山葡萄VaMYB4a-VaPAL模块响应低温胁迫应答的功能研究

doi:10.16420/j.issn.0513-353x.2025-0877

园艺学报 ›› 2026, Vol. 53 ›› Issue (9): 2531-2544.doi: 10.16420/j.issn.0513-353x.2025-0877

• 遗传育种·种质资源·分子生物学 • 下一篇

山葡萄VaMYB4a-VaPAL模块响应低温胁迫应答的功能研究

李佳欣¹, 张震¹^,²^,³, 俞沁含⁵, 吴文康¹, 张宁波¹^,²^,³, 徐伟荣¹^,²^,³^,⁴^,⁵^,^*()

¹ 宁夏大学葡萄酒与园艺学院，银川 750021
² 葡萄与葡萄酒教育部工程研究中心，银川 750021
³ 宁夏优势特色作物现代分子育种重点实验室，银川 750021
⁴ 林木资源高效生产全国重点实验室，银川 750021
⁵ 宁夏大学生命科学学院，银川 750021

收稿日期:2026-03-17 修回日期:2026-05-21 出版日期:2026-09-25 发布日期:2026-09-20
通讯作者:
*E-mail：xuwr@nxu.edu.cn
基金资助:
国家自然科学基金项目(32472711); 宁夏回族自治区科技领军人才培养项目(2025GKLRLX21); 宁夏自然科学基金项目(2024AAC02024); 宁夏回族自治区重点研发计划项目(2025BBF02019)

Functional Study of the VaMYB4a-VaPAL Module in Response to Cold Stress in Vitis amurensis

LI Jiaxin¹, ZHANG Zhen¹^,²^,³, YU Qinhan⁵, WU Wenkang¹, ZHANG Ningbo¹^,²^,³, XU Weirong¹^,²^,³^,⁴^,⁵^,^*()

¹ School of Enology & Horticulture， Ningxia University， Yinchuan 750021， China
² Engineering Research Center of Grape and Wine， Ministry of Education，Ningxia University， Yinchuan 750021， China
³ Key Laboratory of Modern Molecular Breeding for Dominant and Special Crops in Ningxia， Yinchuan 750021， China
⁴ State Key Laboratory of Efficient Production of Forest Resources， Yinchuan 750021， China
⁵ School of Life Science， Ningxia University， Yinchuan 750021， China

Received:2026-03-17 Revised:2026-05-21 Published:2026-09-25 Online:2026-09-20
Contact:
*E-mail：xuwr@nxu.edu.cn

摘要/Abstract

摘要：

低温胁迫是限制葡萄（Vitis spp.）生长发育和产量的重要非生物因子，其感应信号与代谢调控整合机制尚不清晰。以山葡萄转录因子VaMYB4a为研究对象，通过构建VaMYB4a的过表达与CRISPR/Cas9敲除材料并结合生理指标分析表明，VaMYB4a可显著增强光系统Ⅱ的稳定性，降低膜脂过氧化和活性氧积累，提升植株抗寒能力。qRT-PCR检测表明，VaMYB4a激活CBF-COR通路，显著诱导CBF1-CBF4及下游RD29A的表达。酵母单杂交、双荧光素酶报告和电泳迁移率分析表明，VaMYB4a可直接结合VaPAL启动子中的CAACCA顺式作用元件，增强其转录活性，揭示了VaMYB4a可协同调控苯丙烷代谢通路的潜在分子机制。VaPAL过表达显著缓解低温胁迫下的氧化损伤，增强愈伤组织的低温耐受性。启动子活性分析显示，VaPAL是苯丙烷代谢通路中对低温最敏感的关键响应基因。共表达分析揭示，VaPAL可反馈调节CBF通路的响应强度，在信号转导与代谢调节间发挥桥梁作用。VaMYB4a-VaPAL模块在葡萄响应低温胁迫应答中的“信号—代谢”协同调控中发挥重要作用，可为果树作物抗寒性的遗传改良提供参考依据与基因资源。

关键词: 葡萄, 低温胁迫, VaMYB4a, PAL, 抗寒性

Abstract:

Low-temperature stress is a major abiotic factor limiting the growth and productivity of grapevine（Vitis spp.），yet the integrated mechanisms underlying the integration of cold signaling and metabolic regulation remain poorly understood. Here，a wild grapevine（Vitis amurensis）MYB transcriptionfactor（VaMYB4a）were characterized. Through the generation of VaMYB4a overexpression and CRISPR/Cas9 knockout lines，combined with physiological analyses，we found that VaMYB4a significantly enhanced the stability of photosystem Ⅱ，reduced membrane lipid peroxidation and reactive oxygen species accumulation，and thereby improved plant cold tolerance. qRT-PCR analysis showed that VaMYB4a activated the CBF-COR pathway and significantly induced the expression of CBF1-CBF4 and the downstream gene RD29A. Yeast one-hybrid，dual-luciferase reporter，and electrophoretic mobility shift assays demonstrated that VaMYB4a directly binds to the CAACCA cis-element in the VaPAL promoter and enhances its transcriptional activity，revealing a potential molecular mechanism by which VaMYB4a coordinately regulates the phenylpropanoid metabolic pathway. Overexpression of VaPAL markedly alleviated oxidative damage under low-temperature stress and enhanced cold tolerance in callus tissues. Promoter activity analysis further indicated that VaPAL is a key low-temperature-responsive gene with the highest sensitivity within the phenylpropanoid pathway. Co-expression analysis revealed that VaPAL can feedback-regulate the response intensity of the CBF pathway，thereby acting as a bridge between signal transduction and metabolic regulation. Collectively，the VaMYB4a-VaPAL module plays an important role in the coordinated“signal-metabolism”regulation of grapevine responses to low-temperature stress，providing valuable gene resources and a theoretical basis for the genetic improvement of cold tolerance in fruit crops.

Key words: grapevine, low-temperature stress, VaMYB4a, PAL, cold tolerance

中图分类号:

S663.1

李佳欣, 张震, 俞沁含, 吴文康, 张宁波, 徐伟荣. 山葡萄VaMYB4a-VaPAL模块响应低温胁迫应答的功能研究[J]. 园艺学报, 2026, 53(9): 2531-2544.

LI Jiaxin, ZHANG Zhen, YU Qinhan, WU Wenkang, ZHANG Ningbo, XU Weirong. Functional Study of the VaMYB4a-VaPAL Module in Response to Cold Stress in Vitis amurensis[J]. Acta Horticulturae Sinica, 2026, 53(9): 2531-2544.

图/表 8

表1 本研究中互作和qRT-PCR的特异性引物

Table 1 Specific primers for interactions and qRT-PCR

用途 Usage	引物名称 Primer name	引物序列（5′-3′） Primer sequence
双荧光素酶报告实验 Dual-luciferase reporter assay	0800-PAL(P1)-F	ACTCACTATAGGGCGAATTGGGTACCGCGATTCCAATTTTTTCAAAACT
双荧光素酶报告实验 Dual-luciferase reporter assay	0800-PAL(P1)-R	ATATCAAGCTTATCGATACCGTCGACCTCCAAGTGTTTGATCAAATGCC
酵母单杂交 Yeast one-hybrid	pHIS2-PAL(P1)-F	gactcactatagggcgaattcGCGATTCCAATTTTTTCAAAACT
	pHIS2-PAL(P1)-R	gattcgcgaacgcgtgagctcCTCCAAGTGTTTGATCAAATGCC
	pHIS2-PAL(P2)-F	gactcactatagggcgaattcGCAACACTCCACGTCGAAGC
	pHIS2-PAL(P2)-R	gattcgcgaacgcgtgagctcCGGAAACCTAGTTTCAAAAATTAATATG
电泳迁移率实验 Electrophoretic mobility shift assay	marker(PAL-P1)-F	atagaagcctctttaaacaaccaaaaacttataaaaaaaa
	marker(PAL-P1)-R	TTTTTTTTATAAGTTTTTGGTTGTTTAAAGAGGCTTCTAT
	competitor(PAL-P1)-F	atagaagcctctttaaacaaccaaaaacttataaaaaaaa
	competitor(PAL-P1)-R	TTTTTTTTATAAGTTTTTGGTTGTTTAAAGAGGCTTCTAT
	mutation(PAL-P1)-F	atagaagcctctttaaaccccccaaaacttataaaaaaaa
	mutation(PAL-P1)-R	TTTTTTTTATAAGTTTTGGGGGGTTTAAAGAGGCTTCTAT
实时荧光定量PCR qRT-PCR	4CL-RT-F	GATCCTCCTCCAGAACTGCG
	4CL-RT-R	AAGTGTAGAAGGGGTTGGCG
	C4H-RT-F	GAACCACCTGAACCTCTCCG
	C4H-RT-R	ATCCGAACTCCACTCCCTGA
	PAL-RT-F	GACTGGTGAAACGGTGAGGT
	PAL-RT-R	ACCGTTCCAAGCACTGAGAC
	VvCBF1-RT-F	ACGAGGAGGCAATGTTCAATA
	VvCBF1-RT-R	TCATCCCAGCTGAATCCTTTAC
	VvCBF2-RT-F	GAGAAAGCCGGAGATGGTTT
	VvCBF2-RT-R	CACACATAGCAGGTGGAGTAAG
	VvCBF3-RT-F	ATGGATGAGGAGGCATTGTTTA
	VvCBF3-RT-R	AAGTCCGTGTAGGAAGTTGTATC
	VvCBF4-RT-F	ACCCTCACCCGCTCGTATG
	VvCBF4-RT-R	CCGCGTCTCCCGAAACTT
	VvRD29A-RT-F	GGTGATGGTGGTGAAGAAGTGGTA
	VvRD29A-RT-R	AACGGTTGTTCTGGCTTATCCTGTA
内参基因 Reference gene	VvActin1-F	CTATCCTTCGTCTTGACCTTGCTG
内参基因 Reference gene	VvActin1-R	AGTGGTGAACATGTAACCCCTCTC
RUBY报告基因分析启动子活性 RUBY reporter gene analysis of promoter activity	PAL-RUBY-F	attaccgcctttgagtgagctctagaGAGCTAAGGCAACAAAGGTAAAAAT
	PAL-RUBY-R	gcgagggtcgcatgatccatggtaccATCTTAGAATCCCAACAAAGAAACAA
	4CL-RUBY-F	attaccgcctttgagtgagctctagaGAACCCAACCTGTCCAACTTATACT
	4CL-RUBY-R	gcgagggtcgcatgatccatggtaccGGTGGCACAGAGAGAGAGAGAGA
	C4H-RUBY-F	attaccgcctttgagtgagctctagaGAGTGAGTCAGAGTTAGATTATTGTCCG
	C4H-RUBY-R	gcgagggtcgcatgatccatggtaccGGTTGTGGCGGTGGATGG

表1 本研究中互作和qRT-PCR的特异性引物

Table 1 Specific primers for interactions and qRT-PCR

用途 Usage	引物名称 Primer name	引物序列（5′-3′） Primer sequence
双荧光素酶报告实验 Dual-luciferase reporter assay	0800-PAL(P1)-F	ACTCACTATAGGGCGAATTGGGTACCGCGATTCCAATTTTTTCAAAACT
双荧光素酶报告实验 Dual-luciferase reporter assay	0800-PAL(P1)-R	ATATCAAGCTTATCGATACCGTCGACCTCCAAGTGTTTGATCAAATGCC
酵母单杂交 Yeast one-hybrid	pHIS2-PAL(P1)-F	gactcactatagggcgaattcGCGATTCCAATTTTTTCAAAACT
	pHIS2-PAL(P1)-R	gattcgcgaacgcgtgagctcCTCCAAGTGTTTGATCAAATGCC
	pHIS2-PAL(P2)-F	gactcactatagggcgaattcGCAACACTCCACGTCGAAGC
	pHIS2-PAL(P2)-R	gattcgcgaacgcgtgagctcCGGAAACCTAGTTTCAAAAATTAATATG
电泳迁移率实验 Electrophoretic mobility shift assay	marker(PAL-P1)-F	atagaagcctctttaaacaaccaaaaacttataaaaaaaa
	marker(PAL-P1)-R	TTTTTTTTATAAGTTTTTGGTTGTTTAAAGAGGCTTCTAT
	competitor(PAL-P1)-F	atagaagcctctttaaacaaccaaaaacttataaaaaaaa
	competitor(PAL-P1)-R	TTTTTTTTATAAGTTTTTGGTTGTTTAAAGAGGCTTCTAT
	mutation(PAL-P1)-F	atagaagcctctttaaaccccccaaaacttataaaaaaaa
	mutation(PAL-P1)-R	TTTTTTTTATAAGTTTTGGGGGGTTTAAAGAGGCTTCTAT
实时荧光定量PCR qRT-PCR	4CL-RT-F	GATCCTCCTCCAGAACTGCG
	4CL-RT-R	AAGTGTAGAAGGGGTTGGCG
	C4H-RT-F	GAACCACCTGAACCTCTCCG
	C4H-RT-R	ATCCGAACTCCACTCCCTGA
	PAL-RT-F	GACTGGTGAAACGGTGAGGT
	PAL-RT-R	ACCGTTCCAAGCACTGAGAC
	VvCBF1-RT-F	ACGAGGAGGCAATGTTCAATA
	VvCBF1-RT-R	TCATCCCAGCTGAATCCTTTAC
	VvCBF2-RT-F	GAGAAAGCCGGAGATGGTTT
	VvCBF2-RT-R	CACACATAGCAGGTGGAGTAAG
	VvCBF3-RT-F	ATGGATGAGGAGGCATTGTTTA
	VvCBF3-RT-R	AAGTCCGTGTAGGAAGTTGTATC
	VvCBF4-RT-F	ACCCTCACCCGCTCGTATG
	VvCBF4-RT-R	CCGCGTCTCCCGAAACTT
	VvRD29A-RT-F	GGTGATGGTGGTGAAGAAGTGGTA
	VvRD29A-RT-R	AACGGTTGTTCTGGCTTATCCTGTA
内参基因 Reference gene	VvActin1-F	CTATCCTTCGTCTTGACCTTGCTG
内参基因 Reference gene	VvActin1-R	AGTGGTGAACATGTAACCCCTCTC
RUBY报告基因分析启动子活性 RUBY reporter gene analysis of promoter activity	PAL-RUBY-F	attaccgcctttgagtgagctctagaGAGCTAAGGCAACAAAGGTAAAAAT
	PAL-RUBY-R	gcgagggtcgcatgatccatggtaccATCTTAGAATCCCAACAAAGAAACAA
	4CL-RUBY-F	attaccgcctttgagtgagctctagaGAACCCAACCTGTCCAACTTATACT
	4CL-RUBY-R	gcgagggtcgcatgatccatggtaccGGTGGCACAGAGAGAGAGAGAGA
	C4H-RUBY-F	attaccgcctttgagtgagctctagaGAGTGAGTCAGAGTTAGATTATTGTCCG
	C4H-RUBY-R	gcgagggtcgcatgatccatggtaccGGTTGTGGCGGTGGATGG

图1 VaMYB4a转基因盆栽苗的冻害表型（A）、叶绿素荧光成像（B）和最大量子效率（C） UT：未转化的‘赤霞珠’对照。差异显著性采用单因素方差分析，不同小写字母代表在P < 0.05水平差异显著。下同

Fig. 1 Frost damage phenotype（A），chlorophyll fluorescence imaging（B）and maximum quantum efficiency（C） of VaMYB4a transgenic potted seedlings UT：Untransformed‘Cabernet Sauvignon’control. Significant differences were analyzed by One-way ANOVA. Different lowercase letters indicate significant difference at P < 0.05. The same below

图2 VaMYB4a介导CBF-COR低温响应通路相关基因表达水平

Fig. 2 Expression level of VaMYB4a related to the CBF-COR cold pathway mediated

图3 VaMYB4a调控VaPAL、Va4CL和VaC4H基因表达响应低温胁迫 A ~ C：低温胁迫山葡萄‘左山-1’；D ~ F：低温胁迫基因编辑株系和过表达株系。采用Student’s t-test 进行差异显著性分析，*和**表示与对照相比存在显著差异和极显著差异，* P < 0.05，** P < 0.01，下同

Fig. 3 VaMYB4a regulates VaPAL，Va4CL，and VaC4H expression in response to cold stress A-C：‘Zuoshan-1’subjected to cold stress；D-F：Gene-edited and overexpression lines subjected to cold stress. Student’s t-test was used to analyse statistical significance. * and ** indicate the significant differences level and extremely significant differences level compared to the control，* P < 0.05，** P < 0.01. The same below

图4 RUBY报告基因检测VaPAL启动子表型分析（A）和活性检测（B）

Fig. 4 Phenotypic analysis（A）and activity assay（B）of VaPAL promoter using RUBY reporter gene assay

图5 VaMYB4a正向调控VaPAL基因表达 A：VaPAL启动子结构图（P1/P2片段含MYB结合元件）；B：酵母单杂交验证；C：双荧光素酶报告系统载体构建示意图；D：双荧光素酶活性检测；E：电泳迁移率试验（“+”表示添加相应试剂，“-”表示未添加）

Fig. 5 VaMYB4a positively regulates the expression of VaPAL A：Promoter map of VaPAL（P1/P2 represents the fragment containing the MYB binding element）；B：Y1H assay；C：Schematic diagram of dual-luciferase reporter vector and effector vector；D：Dual-luciferase assay；E：EMSA（“+”or“-”indicates the presence or absence of reagents during protein electrophoresis）

图6 VaPAL愈伤组织抗寒性鉴定

Fig. 6 Identification of cold resistance in VaPAL callus

图7 VaMYB4a与VaPAL共表达影响CBF-COR通路

Fig. 7 Co-expression of VaMYB4a and VaPAL affect the CBF-COR pathway

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山葡萄VaMYB4a-VaPAL模块响应低温胁迫应答的功能研究

Functional Study of the VaMYB4a-VaPAL Module in Response to Cold Stress in Vitis amurensis

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山葡萄VaMYB4a-VaPAL模块响应低温胁迫应答的功能研究

Functional Study of the VaMYB4a-VaPAL Module in Response to Cold Stress in Vitis amurensis

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