小麦耐低温关键调控途径解析及<i>TaGGCT18-6A</i>基因功能验证

doi:10.7668/hbnxb.20195902

摘要/Abstract

摘要： 为解析小麦低温胁迫应答机制并挖掘优异抗寒基因资源,通过转录组测序揭示小麦低温响应的关键调控网络,并对候选基因TaGGCT18-6A进行功能验证。结果表明,4 ℃冷处理诱导小麦幼苗分别产生10 893个(6 h处理)和18 784个(24 h处理)差异表达基因,KEGG富集分析显示,差异基因显著富集于MAPK信号转导及谷胱甘肽代谢等通路。基于谷胱甘肽代谢途径关键基因筛选,克隆获得γ-谷氨酰环转移酶基因TaGGCT18-6A。该基因全长1 772 bp,编码218个氨基酸,其编码蛋白具有典型的GGCT-like超家族结构域及核心结构域ChaC。启动子顺式作用元件分析表明,该基因含低温响应元件(LTR)及脱水响应元件(DRE)等胁迫相关元件;表达模式分析发现,TaGGCT18-6A在4 ℃冷处理下呈持续上调趋势。转基因水稻功能验证表明,过表达株系在-4 ℃低温冻害胁迫下相较野生型对照,OE#1、OE#2和OE#3存活率与生物量显著提高,OE#1和OE#2的株高显著提高,OE#1、OE#2和OE#3相对电导率显著降低。经4 ℃冷处理后,过表达株系渗透调节物质(脯氨酸、可溶性糖)积累量较对照显著增加,丙二醛含量降低;SOD、POD和CAT活性显著上升。以上研究表明,TaGGCT18-6A通过调控谷胱甘肽代谢增强抗氧化能力,进而提高植物低温耐受性,为小麦抗寒分子育种提供理论依据和优异基因资源。

关键词: 小麦, 低温胁迫, 转录组分析, γ-谷氨酰环转移酶, TaGGCT18-6A

Abstract:

To elucidate the mechanisms of cold response and explore superior cold-tolerance gene resources in wheat,this study employed transcriptome sequencing to uncover key regulatory networks underlying wheat cold response,and performed functional validation of the candidate gene TaGGCT18-6A.The results demonstrated that 4 ℃ cold treatments induced 10 893 and 18 784 differentially expressed genes(DEGs)in wheat seedlings after 6 h and 24 h cold treatment,respectively.KEGG analysis revealed significant enrichment of DEGs in pathways including MAPK signaling transduction and glutathione metabolism.A γ-glutamyl cyclotransferase gene TaGGCT18-6A was cloned through screening key genes in glutathione metabolism.This gene had a length of 1 772 bp,encoding 218 amino acids with conserved GGCT-like superfamily and ChaC core domain.Promoter cis-acting element analysis identified stress-responsive elements such as low-temperature-responsive(LTR)and dehydration-responsive (DRE)elements; consistently,expression pattern analysis showed sustained upregulation of TaGGCT18-6A under 4 ℃ cold treatments. Functional validation in transgenic rice revealed that overexpression lines OE#1, OE#2 and OE#3 exhibited significantly enhanced survival rate,plant height,and biomass. Overexpression lines OE#1 and OE#2 exhibited significantly enhanced plant height, while overexpression lines OE#1, OE#2 and OE#3 exhibited significantly reduced relative electrolyte leakage under -4 ℃ freezing treatments.After 4 ℃ treatment,overexpression lines accumulated higher levels of osmolytes(proline and soluble sugars),decreased malondialdehyde(MDA)content,and increased activities of superoxide dismutase(SOD),peroxidase(POD),and catalase(CAT).These findings collectively demonstrated that TaGGCT18-6A enhanced plant cold tolerance by regulating glutathione metabolism to improve antioxidant capacity.This study will provide a theoretical foundation and valuable genetic resources for molecular breeding of cold-tolerant wheat.

Key words: Wheat, Low temperature stress, Transcriptome analysis, γ-glutamyl cyclotransferase, TaGGCT18-6A

中图分类号:

成春华, 陈涛, 张龙, 郭利建, 车卓, 马靖福, 杨德龙. 小麦耐低温关键调控途径解析及TaGGCT18-6A基因功能验证[J]. 华北农学报, 2025, 40(5): 1-11. doi: 10.7668/hbnxb.20195902.

CHENG Chunhua, CHEN Tao, ZHANG Long, GUO Lijian, CHE Zhuo, MA Jingfu, YANG Delong. Dissection of Key Regulatory Pathways for Low-temperature Tolerance and Functional Verification of TaGGCT18-6A Gene in Wheat[J]. Acta Agriculturae Boreali-Sinica, 2025, 40(5): 1-11. doi: 10.7668/hbnxb.20195902.

http://www.hbnxb.net/CN/Y2025/V40/I5/1

图/表 8

表1 本研究所用的引物

Tab.1 Primers used in this study

引物名称 Primer name	引物序列(5'—3') Primer sequence(5'—3')	用途 Purpose
TaGGCT18-6A-F	TCAGCAGGATGGTGCTGTG	TaGGCT18-6A基因编码区PCR
TaGGCT18-6A-R	TCAATGGGAGAATGTCAAGG
TaGGCT18-6A-Os-F	ACTATCCTTCGCAAGACCCTT	TaGGCT18-6A转基因水稻PCR
TaGGCT18-6A-Os-R	GGAGTGGATACGAAAACTAATACAT
TaGGCT18-6A-qRT-PCR-F	CGTTCACCGGTGGTCIATTT	TaGGCT18-6A荧光定量PCR
TaGGCT18-6A-qRT-PCR-R	GGCTCCACTGTTCCTTGAGA
TaMAPKKK6-1A -qRT-PCR-F	GATGGTGCACGGGGACGTC	TaMAPKKK6-1A荧光定量PCR
TaMAPKKK6-1A-qRT-PCR-R	ACAACCATGCAGCCGAGCG
TaMAPKK7-4A-qRT-PCR-F	CTTCGCGCTCAAGCTCTTC	TaMAPKK7-4A荧光定量PCR
TaMAPKK7-4A-qRT-PCR-R	ACACGAGCTCCAGCGCCAG
TaMAPK5-4A-qRT-PCR-F	CAACGCCTTCGACAACAACAT	TaMAPK5-4A荧光定量PCR
TaMAPK5-4A-qRT-PCR-R	GTCCGTGTCCATGAGCTCAG
TaGGT-2D-qRT-PCR-F	CTGTGACCTCGCCGCACG	TaGGT-2D荧光定量PCR
TaGGT-2D-qRT-PCR-R	GTGAGCCTGACGAGCATGAA
TaGST6-5A-qRT-PCR-F	CTCGGCGGCAAGGATTTCT	TaGST6-5A荧光定量PCR
TaGST6-5A-qRT-PCR-R	CCTGGGTTTCAGCCTCTCC
TaActin-F	GTGTCGCACCAGAGGATCAT	小麦内参基因荧光定量PCR
TaActin-R	CGCTGGCATACAAGGACAGA

表2 转录组数据统计

Tab.2 Statistics of all the samples of sequencing data

样品 Sample	原始序列数 Raw reads	过滤后序列数 Clean reads	过滤后碱基数/Gb Clean bases	Q30/%	GC含量/% GC content	比对率/% Total mapped ratio
CK1	55 064 908	54 452 354	8.17	94.12	55.78	90.83
CK2	54 300 614	53 691 934	8.05	94.15	56.15	90.99
CK3	55 233 878	54 503 334	8.18	93.91	58.20	89.95
Cold-6 h-1	57 758 566	56 960 080	8.54	94.29	56.38	91.17
Cold-6 h-2	55 003 544	53 988 202	8.10	94.33	56.27	91.64
Cold-6 h-3	55 752 342	54 775 938	8.22	94.20	55.29	89.52
Cold-24 h-1	55 233 518	54 487 646	8.17	93.73	57.33	89.28
Cold-24 h-2	54 561 382	53 976 740	8.10	94.05	56.44	90.17
Cold-24 h-3	53 870 550	53 342 274	8.00	93.77	55.47	90.68

图1 小麦低温处理后的差异表达基因及KEGG分析 A.Cold-6 h与对照的火山图;B.Cold-24 h与对照的火山图;C.Cold-6 h和对照、Cold-24 h和对照比较组之间DEGs的韦恩图;D.6 h和24 h DEGs的KEGG富集分析。

Fig.1 Analysis of differentially expressed genes and KEGG in wheat after low-temperature treatment A.Volcano plot of Cold-6 h vs CK;B.Volcano plot of Cold-24 h vs CK;C.Venn diagram showed the number of DEGs between Cold-6 h vs CK and Cold-24 h vs CK;D.KEGG analysis of DEGs under Cold-6 h vs CK and Cold-24 h vs CK.

图2 MAPK信号通路和谷胱甘肽代谢通路差异表达基因热图 A.MAPK信号通路;B.谷胱甘肽代谢通路。

Fig.2 Heatmap of differentially expressed genes in MAPK signaling pathway and glutathione metabolic pathways A.MAPK signaling pathway;B.Glutathione metabolism pathway.

图3 MAPK信号通路和谷胱甘肽代谢通路差异表达基因的qRT-PCR验证以未做冷处理(0 h)时各个基因的表达量为对照。误差线代表平均值±标准差;不同小写字母表示差异显著(P<0.05,n=3)。图4—6同。

Fig.3 qRT-PCR verification of DEGs in MAPK signaling pathway-plant and glutathione metabolism pathway The expression levels of various genes with untreated group were used as a control.Error bar represents mean±s;different lowercase letters indicate significant differences(P<0.05,n=3).The same as Fig.4—6.

图4 TaGGCT18-6A的序列特征和表达模式分析 A.TaGGCT18-6A基因CDS区的PCR扩增;B.小麦TaGGCT18-6A基因结构分析;C.小麦TaGGCT18-6A蛋白保守基序分析;D.小麦TaGGCT18-6A启动子顺式作用元件分析;E.低温胁迫下TaGGCT18-6A表达模式分析(以未做低温处理时的各个基因的表达量为对照)。

Fig.4 Analysis of sequence characteristics and expression patterns of TaGGCT18-6A A.PCR amplification of TaGGCT18-6A CDS sequence;B.Structural analysis of TaGGCT18-6A gene in wheat;C.Conservative motif analysis of wheat TaGGCT18-6A protein;D.Analysis of cis-acting elements in wheat TaGGCT18-6A promoter;E.Analysis of TaGGCT18-6A expression pattern under low temperature stress(The expression levels of various genes with untreated group were used as a control).

图5 TaGGCT18-6A转基因水稻株系的表型统计 A.TaGGCT18-6A过表达株系的低温处理表型。

Fig.5 Phenotype statistics in the overexpression lines of TaGGCT18-6A A.The phenotype of TaGGCT18-6A overexpression lines under cold treatment.

图6 TaGGCT18-6A转基因水稻株系的生理指标分析

Fig.6 Physiological indices in the overexpression lines of TaGGCT18-6A

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小麦耐低温关键调控途径解析及TaGGCT18-6A基因功能验证

Dissection of Key Regulatory Pathways for Low-temperature Tolerance and Functional Verification of TaGGCT18-6A Gene in Wheat

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小麦耐低温关键调控途径解析及TaGGCT18-6A基因功能验证

Dissection of Key Regulatory Pathways for Low-temperature Tolerance and Functional Verification of TaGGCT18-6A Gene in Wheat

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