小麦转录因子基因TaWRKY65-3A功能解析和分子标记开发

doi:10.3864/j.issn.0578-1752.2026.15.001

中国农业科学 ›› 2026, Vol. 59 ›› Issue (15): 3237-3251.doi: 10.3864/j.issn.0578-1752.2026.15.001

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

小麦转录因子基因TaWRKY65-3A功能解析和分子标记开发

张艳菲¹(), 吴子月¹, 薛志伟⁴, 马冬云¹, 康国章¹, 冯伟¹, 谢迎新¹, 景蕊莲², 王万章³, 毛新国²(), 王晨阳¹()

¹ 河南农业大学农学院/国家小麦工程技术研究中心，郑州 450046
² 中国农业科学院作物科学研究所/作物基因资源与育种全国重点实验室，北京 100081
³ 河南农业大学机电工程学院，郑州 450046
⁴ 安阳市农业科学院，河南安阳 455000

收稿日期:2025-12-12 接受日期:2026-02-06 出版日期:2026-08-01 发布日期:2026-08-03
通信作者:
王晨阳，Tel：0371-56552938；E-mail：xmzxwang@163.com。
毛新国，Tel：010-82105829；E-mail：maoxinguo@caas.cn
联系方式: 张艳菲，E-mail：yfz15838298834@163.com。
基金资助:
河南省科技攻关项目(242102110323); 河南省自然科学基金(242300421321)

Functional Characterization and Molecular Marker Development of the Wheat Transcription Factor Gene TaWRKY65-3A

ZHANG YanFei¹(), WU ZiYue¹, XUE ZhiWei⁴, MA DongYun¹, KANG GuoZhang¹, FENG Wei¹, XIE YingXin¹, JING RuiLian², WANG WanZhang³, MAO XinGuo²(), WANG ChenYang¹()

¹ College of Agriculture, Henan Agricultural University/National Wheat Engineering Technology Research Center, Zhengzhou 450046
² Institute of Crop Sciences, Chinese Academy of Agricultural Sciences/State Key Laboratory of Crop Gene Resources and Breeding, Beijing 100081
³ College of Mechanical & Electrical Engineering, Henan Agricultural University, Zhengzhou 450046
⁴ Anyang Academy of Agricultural Sciences, Anyang 455000, Henan

Received:2025-12-12 Accepted:2026-02-06 Published:2026-08-01 Online:2026-08-03

摘要/Abstract

摘要：

【目的】WRKY转录因子作为植物所特有的转录因子家族，在植物生长、发育、营养吸收、生物和非生物胁迫应答中具有重要作用。揭示小麦转录因子基因TaWRKY65-3A的功能，并开发分子标记，为通过分子育种改良小麦种质提供基因资源。【方法】以小麦品种旱选10号cDNA为模板，克隆TaWRKY65-3A序列；利用SMART网站对其编码蛋白的结构域进行分析；通过qRT-PCR对TaWRKY65-3A在ABA、PEG和NaCl胁迫下的表达模式进行分析；通过小麦基因组变异联合数据库对TaWRKY65-3A序列多态性进行预测，并通过21份遗传多样性丰富的小麦种质进行验证，进一步开发分子标记；以323份小麦构成的自然群体为材料进行TaWRKY65-3A单倍型与表型性状的关联分析；分析189份现代育成小麦品种的单倍型，明确优异单倍型在我国小麦育种过程中受选择的趋势。【结果】TaWRKY65-3A全长1 998 bp，包含2个外显子和1个内含子，具有WRKY转录因子家族所特有的WRKY Domain和C₂H₂型锌指结构域；TaWRKY65-3A启动子区含有多种顺式作用元件，包括脱落酸和茉莉酸甲酯等激素应答元件和干旱、低温胁迫应答元件，且TaWRKY65-3A的表达受ABA、PEG和NaCl胁迫的影响；在TaWRKY65-3A区检测到6个SNP，组成3种单倍型Hap-3A-1、Hap-3A-2和Hap-3A-3；基于TaWRKY65-3A编码区1 265 bp（G/C）和非编码区1 658 bp（T/C）开发dCAPS标记，分别与多种环境下的千粒重和单株产量、每穗小穗数和穗粒数显著关联；在干旱、高温和干旱复合高温等多种不良环境下，单倍型Hap-3A-2与每穗小穗数多、穗粒数多、千粒重高和单株产量高显著相关，为优异单倍型，在我国小麦育种进程中受到正向选择。【结论】TaWRKY65-3A响应植物激素ABA、PEG和NaCl胁迫，在干旱、高温和干旱复合高温等多种极端环境下与产量和产量构成要素显著相关，Hap-3A-2是产量和产量构成要素高的优异单倍型。

关键词: 小麦, 转录因子, TaWRKY65-3A, 产量, dCAPS标记, 单倍型

Abstract:

【Objective】WRKY transcription factors are the plant-specific transcription factor families, and play a crucial role in plant growth, development, nutrient uptake, and response to biotic and abiotic stresses. Characterized the function of the wheat transcription factor gene TaWRKY65-3A and developed the molecular markers, providing genetic resources for wheat germplasm improvement through molecular breeding in this research.【Method】The sequence of TaWRKY65-3A was cloned from the cDNA of wheat variety Hanxuan 10. The domains were analyzed via the SMART website. The expression patterns of TaWRKY65-3A gene were analyzed by qRT-PCR under phytohormone ABA treatments, PEG and NaCl stresses. Predicted the polymorphisms of TaWRKY65-3A through the Wheat Genome Variation Joint Database, and validated them by using 21 wheat germplasms with rich genetic diversity and further developed molecular markers. Association analysis between TaWRKY65-3A haplotypes and phenotypic traits was carried out in a natural population consisting of 323 wheat accessions. The haplotype of 189 modern wheat cultivars from different released years were analyzed, aiming to clarify the selection trends of superior haplotypes during wheat breeding in China.【Result】TaWRKY65-3A is 1 998 bp in full length, comprising 2 exons and 1 intron, and harbors the characteristic WRKY domain and a C₂H₂-type zinc finger motif typical of the WRKY transcription factor family. The promoter region of TaWRKY65-3A contains a variety of cis-acting elements, including hormone-responsive elements, (e.g., to abscisic acid and methyl jasmonate) and stress-responsive elements (e.g., to drought and low temperature). Moreover, the expressions of TaWRKY65-3A was influenced by ABA, PEG and NaCl stresses. Six SNPs were identified in the TaWRKY65-3A region, forming three haplotypes: Hap-3A-1, Hap-3A-2, and Hap-3A-3. dCAPS markers were developed at positions of 1 265 bp (G/C) in the coding sequence and 1 658 bp (T/C) in the non-coding sequence, which were significantly associated with thousand-kernel weight, yield per plant, spikelet number per spike, and kernel number per spike across multiple environments. Notably, the haplotype Hap-3A-2 was demonstrated to be significantly correlated with higher spikelet number per spike, a great kernel number per spike, a higher thousand-kernel weight, and a higher yield per plant in multiple environments, including drought, high temperature, and the combined stress of drought and high temperature, and it was positively selected during wheat breeding in China.【Conclusion】TaWRKY65-3A gene responded to phytohormones ABA, PEG and NaCl stress, and were significantly correlated with spikelet number per spike, kernel number per spike, thousand-kernel weight and yield per plant under various environments such as drought, high temperature, and the combined stress of drought and high temperature. Hap-3A-2 is the favorable haplotype with higher spikelet number per spike, more kernel number per spike, higher thousand-kernel weight, and higher yield per plant. The molecular markers of dCAPS developed in this study provide valuable tools for marker-assisted breeding, facilitating the selection of germplasm with superior yield-related traits.

Key words: wheat, transcription factor, TaWRKY65-3A, yield, dCAPS marker, haplotype

张艳菲, 吴子月, 薛志伟, 马冬云, 康国章, 冯伟, 谢迎新, 景蕊莲, 王万章, 毛新国, 王晨阳. 小麦转录因子基因TaWRKY65-3A功能解析和分子标记开发[J]. 中国农业科学, 2026, 59(15): 3237-3251.

ZHANG YanFei, WU ZiYue, XUE ZhiWei, MA DongYun, KANG GuoZhang, FENG Wei, XIE YingXin, JING RuiLian, WANG WanZhang, MAO XinGuo, WANG ChenYang. Functional Characterization and Molecular Marker Development of the Wheat Transcription Factor Gene TaWRKY65-3A[J]. Scientia Agricultura Sinica, 2026, 59(15): 3237-3251.

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引物名称 Prime name	引物序列 Sequences (5´-3´)	用途 Usage
TaWRKY65-qF	CATCGAGGTGCACGACGAT	TaWRKY65表达 Expression of TaWRKY65
TaWRKY65-qR	GCGGCCGTGTCGACCTC
TaActin-qF	CTCCCTCACAACAACAACCGC
TaActin-qR	TACCAGGAACTTCCATACCAAC
OsActin-qF	AGTAGTCTCGCCCTGCTTCA
OsActin-qR	TTCGCCACTTTGTTCTCAACT
TaWRKY65-1300GFP-F	CCAAATCGACTCTAGAATGGACGGCGAGTGGAGC	亚细胞定位 Subcellular localization
TaWRKY65-1300GFP-R	TGCTCACCATGGTACCCGTCATGGCTGTGCCCTC	亚细胞定位 Subcellular localization
TaWRKY65-3A-1265F	TTGACCCCTCCTGCCATT	开发分子标记 Developing the molecular markers
TaWRKY65-3A-1265R	GCTCGTTCTTCCGACTCCAT
TaWRKY65-3A-1658F	GCTTTTGTTGGTTAGTTTTCTCAC
TaWRKY65-3A-1658R	GGGTTTGTATCCCTCTCAATCTC
TaWRKY65-3A-dCAPS-Sma I-1265F	CTCGTCCGGAAGGAGCCCGG
TaWRKY65-3A-dCAPS-Sma I-1265R	GTCGCCCGAGCCAGC
TaWRKY65-3A-dCAPS-Sac I-1658F	AAGCATCTTAGAGGCGGAGCT
TaWRKY65-3A-dCAPS-Sac I-1658R	TGGTCCAAACAAATGAATAAAAAC

顺式作用元件 Cis-acting element	数量 Numbers	生物学功能 Biological function
TGACG-motif	5	茉莉酸甲酯应答元件Cis-acting regulatory element involved in the MeJA-responsiveness
CGTCA-motif	5	茉莉酸甲酯应答元件Cis-acting regulatory element involved in the MeJA-responsiveness
ABRE	2	脱落酸应答元件Cis-acting element involved in the abscisic acid responsiveness
TC-rich repeats	1	防御和胁迫应答元件Cis-acting element involved in defense and stress responsiveness
LTR	1	低温应答元件Cis-acting element involved in low-temperature responsiveness
MBS	1	MYB蛋白结合元件MYB protein binding element

位置Sites (bp)						样本数量 Number of samples	占总样本百分比 Percentage of total samples (%)
809	1265	1658	1684	1758	1887	样本数量 Number of samples	占总样本百分比 Percentage of total samples (%)
T	G	T	T	G	A	40	27.6
T	C	T	C	A	A	58	40.0
C	G	C	C	G	G	24	16.6
T	C	T	C	G	A	4	2.8
T	C	T	T	G	A	2	1.4
T	C	T	T	A	A	2	1.4
C	G	T	T	G	G	2	1.4
T	G	C	C	G	G	1	0.7
C	G	T	C	G	G	1	0.7
C	G	C	C	G	A	1	0.7
/	/	/	/	/	/	10	6.9
合计Total						145	100

环境 Environments	年份 Years	地点 Sites	处理 Treatments	性状（P值）Traits (P value)
				dCAPS-1265		dCAPS-1658
				千粒重 Thousand grain weight	单株产量 Grain yield per plant	穗粒数 Grain number per spike	每穗小穗数 Spikelets per spike
E1	2015	顺义 SY	DS+HT	0.0030**	0.6016ns	0.1759ns	0.0005***
E2			DS	0.0107*	0.5323ns	0.0043**	0.0035**
E3			WW+HT	0.0772ns	0.3067ns	0.0002***	0.0007***
E4			WW	0.0001***	0.1644ns	0.0347*	0.0008***
E5	2016	顺义 SY	DS+HT	0.0433*	0.0001***	0.0086**	0.0006***
E6			DS	0.0007***	0.0019**	0.0044**	0.0013**
E7			WW+HT	0.0624ns	0.0075**	0.0547ns	0.0037**
E8			WW	0.5628ns	0.0274*	0.2390ns	0.0179*
E9		昌平 CP	WW	0.0003***	0.0088**	0.0053**	0.0149*
E10		昌平 CP	DS	0.0010**	0.1856ns	0.0260*	0.1182ns
E11	2017	顺义 SY	DS+HT	0.3325ns	0.0035**	0.0011**	0.0032**
E12			DS	0.0010**	0.0055**	0.1369ns	0.0069**
E13			WW+HT	0.1806ns	0.0054**	0.0025**	0.0133*
E14			WW	0.0001***	0.0549ns	0.0040**	0.1279ns
E15		昌平 CP	DS	0.0017**	0.0169*	0.0008***	0.0020**
E16		昌平 CP	WW	0.0001***	0.0264*	0.0104*	0.0072**

小麦转录因子基因TaWRKY65-3A功能解析和分子标记开发

Functional Characterization and Molecular Marker Development of the Wheat Transcription Factor Gene TaWRKY65-3A

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