绣球WRKY基因家族全基因组鉴定及其响应叶斑病的表达分析

doi:10.7668/hbnxb.20195347

摘要/Abstract

摘要：

为了解析绣球WRKY转录因子家族的成员特征及其在绣球叶斑病应答中的作用,利用生物信息学方法,对绣球无尽夏基因组中的WRKY家族成员进行全基因组鉴定,并系统分析了蛋白质理化特征、基因结构、系统进化、共线性和家族成员在多主棒孢菌侵染下的表达模式。结果表明:绣球全基因组中共鉴定到84个非冗余的HmWRKY成员,均属于亲水性蛋白,在绣球18条染色体上呈现不均匀分布,编码氨基酸112~1 046个;所有成员可分为3个亚组,即Group Ⅰ~Group Ⅲ,均含有WRKYGQK和C2H2组成的DNA结合域;HmWRKY成员的序列长度变化较大,从512 bp到40 338 bp,并且检测到8个存在共线性的基因对,Ka/Ks的比值均小于1,说明HmWRKY家族在进化中趋于纯化选择;同时,18个HmWRKY成员在绣球叶斑病菌侵染后表现出显著差异表达的特性,其中9个上调表达、9个下调表达。以上结果表明,HmWRKY基因在绣球响应叶斑病中可能具有重要作用。

关键词: 绣球, WRKY基因家族, 转录因子, 叶斑病, 多主棒孢菌

Abstract:

To understand the member characteristics of the WRKY transcription factor family in hydrangea and their roles in response to leaf spot disease,this study utilized the bioinformatics method to characterize WRKY family members in Hydrangea macrophylla Endless Summer,and systematically analyzed the protein physicochemical features,gene structure,systematic evolution,collinearity and expression patterns under the infection with Corynespora cassiicola of WRKY family members.The results showed that there were 84 non-redundant HmWRKY members in H.macrophylla genome.All of the HmWRKYs were hydrophilic proteins and unevenly distributed on 18 chromosomes of H.macrophylla,encoding 112—1 046 amino acids.In addition,HmWRKY members were divided into 3 subgroups(Group Ⅰ—Group Ⅲ),which contained a conserved DNA binding domain composed of WRKYGQK and C2H2.The sequence length of HmWRKY varies greatly from 512 bp to 40 338 bp,and 8 collinear gene pairs with Ka/Ks ratios less than 1 were detected,indicating that the HmWRKY family experienced purification selection in evolution.18 HmWRKY members showed significantly differential expression after infection with C.cassiicola,of which 9 up-regulated and 9 down-regulated.The results indicated that these HmWRKY genes might play important roles in response to leaf spot disease of hydrangea.

Key words: Hydrangea macrophylla, WRKY gene family, Transcription factor, Leaf spot disease, Corynespora cassiicola

刘欣童, 陈慧杰, 陈双双, 冯景, 齐香玉, 周惠民, 金玉妍, 孙明, 邓衍明. 绣球WRKY基因家族全基因组鉴定及其响应叶斑病的表达分析[J]. 华北农学报, 2025, 40(1): 93-103. doi: 10.7668/hbnxb.20195347.

LIU Xintong, CHEN Huijie, CHEN Shuangshuang, FENG Jing, QI Xiangyu, ZHOU Huimin, JIN Yuyan, SUN Ming, DENG Yanming. Genome-wide Characterization of WRKY Gene Family in Hydrangea macrophylla and Expression Analysis in Response to Leaf Spot Disease[J]. Acta Agriculturae Boreali-Sinica, 2025, 40(1): 93-103. doi: 10.7668/hbnxb.20195347.

http://www.hbnxb.net/CN/Y2025/V40/I1/93

图/表 10

参考文献 46

[1]	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 S, Ge C H, Shi H F, Min Z, Wang G P, Li W M. 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
[2]	梁细妹, 秦双双, 韦范, 韦桂丽, 林泉, 梁莹. 转录因子在植物干旱应激中的功能研究进展[J]. 生物资源, 2024, 46(3):220-230.doi:10.14188/j.ajsh.2024.03.002.
	Liang X M, Qin S S, Wei F, Wei G L, Lin Q, Liang Y. Research progress on the function of transcription factors in plant drought stress[J]. Biotic Resources, 2024, 46(3):220-230.
[3]	Li W X, Pang S Y, Lu Z G, Jin B. Function and mechanism of WRKY transcription factors in abiotic stress responses of plants[J]. Plants, 2020, 9(11):1515.doi:10.3390/plants9111515.
[4]	Rushton P J, Somssich I E, Ringler P, Shen Q J. WRKY transcription factors[J]. Trends in Plant Science, 2010, 15(5):247-258.doi:10.1016/j.tplants.2010.02.006. pmid: 20304701
[5]	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.
[6]	Huang H, Zhao W C, Li C H, Qiao H, Song S S, Yang R, Sun L L, Ma J L, Ma X C, Wang S H. SlVQ15 interacts with jasmonate-ZIM domain proteins and SlWRKY31 to regulate defense response in tomato[J]. Plant Physiology, 2022, 190(1):828-842.doi:10.1093/plphys/kiac275. pmid: 35689622
[7]	Xie W Y, Ke Y G, Cao J B, Wang S P, Yuan M. Knock out of transcription factor WRKY53 thickens sclerenchyma cell walls,confers bacterial blight resistance[J]. Plant Physiology, 2021, 187(3):1746-1761.doi:10.1093/plphys/kiab400.
[8]	李辉, 康泽培, 邱财生, 戴志刚, 邱化蛟. 红麻WRKY基因家族鉴定及其盐胁迫下的表达分析[J]. 华北农学报, 2022, 37(4):71-81.doi:10.7668/hbnxb.20192961.
	Li H, Kang Z P, Qiu C S, Dai Z G, Qiu H J. Identification of kenaf WRKY gene family and its expression analysis under salt stress[J]. Acta Agriculturae Boreali-Sinica, 2022, 37(4):71-81. doi: 10.7668/hbnxb.20192961
[9]	温海洋, 朱紫童, 湛佳伟, 李长, 武博寒, 杨永霞, 张松涛, 贾宏昉. 烟草转录因子NtWRKY11基因克隆、亚细胞定位及表达模式分析[J]. 华北农学报, 2024, 39(3):60-66.doi:10.7668/hbnxb.20194660.
	Wen H Y, Zhu Z T, Zhan J W, Li C, Wu B H, Yang Y X, Zhang S T, Jia H F. Cloning,subcellular localization and expression pattern analysis of tobacco transcription factor NtWRKY11 gene[J]. Acta Agriculturae Boreali-Sinica, 2024, 39(3):60-66.
[10]	Yu Y A, He L Y, Wu Y X. Wheat WRKY transcription factor TaWRKY24 confers drought and salt tolerance in transgenic plants[J]. Plant Physiology and Biochemistry, 2023, 205:108137.doi:10.1016/j.plaphy.2023.108137.
[11]	Ge M M, Tang Y, Guan Y J, Lyu M C, Zhou C, Ma H L, Lü J Y. TaWRKY31,a novel WRKY transcription factor in wheat,participates in regulation of plant drought stress tolerance[J]. BMC Plant Biology, 2024, 24(1):27.doi:10.1186/s12870-023-04709-7.
[12]	陈慧杰, 严子桢, 齐香玉, 陈双双, 冯景, 金玉妍, 苗艳华, 邓衍明. 不同绣球品种耐寒性评价及主要指标筛选[J]. 植物资源与环境学报, 2024, 33(2):41-49.doi:3969/j.issn.1674-7895.2024.02.05.
	Chen H J, Yan Z Z, Qi X Y, Chen S S, Feng J, Jin Y Y, Miao Y H, Deng Y M. Evaluation of cold tolerance and screening of main indexes of different cultivars of Hydrangea macrophylla[J]. Journal of Plant Resources and Environment, 2024, 33(2):41-49.
[13]	毛俊丹, 陈慧杰, 齐香玉, 陈双双, 冯景, 金玉妍, 邓衍明, 张浩. 钢渣混合基质对大花绣球扦插生根的影响[J]. 应用生态学报, 2024, 35(8):2150-2158.doi:10.13287/j.1001-9332.202408.015.
	Mao J D, Chen H J, Qi X Y, Chen S S, Feng J, Jin Y Y, Deng Y M, Zhang H. Effects of steel slag mixed substrate on rooting of Hydrangea macrophylla cuttings[J]. Chinese Journal of Applied Ecology, 2024, 35(8):2150-2158.
[14]	王晓玥, 陈双双, 齐香玉, 冯景, 陈慧杰, 孙明, 邓衍明. 绣球花铝转运蛋白HmALMT11的生物信息学及其表达特性分析[J]. 华北农学报, 2024, 39(4):94-101.doi:10.7668/hbnxb.20194790.
	Wang X Y, Chen S S, Qi X Y, Feng J, Chen H J, Sun M, Deng Y M. Bioinformatics analysis and expression profiling of the aluminum transporter HmALMT11 in Hydrangea macrophylla[J]. Acta Agriculturae Boreali-Sinica, 2024, 39(4):94-101.
[15]	Chen S S, Qi X Y, Feng J, Chen H J, Qin Z Y, Wang H D, Deng Y M. Biochemistry and transcriptome analyses reveal key genes and pathways involved in high-aluminum stress response and tolerance in hydrangea sepals[J]. Plant Physiology and Biochemistry, 2022, 185:268-278.doi:10.1016/j.plaphy.2022.06.008. pmid: 35724621
[16]	邓衍明, 韩勇, 齐香玉, 孙晓波, 贾新平, 陈双双. 绣球属植物种质资源分析及其花色可调性和叶斑病抗性比较[J]. 植物资源与环境学报, 2018, 27(4):90-100.doi:10.3969/j.issn.1674-7895.2018.04.10.
	Deng Y M, Han Y, Qi X Y, Sun X B, Jia X P, Chen S S. Analysis on germplasm resources of species in Hydrangea Linn.and comparisons on their flower color variability and resistance to leaf spot disease[J]. Journal of Plant Resources and Environment, 2018, 27(4):90-100.
[17]	陈慧杰, 邓衍明, 齐香玉, 陈双双, 冯景, 韩勇, 王华娣, 秦紫艺. 绣球叶斑病病原鉴定及其生物学特性[J]. 江苏农业科学, 2022, 50(7):106-111.doi:10.15889/j.issn.1002-1302.2022.07.016.
	Chen H J, Deng Y M, Qi X Y, Chen S S, Feng J, Han Y, Wang H D, Qin Z Y. Identification of pathogen from hydrangea leaf spot disease and its biological characteristics[J]. Jiangsu Agricultural Sciences, 2022, 50(7):106-111.
[18]	Zhu J Z, Chen J, Wang Y, Li C X, Zhang C J, He A G, Zhong J. Leaf spot of Hydrangea macrophylla caused by Corynespora cassiicola in China[J]. Canadian Journal of Plant Pathology, 2020, 42(1):125-132.doi:10.1080/07060661.2019.1632934.
[19]	Wu X B, Alexander L W. Genome-wide association studies for inflorescence type and remontancy in Hydrangea macrophylla[J]. Horticulture Research, 2020, 7:27.doi:10.1038/s41438-020-0255-y.
[20]	Chen C J, Chen H, Zhang Y, Thomas H R, Frank M H, He Y H, Xia R. TBtools:an integrative toolkit developed for interactive analyses of big biological data[J]. Molecular Plant, 2020, 13(8):1194-1202.doi:10.1016/j.molp.2020.06.009.
[21]	Crooks G E, Hon G, Chandonia J M, Brenner S E. WebLogo:a sequence logo generator[J]. Genome Research, 2004, 14(6):1188-1190.doi:10.1101/gr.849004. pmid: 15173120
[22]	Tamura K, Stecher G, Peterson D, Filipski A, Kumar S. MEGA6:molecular evolutionary genetics analysis version 6.0[J]. Molecular Biology and Evolution, 2013, 30(12):2725-2729.doi:10.1093/molbev/mst197.
[23]	陈双双, 齐香玉, 冯景, 陈慧杰, 王华娣, 秦紫艺, 邓衍明. 铝处理下绣球实时荧光定量PCR内参基因筛选及验证[J]. 华北农学报, 2021, 36(2):9-18.doi:10.7668/hbnxb.20191716.
	Chen S S, Qi X Y, Feng J, Chen H J, Wang H D, Qin Z Y, Deng Y M. Selection and validation of reference genes for qRT-PCR gene expression analysis in Hydrangea macrophylla under aluminum treatment[J]. Acta Agriculturae Boreali-Sinica, 2021, 36(2):9-18.
[24]	Wang Q S, Wang M H, Zhang X Z, Hao B J, Kaushik S K, Pan Y C. WRKY gene family evolution in Arabidopsis thaliana[J]. Genetica, 2011, 139(8):973-983.doi:10.1007/s10709-011-9599-4.
[25]	Chen L G, Zhang L P, Xiang S Y, Chen Y L, Zhang H Y, Yu D Q. The transcription factor WRKY75 positively regulates jasmonate-mediated plant defense to necrotrophic fungal pathogens[J]. Journal of Experimental Botany, 2021, 72(4):1473-1489.doi:10.1093/jxb/eraa529. pmid: 33165597
[26]	Wani S H, Anand S, Singh B, Bohra A, Joshi R. WRKY transcription factors and plant defense responses:latest discoveries and future prospects[J]. Plant Cell Reports, 2021, 40(7):1071-1085.doi:10.1007/s00299-021-02691-8.
[27]	Kim K C, Fan B F, Chen Z X. Pathogen-induced Arabidopsis WRKY7 is a transcriptional repressor and enhances plant susceptibility to Pseudomonas syringae[J]. Plant Physiology, 2006, 142(3):1180-1192.doi:10.1104/pp.106.082487.
[28]	Liu X F, Song Y Z, Xing F Y, Wang N, Wen F J, Zhu C X. GhWRKY25,a group I WRKY gene from cotton,confers differential tolerance to abiotic and biotic stresses in transgenic Nicotiana benthamiana[J]. Protoplasma, 2016, 253(5):1265-1281.doi:10.1007/s00709-015-0885-3.
[29]	Wang H H, Meng J, Peng X X, Tang X K, Zhou P L, Xiang J H, Deng X B. Rice WRKY4 acts as a transcriptional activator mediating defense responses toward Rhizoctonia solani,the causing agent of rice sheath blight[J]. Plant Molecular Biology, 2015, 89(1/2):157-171.doi:10.1007/s11103-015-0360-8.
[30]	Liu X Q, Bai X Q, Qian Q, Wang X J, Chen M S, Chu C C. OsWRKY03,a rice transcriptional activator that functions in defense signaling pathway upstream of OsNPR1[J]. Cell Research, 2005, 15(8):593-603.doi:10.1038/sj.cr.7290329.
[31]	Han M, Ryu H S, Kim C Y, Park D S, Ahn Y K, Jeon J S. OsWRKY30 is a transcription activator that enhances rice resistance to the Xanthomonas oryzae pathovar oryzae[J]. Journal of Plant Biology, 2013, 56(4):258-265.doi:10.1007/s12374-013-0160-0.
[32]	Chujo T, Miyamoto K, Ogawa S, Masuda Y, Shimizu T, Kishi-Kaboshi M, Takahashi A, Nishizawa Y, Minami E, Nojiri H, Yamane H, Okada K. Overexpression of phosphomimic mutated OsWRKY53 leads to enhanced blast resistance in rice[J]. PLoS One, 2014, 9(6):e98737.doi:10.1371/journal.pone.0098737.
[33]	Yang B, Jiang Y Q, Rahman M H, Deyholos M K, Kav N N V. Identification and expression analysis of WRKY transcription factor genes in canola(Brassica napus L.)in response to fungal pathogens and hormone treatments[J]. BMC Plant Biology, 2009, 9(1):68.doi:10.1186/1471-2229-9-68.
[34]	Wu W, Fu P N, Lu J. Grapevine WRKY transcription factors[J]. Fruit Research, 2022, 2(1):1-8.doi:10.48130/frures-2022-0010.
[35]	Guo R R, Qiao H B, Zhao J, Wang X H, Tu M X, Guo C L, Wan R, Li Z, Wang X P. The grape VlWRKY3 gene promotes abiotic and biotic stress tolerance in transgenic Arabidopsis thaliana[J]. Frontiers in Plant Science, 2018, 9:545.doi:10.3389/fpls.2018.00545.
[36]	Merz P R, Moser T, Höll J, Kortekamp A, Buchholz G, Zyprian E, Bogs J. The transcription factor VvWRKY33 is involved in the regulation of grapevine(Vitis vinifera)defense against the oomycete pathogen Plasmopara viticola[J]. Physiologia Plantarum, 2015, 153(3):365-380.doi:10.1111/ppl.12251.
[37]	Liu W D, Yan C H, Li R M, Chen G Y, Wang X Q, Wen Y Q, Zhang C H, Wang X P, Xu Y, Wang Y J. VqMAPK3/VqMAPK6,VqWRKY33,and VqNSTS3 constitute a regulatory node in enhancing resistance to powdery mildew in grapevine[J]. Horticulture Research, 2023, 10(7):uhad116.doi:10.1093/hr/uhad116.
[38]	Yu D S, Fan R C, Zhang L, Xue P Y, Liao L B, Hu M Z, Cheng Y J, Li J E, Qi T, Jing S J, Wang Q Y, Bhatt A, Shen Q H. HvWRKY2 acts as an immunity suppressor and targets HvCEBiP to regulate powdery mildew resistance in barley[J]. The Crop Journal, 2023, 11(1):99-107.doi:10.1016/j.cj.2022.05.010.
[39]	Adachi H, Ishihama N, Nakano T, Yoshioka M, Yoshioka H. Nicotiana benthamiana MAPK-WRKY pathway confers resistance to a necrotrophic pathogen Botrytis cinerea[J]. Plant Signaling & Behavior, 2016, 11(6):e1183085.doi:10.1080/15592324.2016.1183085.
[40]	Sun S S, Ren Y X, Wang D X, Farooq T, He Z F, Zhang C, Li S F, Yang X L, Zhou X P. A group I WRKY transcription factor regulates mulberry mosaic dwarf-associated virus-triggered cell death in Nicotiana benthamiana[J]. Molecular Plant Pathology, 2022, 23(2):237-253.doi:10.1111/mpp.13156.
[41]	Oh S K, Baek K H, Park J M, Yi S Y, Yu S H, Kamoun S, Choi D. Capsicum annuum WRKY protein CaWRKY1 is a negative regulator of pathogen defense[J]. New Phytologist, 2008, 177(4):977-989.doi:10.1111/j.1469-8137.2007.02310.x.
[42]	Liu X T, Li D D, Zhang S Y, Xu Y L, Zhang Z. Genome-wide characterization of the rose(Rosa chinensis)WRKY family and role of RcWRKY41 in gray mold resistance[J]. BMC Plant Biology, 2019, 19(1):522.doi:10.1186/s12870-019-2139-6.
[43]	丁夏威, 田沺, 陈健文, 陈景益, 王章英, 邹宏达, 黄立飞. 甘薯WRKY基因家族鉴定及其响应茎腐病侵染的表达分析[J]. 江苏农业科学, 2024, 52(21): 25-39. doi:10.15889/j.issn.1002-1302.2024.21.004.
	Ding X W, Tian T, Chen J W, Chen J Y, Wang Z Y, Zou H D, Huang L F. Genome-wide characterization of WRKY gene family and the expression pattern during stem rot disease inoculation in sweet potato[J]. Jiangsu Agricultural Sciences, 2024, 52(21): 25-39.
[44]	Dou L L, Zhang X H, Pang C Y, Song M Z, Wei H L, Fan S L, Yu S X. Genome-wide analysis of the WRKY gene family in cotton[J]. Molecular Genetics and Genomics, 2014, 289(6):1103-1121.doi:10.1007/s00438-014-0872-y. pmid: 24942461
[45]	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.
[46]	Ye H, Qiao L Y, Guo H Y, Guo L P, Ren F, Bai J F, Wang Y K. Genome-wide identification of wheat WRKY gene family reveals that TaWRKY75-a is referred to drought and salt resistances[J]. Frontiers in Plant Science, 2021, 12:663118.doi:10.3389/fpls.2021.663118.

基因名称 Gene name	正向引物 (5'—3') Forward primer (5'—3')	反向引物 (5'—3') Reverse primer (5'—3')
HmWRKY13	CTCTTCGCTTGGGATTTGGCTCAT	TGGTTCGCAGGGTCAGATAGTGC
HmWRKY15	AGACATTAGTTGATTGTTCGGTGGG	AATAAGGGTCGTGCCTTGGCTGA
HmWRKY18	ACCAGCCGAGGAGTGGGTGAACT	GCTACCGCCGAGCAGACAAGAAT
HmWRKY19	TTACCATCCCGATCATTTGACGA	GACGAGGGTTGTTGCTTGCTGTC
HmWRKY36	GCTGAAGTTGATCCCTTGATGGC	CGGTCGAAACCCGAGAATTGTTA
HmWRKY40	CCATTCCCTACTATCACATTGGACC	GCCGTTGCTTCCCTCATCATCTC
HmWRKY71	ACAGAACTGCCACTCCGCAGAAA	GAAGAAGGCTCCGATGAAGACGA
HmWRKY79	GCACCCACCATTGATTCACAGAG	GGCACCCACAAAGGCAAGTAAGA

基因名称 Gene name	正向引物 (5'—3') Forward primer (5'—3')	反向引物 (5'—3') Reverse primer (5'—3')
HmWRKY13	CTCTTCGCTTGGGATTTGGCTCAT	TGGTTCGCAGGGTCAGATAGTGC
HmWRKY15	AGACATTAGTTGATTGTTCGGTGGG	AATAAGGGTCGTGCCTTGGCTGA
HmWRKY18	ACCAGCCGAGGAGTGGGTGAACT	GCTACCGCCGAGCAGACAAGAAT
HmWRKY19	TTACCATCCCGATCATTTGACGA	GACGAGGGTTGTTGCTTGCTGTC
HmWRKY36	GCTGAAGTTGATCCCTTGATGGC	CGGTCGAAACCCGAGAATTGTTA
HmWRKY40	CCATTCCCTACTATCACATTGGACC	GCCGTTGCTTCCCTCATCATCTC
HmWRKY71	ACAGAACTGCCACTCCGCAGAAA	GAAGAAGGCTCCGATGAAGACGA
HmWRKY79	GCACCCACCATTGATTCACAGAG	GGCACCCACAAAGGCAAGTAAGA

基因名 Gene name	染色体 Chromosome	起始/Mb Start	终止/Mb End	编码氨基酸数 Number of amino acids	分子质量/ku Molecular weight	等电点 Isoelectric point	疏水系数 Hydrophobic coefficient
HmWRKY1	Chr01	13.683	13.695	732	80.782	6.11	-0.704
HmWRKY2	Chr01	79.282	79.287	364	40.177	8.78	-0.698
HmWRKY3	Chr01	95.448	95.452	565	62.043	8.34	-0.918
HmWRKY4	Chr01	115.941	115.943	256	29.060	5.04	-1.012
HmWRKY5	Chr01	116.506	116.507	283	31.296	6.25	-0.739
HmWRKY6	Chr02	83.419	83.429	507	55.480	6.82	-0.881
HmWRKY7	Chr02	86.751	86.767	1 046	113.474	6.25	-0.372
HmWRKY8	Chr02	111.319	111.323	466	51.692	9.01	-0.949
HmWRKY9	Chr02	114.086	114.089	112	12.937	9.59	-1.220
HmWRKY10	Chr02	120.571	120.573	604	66.932	7.69	-0.939
HmWRKY11	Chr03	89.989	89.990	348	37.964	9.57	-0.669
HmWRKY12	Chr03	91.032	91.033	348	38.032	9.57	-0.676
HmWRKY13	Chr04	12.983	13.000	525	57.528	8.15	-0.740
HmWRKY14	Chr04	22.807	22.815	239	27.579	8.99	-0.855
HmWRKY15	Chr04	92.267	92.269	281	31.524	8.37	-0.626
HmWRKY16	Chr04	110.396	110.404	543	59.178	7.10	-0.722
HmWRKY17	Chr05	1.284	1.286	428	46.985	6.70	-0.666
HmWRKY18	Chr05	1.539	1.540	253	29.168	5.02	-1.028
HmWRKY19	Chr05	83.729	83.735	332	36.473	9.52	-0.742
HmWRKY20	Chr05	101.262	101.264	158	18.187	9.43	-0.969
HmWRKY21	Chr06	0.794	0.797	224	25.434	6.55	-0.975
HmWRKY22	Chr06	8.941	8.946	468	51.251	6.45	-0.842
HmWRKY23	Chr06	9.292	9.295	306	33.568	5.70	-0.698
HmWRKY24	Chr06	56.424	56.428	292	32.865	8.42	-0.683
HmWRKY25	Chr07	22.245	22.250	483	53.027	6.68	-0.845
HmWRKY26	Chr07	33.488	33.489	284	31.865	5.39	-0.923
HmWRKY27	Chr07	42.898	42.899	285	31.932	5.39	-0.916
HmWRKY28	Chr07	53.864	53.868	313	35.034	6.60	-0.811
HmWRKY29	Chr07	59.481	59.485	309	34.531	6.56	-0.795
HmWRKY30	Chr07	110.028	110.031	593	64.482	6.22	-0.644
HmWRKY31	Chr07	119.099	119.100	318	34.535	9.56	-0.667
HmWRKY32	Chr07	121.753	121.754	312	34.110	9.49	-0.720
HmWRKY33	Chr07	132.340	132.355	512	56.360	5.20	-0.828
HmWRKY34	Chr08	115.250	115.251	134	15.508	5.74	-0.723
HmWRKY35	Chr08	117.781	117.783	196	22.184	9.30	-0.854
HmWRKY36	Chr08	122.276	122.278	337	38.202	5.29	-0.734
HmWRKY37	Chr09	83.549	83.550	124	13.819	7.79	-0.737
HmWRKY38	Chr09	86.770	86.773	638	70.111	6.09	-0.852
HmWRKY39	Chr09	88.002	88.006	638	70.045	6.06	-0.851
HmWRKY40	Chr09	102.949	102.957	510	56.144	8.05	-0.652
HmWRKY41	Chr09	104.708	104.713	610	66.257	6.71	-0.736
HmWRKY42	Chr09	107.864	107.868	738	80.527	5.90	-0.783
HmWRKY43	Chr10	88.252	88.254	181	20.576	9.44	-0.824
HmWRKY44	Chr10	93.157	93.160	181	20.576	9.44	-0.824
HmWRKY45	Chr10	115.441	115.443	331	36.416	6.25	-0.674
HmWRKY46	Chr10	119.771	119.772	329	36.320	6.25	-0.697
HmWRKY47	Chr11	49.019	49.021	355	39.403	5.74	-0.762
HmWRKY48	Chr11	88.046	88.050	444	50.000	5.59	-0.905
HmWRKY49	Chr12	69.607	69.615	211	24.342	6.50	-0.399
HmWRKY50	Chr12	87.386	87.389	351	38.375	9.74	-0.621
HmWRKY51	Chr13	27.441	27.446	505	54.716	5.59	-0.565
HmWRKY52	Chr13	36.015	36.026	415	46.128	6.61	-0.962
HmWRKY53	Chr13	41.469	41.474	505	54.737	5.51	-0.584
HmWRKY54	Chr13	119.199	119.201	335	36.514	9.51	-0.637
HmWRKY55	Chr13	137.123	137.125	336	36.595	9.57	-0.627
HmWRKY56	Chr14	8.248	8.252	224	25.620	6.15	-0.926
HmWRKY57	Chr14	12.296	12.298	185	21.330	5.29	-1.147
HmWRKY58	Chr14	47.904	47.910	232	26.352	6.91	-0.625
HmWRKY59	Chr14	49.227	49.232	300	34.243	6.77	-0.785
HmWRKY60	Chr14	105.534	105.536	316	35.538	5.94	-0.566
HmWRKY61	Chr14	107.698	107.699	308	34.660	5.44	-0.658
HmWRKY62	Chr15	16.795	16.797	307	33.868	5.23	-0.740
HmWRKY63	Chr15	18.197	18.209	488	52.812	6.08	-0.716
HmWRKY64	Chr15	62.669	62.681	211	24.562	5.98	-0.951
HmWRKY65	Chr15	84.245	84.248	495	55.368	5.27	-0.806
HmWRKY66	Chr16	5.706	5.710	303	34.270	5.63	-0.713
HmWRKY67	Chr16	7.562	7.567	182	21.128	9.26	-0.911
HmWRKY68	Chr16	7.588	7.590	303	33.513	5.28	-0.825
HmWRKY69	Chr16	9.131	9.136	183	21.188	9.16	-0.931
HmWRKY70	Chr16	9.159	9.161	303	33.495	5.27	-0.828
HmWRKY71	Chr16	21.874	21.895	420	47.909	9.22	-0.660
HmWRKY72	Chr16	54.006	54.046	230	26.153	8.53	-1.081
HmWRKY73	Chr17	19.090	19.093	315	35.130	6.27	-0.885
HmWRKY74	Chr17	29.651	29.653	267	29.792	5.63	-0.688
HmWRKY75	Chr17	60.788	60.793	535	58.326	6.32	-0.651
HmWRKY76	Chr17	66.838	66.844	526	57.605	6.39	-0.657
HmWRKY77	Chr17	78.718	78.719	124	13.916	7.79	-0.675
HmWRKY78	Chr17	78.765	78.765	124	13.916	7.79	-0.675
HmWRKY79	Chr17	78.811	78.812	124	13.916	7.79	-0.675
HmWRKY80	Chr17	105.308	105.310	322	36.103	6.67	-0.847
HmWRKY81	Chr18	4.759	4.761	308	34.809	5.73	-0.732
HmWRKY82	Chr18	13.134	13.139	322	36.188	6.86	-0.683
HmWRKY83	Chr18	13.154	13.161	355	39.032	5.64	-0.728
HmWRKY84	Chr18	107.003	107.008	297	33.288	5.05	-0.740

基因名 Gene name	染色体 Chromosome	起始/Mb Start	终止/Mb End	编码氨基酸数 Number of amino acids	分子质量/ku Molecular weight	等电点 Isoelectric point	疏水系数 Hydrophobic coefficient
HmWRKY1	Chr01	13.683	13.695	732	80.782	6.11	-0.704
HmWRKY2	Chr01	79.282	79.287	364	40.177	8.78	-0.698
HmWRKY3	Chr01	95.448	95.452	565	62.043	8.34	-0.918
HmWRKY4	Chr01	115.941	115.943	256	29.060	5.04	-1.012
HmWRKY5	Chr01	116.506	116.507	283	31.296	6.25	-0.739
HmWRKY6	Chr02	83.419	83.429	507	55.480	6.82	-0.881
HmWRKY7	Chr02	86.751	86.767	1 046	113.474	6.25	-0.372
HmWRKY8	Chr02	111.319	111.323	466	51.692	9.01	-0.949
HmWRKY9	Chr02	114.086	114.089	112	12.937	9.59	-1.220
HmWRKY10	Chr02	120.571	120.573	604	66.932	7.69	-0.939
HmWRKY11	Chr03	89.989	89.990	348	37.964	9.57	-0.669
HmWRKY12	Chr03	91.032	91.033	348	38.032	9.57	-0.676
HmWRKY13	Chr04	12.983	13.000	525	57.528	8.15	-0.740
HmWRKY14	Chr04	22.807	22.815	239	27.579	8.99	-0.855
HmWRKY15	Chr04	92.267	92.269	281	31.524	8.37	-0.626
HmWRKY16	Chr04	110.396	110.404	543	59.178	7.10	-0.722
HmWRKY17	Chr05	1.284	1.286	428	46.985	6.70	-0.666
HmWRKY18	Chr05	1.539	1.540	253	29.168	5.02	-1.028
HmWRKY19	Chr05	83.729	83.735	332	36.473	9.52	-0.742
HmWRKY20	Chr05	101.262	101.264	158	18.187	9.43	-0.969
HmWRKY21	Chr06	0.794	0.797	224	25.434	6.55	-0.975
HmWRKY22	Chr06	8.941	8.946	468	51.251	6.45	-0.842
HmWRKY23	Chr06	9.292	9.295	306	33.568	5.70	-0.698
HmWRKY24	Chr06	56.424	56.428	292	32.865	8.42	-0.683
HmWRKY25	Chr07	22.245	22.250	483	53.027	6.68	-0.845
HmWRKY26	Chr07	33.488	33.489	284	31.865	5.39	-0.923
HmWRKY27	Chr07	42.898	42.899	285	31.932	5.39	-0.916
HmWRKY28	Chr07	53.864	53.868	313	35.034	6.60	-0.811
HmWRKY29	Chr07	59.481	59.485	309	34.531	6.56	-0.795
HmWRKY30	Chr07	110.028	110.031	593	64.482	6.22	-0.644
HmWRKY31	Chr07	119.099	119.100	318	34.535	9.56	-0.667
HmWRKY32	Chr07	121.753	121.754	312	34.110	9.49	-0.720
HmWRKY33	Chr07	132.340	132.355	512	56.360	5.20	-0.828
HmWRKY34	Chr08	115.250	115.251	134	15.508	5.74	-0.723
HmWRKY35	Chr08	117.781	117.783	196	22.184	9.30	-0.854
HmWRKY36	Chr08	122.276	122.278	337	38.202	5.29	-0.734
HmWRKY37	Chr09	83.549	83.550	124	13.819	7.79	-0.737
HmWRKY38	Chr09	86.770	86.773	638	70.111	6.09	-0.852
HmWRKY39	Chr09	88.002	88.006	638	70.045	6.06	-0.851
HmWRKY40	Chr09	102.949	102.957	510	56.144	8.05	-0.652
HmWRKY41	Chr09	104.708	104.713	610	66.257	6.71	-0.736
HmWRKY42	Chr09	107.864	107.868	738	80.527	5.90	-0.783
HmWRKY43	Chr10	88.252	88.254	181	20.576	9.44	-0.824
HmWRKY44	Chr10	93.157	93.160	181	20.576	9.44	-0.824
HmWRKY45	Chr10	115.441	115.443	331	36.416	6.25	-0.674
HmWRKY46	Chr10	119.771	119.772	329	36.320	6.25	-0.697
HmWRKY47	Chr11	49.019	49.021	355	39.403	5.74	-0.762
HmWRKY48	Chr11	88.046	88.050	444	50.000	5.59	-0.905
HmWRKY49	Chr12	69.607	69.615	211	24.342	6.50	-0.399
HmWRKY50	Chr12	87.386	87.389	351	38.375	9.74	-0.621
HmWRKY51	Chr13	27.441	27.446	505	54.716	5.59	-0.565
HmWRKY52	Chr13	36.015	36.026	415	46.128	6.61	-0.962
HmWRKY53	Chr13	41.469	41.474	505	54.737	5.51	-0.584
HmWRKY54	Chr13	119.199	119.201	335	36.514	9.51	-0.637
HmWRKY55	Chr13	137.123	137.125	336	36.595	9.57	-0.627
HmWRKY56	Chr14	8.248	8.252	224	25.620	6.15	-0.926
HmWRKY57	Chr14	12.296	12.298	185	21.330	5.29	-1.147
HmWRKY58	Chr14	47.904	47.910	232	26.352	6.91	-0.625
HmWRKY59	Chr14	49.227	49.232	300	34.243	6.77	-0.785
HmWRKY60	Chr14	105.534	105.536	316	35.538	5.94	-0.566
HmWRKY61	Chr14	107.698	107.699	308	34.660	5.44	-0.658
HmWRKY62	Chr15	16.795	16.797	307	33.868	5.23	-0.740
HmWRKY63	Chr15	18.197	18.209	488	52.812	6.08	-0.716
HmWRKY64	Chr15	62.669	62.681	211	24.562	5.98	-0.951
HmWRKY65	Chr15	84.245	84.248	495	55.368	5.27	-0.806
HmWRKY66	Chr16	5.706	5.710	303	34.270	5.63	-0.713
HmWRKY67	Chr16	7.562	7.567	182	21.128	9.26	-0.911
HmWRKY68	Chr16	7.588	7.590	303	33.513	5.28	-0.825
HmWRKY69	Chr16	9.131	9.136	183	21.188	9.16	-0.931
HmWRKY70	Chr16	9.159	9.161	303	33.495	5.27	-0.828
HmWRKY71	Chr16	21.874	21.895	420	47.909	9.22	-0.660
HmWRKY72	Chr16	54.006	54.046	230	26.153	8.53	-1.081
HmWRKY73	Chr17	19.090	19.093	315	35.130	6.27	-0.885
HmWRKY74	Chr17	29.651	29.653	267	29.792	5.63	-0.688
HmWRKY75	Chr17	60.788	60.793	535	58.326	6.32	-0.651
HmWRKY76	Chr17	66.838	66.844	526	57.605	6.39	-0.657
HmWRKY77	Chr17	78.718	78.719	124	13.916	7.79	-0.675
HmWRKY78	Chr17	78.765	78.765	124	13.916	7.79	-0.675
HmWRKY79	Chr17	78.811	78.812	124	13.916	7.79	-0.675
HmWRKY80	Chr17	105.308	105.310	322	36.103	6.67	-0.847
HmWRKY81	Chr18	4.759	4.761	308	34.809	5.73	-0.732
HmWRKY82	Chr18	13.134	13.139	322	36.188	6.86	-0.683
HmWRKY83	Chr18	13.154	13.161	355	39.032	5.64	-0.728
HmWRKY84	Chr18	107.003	107.008	297	33.288	5.05	-0.740

基因1 Gene 1	基因2 Gene 2	非同义突变 (Ka) Non synonymous mutation(Ka)	同义突变 (Ks) Synonymous mutation(Ks)	Ka/Ks	有效长度/bp Effective length	平均同义突变位点 Average S-sites	平均非同义突变位点 Average N-sites
HmWRKY3	HmWRKY10	0.272 1	1.742 1	0.156 2	1 599	372.75	1 226.25
HmWRKY31	HmWRKY32	0.008 5	0.051 1	0.165 4	936	222.50	713.50
HmWRKY38	HmWRKY39	0.004 1	0.009 3	0.437 1	1 914	433.08	1 480.92
HmWRKY38	HmWRKY53	0.200 0	0.837 7	0.238 7	1 512	348.33	1 163.67
HmWRKY38	HmWRKY51	0.205 9	0.832 2	0.247 4	1 512	349.42	1 162.58
HmWRKY55	HmWRKY54	0.005 2	0.012 6	0.416 6	1 005	240.17	764.83
HmWRKY65	HmWRKY48	0.164 8	0.657 9	0.250 5	1 284	269.00	1 015.00
HmWRKY75	HmWRKY76	0.025 9	0.073 6	0.351 8	1 575	356.67	1 218.33

选择文件类型/文献管理软件名称

选择包含的内容