159	18.470 75	9.54	477	WRKYGQK	C2H2	Ⅱc
CmWRKY35	CMHBY209582	ctg0217	652 446	656 055	365	40.350 72	5.89	1 095	WRKYGQK	C2HC	Ⅲ
CmWRKY36	CMHBY209993	ctg0235	303 630	305 981	385	42.936 53	9.32	1 155	WRKYGQK	C2H2	Ⅱd
CmWRKY37	CMHBY210429	ctg0253	734 526	737 310	284	32.208 86	5.17	852	WRKYGQK	C2H2	Ⅰ
CmWRKY38	CMHBY210481	ctg0256	435 954	437 364	366	40.592 41	5.29	1 098	WRKYGQK	C2H2	Ⅱe
CmWRKY39	CMHBY210484	ctg0256	522 779	526 722	185	20.884 72	9.56	555	WRKYGQK	C2H2	Ⅱc
CmWRKY40	CMHBY210634	ctg0263	447 165	448 857	399	44.613 04	7.19	1 197	WRIYGRK	C2H2	Ⅰ
CmWRKY41	CMHBY211173	ctg0288	432 976	434 772	342	36.932 20	9.51	1 026	WRKYGQK	C2H2	Ⅱd
CmWRKY42	CMHBY211728	ctg0319	255 679	256 903	166	19.130 71	5.39	498	WRKYGKK	C2H2	Ⅱc
CmWRKY43	CMHBY211892	ctg0329	172 606	178 898	233	26.306 61	9.00	699	WRKYGQK	C2H2	Ⅱc
CmWRKY44	CMHBY211905	ctg0329	467 919	471 452	192	21.648 77	6.75	576	WRKYGKK	C2H2	Ⅱc
CmWRKY45	CMHBY212107	ctg0344	248 488	253 413	837	89.945 94	6.14	2 511	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY46	CMHBY212251	ctg0350	663 296	665 093	342	36.946 23	9.57	1 026	WRKYGQK	C2H2	Ⅱd
CmWRKY47	CMHBY212900	ctg0389	183 380	189 150	498	54.510 02	6.09	1 494	WRKYGQK	C2H2	Ⅱb
CmWRKY48	CMHBY212955	ctg0392	7 310	8 467	265	29.731 81	5.37	795	WRKYGQK	C2H2	Ⅱe
CmWRKY49	CMHBY213251	ctg0409	376 075	378 121	335	37.527 69	9.69	1 005	WRKYGQK	C2H2	Ⅱd
CmWRKY50	CMHBY213285	ctg0411	50 771	56 127	582	61.677 63	6.45	1 746	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY51	CMHBY213532	ctg0428	216 333	219 113	463	49.866 88	5.25	1 389	WRKYGQK	C2H2	Ⅱe
CmWRKY52	CMHBY213833	ctg0451	36 690	38 098	384	41.998 13	6.08	1 152	WRKYGQK	C2H2	Ⅱe
CmWRKY53	CMHBY214338	ctg0484	478 121	481 223	309	34.016 11	5.66	927	WRKYGQK	C2H2	Ⅱc
CmWRKY54	CMHBY214647	ctg0511	328 965	330 843	322	36.354 23	8.07	966	WRKYGQK	C2H2	Ⅱc
CmWRKY55	CMHBY214751	ctg0521	162 308	166 769	524	56.661 59	8.06	1 572	WRKYGQK	C2H2	Ⅱb
CmWRKY56	CMHBY214924	ctg0536	344 729	350 993	289	31.193 52	5.47	867	WRKYGQK	C2H2	Ⅱe
CmWRKY57	CMHBY215009	ctg0544	410 481	411 631	265	29.759 86	5.48	795	WRKYGQK	C2H2	Ⅱe
CmWRKY58	CMHBY215307	ctg0577	311 202	317 323	228	25.567 96	7.04	684	WRKYGQK	C2H2	Ⅱc
CmWRKY59	CMHBY216199	ctg0692	134 992	136 603	310	34.240 94	6.10	930	WRKYGQK	C2H2	Ⅱc
CmWRKY60	CMHBY216832	ctg0822	18 701	19 650	112	12.189 29	10.05	336	WRKYGQK	C2QXH	Ⅱb
CmWRKY61	CMHBY217264	ctg0960	105 665	110 232	581	63.010 46	6.66	1 743	WRKYGQK	C2H2	Ⅱb
CmWRKY62	CMHBY226653	ctg0263	435 559	446 506	1 243	139.990 50	5.99	3 729	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY63	CMHBY226659	ctg0263	500 247	500 879	211	23.504 68	9.54	633	WRKYGRK	None	Ⅰ
CmWRKY64	CMHBY228900	ctg0411	9 315	9 470	52	5.833 47	8.09	156	WRKYGQK	None	Ⅰ
CmWRKY65	CMHBY228901	ctg0411	10 343	10 560	73	7.935 91	9.47	218	None	C2H2	Ⅱd

基因名称 Gene name	基因号 Gene ID	序列 Scaffold			氨基酸长度/aa Amino acid length	分子质量/ku MW	等电点 pI	CDS长度/bp CDS length	七肽域 Heptapeptide	锌指结构 Zinc finger	组别 Group
		序列号 No.	位置/bp Position
		序列号 No.	From	To
CmWRKY1	CMHBY200296	ctg0002	3 819 280	3 820 865	358	40.032 24	5.42	1 074	WRKYGQK	C2HC	Ⅲ
CmWRKY2	CMHBY200841	ctg0006	3 189 450	3 192 831	178	20.104 56	9.45	534	WRKYGQK	C2H2	Ⅱc
CmWRKY3	CMHBY201642	ctg0015	551 307	552 000	187	21.099 93	9.74	561	WRKYGQK	C2H2	Ⅱc
CmWRKY4	CMHBY201644	ctg0015	563 993	565 021	297	33.317 43	5.69	891	WRKYGQK	C2H2	Ⅱe
CmWRKY5	CMHBY201907	ctg0017	3 084 906	3 086 292	337	37.165 88	9.61	1 011	WRKYGQK	C2H2	Ⅱd
CmWRKY6	CMHBY201976	ctg0018	2 011 234	2 013 043	319	34.602 16	9.48	957	WRKYGQK	C2H2	Ⅱd
CmWRKY7	CMHBY202049	ctg0019	672 049	678 460	511	56.155 34	8.12	1 533	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY8	CMHBY202109	ctg0019	2 421 959	2 425 973	560	60.871 58	6.02	1 680	None	C2H2	Ⅱb
CmWRKY9	CMHBY202144	ctg0020	466 917	469 565	562	61.015 00	8.27	1 686	WRKYGQK	C2H2	Ⅱb
CmWRKY10	CMHBY202286	ctg0022	658 043	660 044	311	34.494 63	8.63	933	WRKYGQK	C2H2	Ⅱa
CmWRKY11	CMHBY202726	ctg0028	1 607 424	1 609 926	631	68.497 65	6.41	1 893	WRKYGQK	C2H2	Ⅱb
CmWRKY12	CMHBY202917	ctg0031	1 368 429	1 370 657	415	45.247 92	6.25	1 245	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY13	CMHBY203010	ctg0033	78 218	80 958	607	65.585 85	5.93	1 821	WRKYGQK	C2H2	Ⅱb
CmWRKY14	CMHBY203172	ctg0036	1 788 738	1 791 911	125	14.509 46	9.84	375	WRKYGQK	C2H2	Ⅱc
CmWRKY15	CMHBY203286	ctg0038	575 897	577 764	323	36.224 96	5.90	969	WRKYGQK	C2HC	Ⅲ
CmWRKY16	CMHBY203524	ctg0043	723 095	724 650	348	38.529 42	6.18	1 044	WRKYGQK	C2H2	Ⅱc
CmWRKY17	CMHBY204080	ctg0054	757 019	759 176	517	57.331 39	5.19	1 551	WRKYGQK	C2H2	Ⅱb
CmWRKY18	CMHBY204310	ctg0059	705 525	716 193	532	58.049 02	5.02	1 596	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY19	CMHBY205107	ctg0077	1 155 849	1 158 176	317	35.610 91	7.65	951	WRKYGQK	C2H2	Ⅱa
CmWRKY20	CMHBY205108	ctg0077	1 195 492	1 196 805	266	29.940 01	8.95	798	WRKYGQK	C2H2	Ⅱa
CmWRKY21	CMHBY205461	ctg0084	100 470	103 546	601	66.277 11	6.71	1 803	WRKYGQK	C2H2/C2H2	Ⅰ
									/WRKYGQK
CmWRKY22	CMHBY206287	ctg0106	976 128	982 839	716	77.723 59	6.13	2 148	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY23	CMHBY206346	ctg0108	754 075	754 711	104	11.708 08	9.61	312	WRKYGQK	C2H2	Ⅰ
CmWRKY24	CMHBY206743	ctg0119	823 860	829 392	610	67.028 55	6.85	1 830	WRKYGQK	C2H2	Ⅱb
CmWRKY25	CMHBY207206	ctg0132	474 665	478 302	370	40.334 03	6.01	1 110	WRKYGQK	C2HC	Ⅲ
CmWRKY26	CMHBY207207	ctg0132	480 307	481 816	328	37.069 01	5.54	984	WRKYGQK	C2HC	Ⅲ
CmWRKY27	CMHBY207827	ctg0156	185 305	187 918	345	38.637 93	5.28	1 035	WRKYGQK	C2HC	Ⅲ
CmWRKY28	CMHBY208621	ctg0184	331 282	334 536	587	65.384 84	8.05	1 761	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY29	CMHBY208659	ctg0185	160 154	163 898	479	52.124 97	9.15	1 437	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY30	CMHBY208739	ctg0187	40 062	41 344	308	34.164 37	6.67	924	WRKYGQK	C2H2	Ⅱc
CmWRKY31	CMHBY208751	ctg0187	514 452	515 760	262	29.673 23	5.21	786	WRKYGQK	C2H2	Ⅱe
CmWRKY32	CMHBY209160	ctg0201	231 216	233 088	153	17.033 46	9.10	459	WRKYGQK	C2H2	Ⅱd
CmWRKY33	CMHBY209246	ctg0205	155 844	161 515	602	65.198 13	7.10	1 806	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY34	CMHBY209375	ctg0209	601 155	60 3187	159	18.470 75	9.54	477	WRKYGQK	C2H2	Ⅱc
CmWRKY35	CMHBY209582	ctg0217	652 446	656 055	365	40.350 72	5.89	1 095	WRKYGQK	C2HC	Ⅲ
CmWRKY36	CMHBY209993	ctg0235	303 630	305 981	385	42.936 53	9.32	1 155	WRKYGQK	C2H2	Ⅱd
CmWRKY37	CMHBY210429	ctg0253	734 526	737 310	284	32.208 86	5.17	852	WRKYGQK	C2H2	Ⅰ
CmWRKY38	CMHBY210481	ctg0256	435 954	437 364	366	40.592 41	5.29	1 098	WRKYGQK	C2H2	Ⅱe
CmWRKY39	CMHBY210484	ctg0256	522 779	526 722	185	20.884 72	9.56	555	WRKYGQK	C2H2	Ⅱc
CmWRKY40	CMHBY210634	ctg0263	447 165	448 857	399	44.613 04	7.19	1 197	WRIYGRK	C2H2	Ⅰ
CmWRKY41	CMHBY211173	ctg0288	432 976	434 772	342	36.932 20	9.51	1 026	WRKYGQK	C2H2	Ⅱd
CmWRKY42	CMHBY211728	ctg0319	255 679	256 903	166	19.130 71	5.39	498	WRKYGKK	C2H2	Ⅱc
CmWRKY43	CMHBY211892	ctg0329	172 606	178 898	233	26.306 61	9.00	699	WRKYGQK	C2H2	Ⅱc
CmWRKY44	CMHBY211905	ctg0329	467 919	471 452	192	21.648 77	6.75	576	WRKYGKK	C2H2	Ⅱc
CmWRKY45	CMHBY212107	ctg0344	248 488	253 413	837	89.945 94	6.14	2 511	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY46	CMHBY212251	ctg0350	663 296	665 093	342	36.946 23	9.57	1 026	WRKYGQK	C2H2	Ⅱd
CmWRKY47	CMHBY212900	ctg0389	183 380	189 150	498	54.510 02	6.09	1 494	WRKYGQK	C2H2	Ⅱb
CmWRKY48	CMHBY212955	ctg0392	7 310	8 467	265	29.731 81	5.37	795	WRKYGQK	C2H2	Ⅱe
CmWRKY49	CMHBY213251	ctg0409	376 075	378 121	335	37.527 69	9.69	1 005	WRKYGQK	C2H2	Ⅱd
CmWRKY50	CMHBY213285	ctg0411	50 771	56 127	582	61.677 63	6.45	1 746	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY51	CMHBY213532	ctg0428	216 333	219 113	463	49.866 88	5.25	1 389	WRKYGQK	C2H2	Ⅱe
CmWRKY52	CMHBY213833	ctg0451	36 690	38 098	384	41.998 13	6.08	1 152	WRKYGQK	C2H2	Ⅱe
CmWRKY53	CMHBY214338	ctg0484	478 121	481 223	309	34.016 11	5.66	927	WRKYGQK	C2H2	Ⅱc
CmWRKY54	CMHBY214647	ctg0511	328 965	330 843	322	36.354 23	8.07	966	WRKYGQK	C2H2	Ⅱc
CmWRKY55	CMHBY214751	ctg0521	162 308	166 769	524	56.661 59	8.06	1 572	WRKYGQK	C2H2	Ⅱb
CmWRKY56	CMHBY214924	ctg0536	344 729	350 993	289	31.193 52	5.47	867	WRKYGQK	C2H2	Ⅱe
CmWRKY57	CMHBY215009	ctg0544	410 481	411 631	265	29.759 86	5.48	795	WRKYGQK	C2H2	Ⅱe
CmWRKY58	CMHBY215307	ctg0577	311 202	317 323	228	25.567 96	7.04	684	WRKYGQK	C2H2	Ⅱc
CmWRKY59	CMHBY216199	ctg0692	134 992	136 603	310	34.240 94	6.10	930	WRKYGQK	C2H2	Ⅱc
CmWRKY60	CMHBY216832	ctg0822	18 701	19 650	112	12.189 29	10.05	336	WRKYGQK	C2QXH	Ⅱb
CmWRKY61	CMHBY217264	ctg0960	105 665	110 232	581	63.010 46	6.66	1 743	WRKYGQK	C2H2	Ⅱb
CmWRKY62	CMHBY226653	ctg0263	435 559	446 506	1 243	139.990 50	5.99	3 729	WRKYGQK	C2H2	Ⅰ
									/WRKYGQK	/C2H2
CmWRKY63	CMHBY226659	ctg0263	500 247	500 879	211	23.504 68	9.54	633	WRKYGRK	None	Ⅰ
CmWRKY64	CMHBY228900	ctg0411	9 315	9 470	52	5.833 47	8.09	156	WRKYGQK	None	Ⅰ
CmWRKY65	CMHBY228901	ctg0411	10 343	10 560	73	7.935 91	9.47	218	None	C2H2	Ⅱd

基序 Motif	宽度 Width	E值 E value	位点 Sites	最佳匹配 Best possible match
1	29	4.5e-1397	63	I[LP]DDGYRWRKYGQK[PV][VI]K[GN][SN]P[YH]PR[SGA]YY[RK]C[TS]
2	29	2.7e-1060	60	GCP[VA][RK]K[QH]V[EQ]R[SAC]S[ED]DP[ST][IM][LV][IV][TV]TYEGEHNHP
3	50	2.3e-355	11	DGYNWRKYGQKQVKGSE[FY]PRSYYKCT[HY][PS]NCPVKKKVERS[HL]DGQITEI[IV]YK
4	50	7.80e-111	11	XKRXKIELXX[LA][KR]AE[LM]G[RE][MV][KS]EEN[EQ][RK]L[KR]E[ML]LDQ[MV]TKNY[NQ][AS]LQ[MS][HQ][LF]X[DT][IL]MQQ[EQ]
5	15	2.20e-139	32	[VI]R[EK][PA]R[VF][ASV][VF][QR][TA]RS[ED]VD
6	29	4.20e-88	10	L[PS][PV][AS]A[TM]AMASTTS[AS]A[AV][SR]MLLSGS[MST][ST]S[ASK][DQ][GA]
7	33	4.60e-85	16	L[VI]SAATAAI[AT]SDPNFT[AS]ALAAAISSI[IM][GL][AK]PXX[NP]
8	21	3.20e-52	17	CHCSK[KR][RK]KN[RL]QKR[VT][VI]RVP[AI]IS
9	15	3.80e-27	10	S[AST]S[ASP][PS]FPT[IV]TLDLT[QHT]
10	18	4.80e-26	16	[GH][ST]HNHPKPQPN[RK]RAXSGS

选择文件类型/文献管理软件名称

选择包含的内容