葡萄DGK基因家族鉴定及VvDGK1在盐胁迫下的功能研究

doi:10.16420/j.issn.0513-353x.2025-0766

摘要/Abstract

摘要：

甘油二酯激酶（DGK）是逆境胁迫下催化合成磷脂酸（PA）的关键信号酶，在细胞信号转导过程中发挥重要的调控作用。为系统分析葡萄中DGK家族基因成员并开展生物学功能研究，本研究中基于葡萄基因组数据库，查找和鉴定了葡萄DGK基因家族成员，并进行了系统进化、理化性质、基因结构、蛋白和启动子序列、基因表达水平等分析。将VvDGK1在拟南芥中进行遗传转化，并进行盐胁迫处理，对VvDGK1的互作蛋白进行筛选和鉴定。结果显示，共鉴定到5个DGK基因家族成员，其中VvDGK1和VvDGK2属于亚族Ⅰ，VvDGK3属于亚族Ⅱ，VvDGK4和VvDGK5属于亚族Ⅲ。系统进化和共线性分析发现，葡萄中5个VvDGK基因与苹果MdDGK基因同源性较高，VvDGK1与MdDGK3、MdDGK7和AtDGK1之间的共线性关系最强。VvDGK基因启动子区存在大量与生长发育、植物激素和逆境胁迫响应相关的顺式作用元件。VvDGK在茎和叶中表达量较高，在根、花和果实中表达量较低，呈现明显的组织特异性；在干旱、盐和脱落酸（ABA）等胁迫处理下呈现不同的表达模式，其中VvDGK1在不同胁迫处理下均被显著诱导。将VvDGK1在拟南芥中过表达后发现，与野生型相比，转化株系增强了对盐胁迫的抗性，减缓了胁迫下叶片光系统Ⅱ最大光化学效率（F_v/F_m）的下降和叶绿素的降解，降低了胁迫下活性氧的积累，增强了抗氧化酶（如CAT、POD和APX）的活性。酵母双杂交筛选到VvFd和VvRCA，能够与VvDGK1发生蛋白互作，双分子荧光互补（BiFC）实验进一步验证了VvDGK1和VvFd的体内互作关系，推测VvDGK1通过与VvFd和VvRCA蛋白互作影响光合作用，参与盐胁迫下的信号转导。

关键词: 葡萄, 甘油二酯激酶, VvDGK1, 盐胁迫, 蛋白质互作

Abstract:

Diacylglycerol kinase（DGK）is a key signaling enzyme that induces the synthesis of phosphatidic acid（PA）under adverse stress and plays an important regulatory role in the process of cellular signal transduction. To systematically analyze the DGK family gene members in grapes and conduct biological function research，based on the grape genome database，searched for and identified the DGK gene family members in grapes，and conducted analyses on their phylogenetic evolution，physicochemical properties，gene structure，protein and promoter sequences，and gene expression levels. In addition，VvDGK1 was genetically transformed into Arabidopsis thaliana and subjected to salt stress treatment，and the interacting proteins of VvDGK1 were screened and identified. The results showed that a total of five DGK family members were identified in the grape genome database. Among them，VvDGK1 and VvDGK2 belong to ClusterⅠ，VvDGK3 to ClusterⅡ，and VvDGK4 and VvDGK5 to Cluster Ⅲ. Systematic evolution and collinearity analysis revealed that the five VvDGKs in grapes have a relatively high homology with the DGK genes in apples，and the collinearity relationship between VvDGK1 and MdDGK3，MdDGK7，and AtDGK1 is the strongest. The promoter region of the VvDGKs gene contains a large number of cis-acting elements related to growth and development，plant hormones，and responses to adverse stress. The VvDGK gene has a relatively high expression level in stems and leaves，but a relatively low expression level in roots，flowers and fruits，showing obvious tissue specificity. The VvDGK gene shows different expression patterns under stress treatments such as drought，salt，and abscisic acid. Among them，VvDGK1 gene expression was significantly induced under different stress treatments. Overexpression of VvDGK1 in Arabidopsis revealed that the transformed strain enhanced resistance to salt stress，slowed the decline of the maximum photochemical efficiency of leaf photosystemⅡ（F_v/F_m）and the degradation of chlorophyll under stress，reduced the accumulation of reactive oxygen species under stress，and increased the activity of antioxidant enzymes（such as CAT，POD and APX）. The yeast two-hybrid system screened out VvFd and VvRCA，which can interact with VvDGK1. The bimolecular fluorescent complimentary（BiFC）further verified the in vivo interaction between VvDGK1 and VvFd. It is speculated that VvDGK1 participates in signal transduction under salt stress by influencing photosynthesis through interaction with VvFd and VvRCA.

Key words: grapevine, diacylglycerol kinase, VvDGK1, salt stress, protein-protein interaction

谭延肖, 韩双, 冷凇凝, 陈晓乐, 刘佰霖, 史红梅, 何维华, 张煜. 葡萄DGK基因家族鉴定及VvDGK1在盐胁迫下的功能研究[J]. 园艺学报, 2026, 53(7): 2034-2052.

TAN Yanxiao, HAN Shuang, LENG Songning, CHEN Xiaole, LIU Bailin, SHI Hongmei, HE Weihua, ZHANG Yu. Identification of the DGK Family Genes in Vitis vinifera and Functional Study of VvDGK1 Under Salt Stress[J]. Acta Horticulturae Sinica, 2026, 53(7): 2034-2052.

图/表 14

表1 本研究中所用的引物

Table 1 Primers used in the experiment

用途 Purpose	名称 Name	引物序列（5′-3′） Primer sequence
qRT-PCR	VvDGK1-F	ATTCTGATATGGGGGCCAAA
	VvDGK1-R	GCCAAGCTTTGGTTCCAA
	VvDGK2-F	TAACATTCCTCTTGACCGT
	VvDGK2-R	TCCATCACCAACCAGCAGT
	VvDGK3-F	ATGAAGCCGAGGGAAAATGA
	VvDGK3-R	TTCCCATACTGACCATGCAT
	VvDGK4-F	GCAACAAAGCTCCAATCGC
	VvDGK4-R	AACTTTACAAAAATCAGCAAG
	VvDGK5-F	AACAGGGTCACGTCGCAGAA
	VvDGK5-R	AACTAAAGCCTTTCCGATG
	VvActin1-F	GATTCTGGTGATGGTGTGAGT
	VvActin1-R	GACAATTTCCCGTTCAGCAGT
基因扩增 Gene amplification	VvDGK1-F	AATGGATGATGATGGACTTTGCAACGAGATCTTTATG
基因扩增 Gene amplification	VvDGK1-R	CGCTTCTAGCGCCATTTCTTCTACATGTGCTGCAGAAG
遗传转化载体构建 Construction of genetic transformation vector	VvDGK1-G-F	GCTAGTCTAGAATGGATGATGATGGACTTTGC
遗传转化载体构建 Construction of genetic transformation vector	VvDGK1-G-R	ATAGAGCTCCTAGCGCCATTTCTTCTA
酵母双杂交筛库 Yeast two-hybrid screening	pGBKT7-VvDGK1-F	AGGGCCAGAATTCCTGCTTAAGGGATGATGATATGGACTTATGTTGCCTTGA
	pGBKT7-VvDGK1-R	TAAGGCCATGGGGCCTATGTCTTACGCAGCTCATTCAGCGCTTCTTG
	pGADT7-T-F	GGAGTACCCATACGACGTACC
	pGADT7-T-R	TATCTACGATTCATCTGCAGC
酵母双杂交验证 Yeast two-hybrid verification	pGADT7-VvRCA-F	TCAACGCAGAGTGGCCATTAATGAGCTCAAACCCTAGCATAAAAAGAT
	pGADT7-VvRCA-R	GAATTCTCGAGAGGCCGAGGTTAAGTCATTTGAAGGTTGAGGCTATTGAG
	pGADT7-VvFd-F	CCGGGGCATCGATACGTGGGCATGACTAAGAACCAAGAAACTCTTGT
	pGADT7-VvFd-R	CCTGAGCTCGACGATGGATCGCTCATACGGGACTGCACTCTC
BiFC验证 Bimolecular fluorescent complimentary	pSm35s-VvDGK1-cYFP-F	ATTACAGGTACCCGGGGATCATGGATGGACTTATGTTGCCT
	pSm35s-VvDGK1-cYFP-R	GCCACCGCCGTCGACTCTAGTCTTCTACATGTGCTGCAGAAG
	pSm35s-VvFd-nYFP-F	TCAACGCAGAGTGGCCATTAATGACTAAGAACCAAGAAACTCT
	pSm35s-VvFd-nYFP-R	GAATTCTCGAGAGGCCGAGGCGCTCATACGGGACTGCACTCTC

图1 葡萄VvDGK家族基因成员染色体定位

Fig. 1 Chromosomal localization of VvDGK gene family members in Vitis vinifera

表2 葡萄VvDGK家族基因成员基本理化性质

Table 2 Physicochemical properties of VvDGK gene family members in Vitis vinifera

基因 Gene	基因ID Gene ID	染色体 Chromosome number	编码序列/ bp CDS	氨基酸残基数 Amino acidresidue number	相对分子量/ kD Mw	等电点 pI	不稳定指数 Instability index	亲水性 Gravy	跨膜区 Trans- membrane region	亚细胞定位 Subcellular location
VvDGK1	VIT_17s0000g06970	17	2 196	731	81.01	6.28	52.68	-0.26	1	细胞核Nucleus
VvDGK2	VIT_07s0031g02840	7	2 145	714	79.81	8.53	52.57	-0.23	1	细胞核Nucleus
VvDGK3	VIT_11s0052g01840	11	1 539	512	56.85	6.58	50.72	-0.31	0	叶绿体Chloroplast
VvDGK4	VIT_19s0014g03790	19	1 440	479	53.80	6.66	49.28	-0.36	0	叶绿体，细胞质，细胞核 Chloroplast，cytoplasm，nucleus
VvDGK5	VIT_12s0057g00010	12	1 503	500	55.78	5.82	41.93	-0.43	0	叶绿体，细胞质，细胞核 Chloroplast，cytoplasm，nucleus

图2 葡萄VvDGK家族基因保守基序（A）、功能结构域（B）和基因结构（C）分析

Fig. 2 Analysis of the conserved motifs（A），functional domain（B），and gene structure（C）of VvDGK genes in Vitis vinifera

表3 葡萄VvDGK基因8个保守基序信息

Table 3 Eight conserved motif information of VvDGK genes in Vitis vinifera

基序编号Code	长度/aa Length	保守氨基酸序列Conserved amino acid sequence
1	50	KDRBFTPQFVDDGLLEVVGFRDAWHGEKLLVPLGHGTRLAQAHQIRFELH
2	50	FWNYFSIGMDAKVAYAFHTLREEHPEKFYNQLVNKIIYAKLGCKQGWFRT
3	26	PKSIRSIVCLNJPSFPGGLDPWGNPN
4	21	PPIRIIPAGTGNTLAWVLGWG
5	37	PARPVJVFINSKSGGQLGGELRQRLNALLNPVQVFDL
6	50	NKFMDELEEAEEEHPPPHDIPHHSYFLQIRELLETKSJTPSTGMKDCLYD
7	27	AKHVYMRIDGEPWKQPLPIDDDTVVVE
8	29	EFYLPKLRDAEEIKIDSWHFLMRTSIPDC

图3 拟南芥（At）、水稻（Os）、番茄（Sl）、苹果（Md）和葡萄（Vv）DGK家族基因成员的系统进化树

Fig. 3 Phylogenetic tree of DGK gene family members in Arabidopsis thaliana（At），Oryza sativa（Os），Solanum lycopersicum（Sl），Malus × domestica（Md），and Vitis vinifera（Vv）

图4 拟南芥、水稻、番茄、苹果和葡萄DGK基因之间的共线性关系分析绿色：两对基因之间存在 ≤ 50%的共线性；棕色：两对基因之间存在 ≤ 75%的共线性；红色：两对基因之间存在 ≤ 99%的共线性

Fig. 4 Co-linear relationships among DGK genes of Arabidopsis thaliana，Oryza sativa，Solanum lycopersicum，Malus × domestica，and Vitis vinifera Green：Co-linear relationships ≤ 50% between the two pairs of genes；Brown：Co-linear relationships ≤ 75% between the two pairs of genes；Red：Co-linear relationships ≤ 99% between the two pairs of genes

图5 葡萄VvDGK家族基因启动子顺式作用元件分析

Fig. 5 Analysis of cis-acting elements in the promoters of the VvDGK family genes in Vitis vinifera

图6 VvDGK家族基因在葡萄不同组织和不同胁迫处理下的表达量变化内参基因：Actin1；n = 3。不同小写字母表示在不同组织间差异显著（P < 0.05）；以各处理起始时间点作为对照，对照表达量为1。* 表示与对照相比在0.05水平上差异显著

Fig. 6 Analysis of the expression patterns of VvDGK family genes in different tissues of grapes and under different stress treatments Reference gene：Actin1；n = 3. Different lowercase letter indicates significance level（P < 0.05）of each gene at different tissues. Using the starting time point of each treatment as the control，and the expression level of the control was 1. * indicates the significant difference compared with control at the 0.05 level

图7 VvDGK1转基因拟南芥的耐盐性鉴定结果 WT：拟南芥Columbia野生型；#1 ~ #8：VvDGK1转基因株系。*表示相同处理条件下转基因株系与野生型在P < 0.05水平上差异显著；**表示同处理条件下转基因株系与野生型在P < 0.01水平上差异显著；*** 表示同处理条件下转基因株系与野生型在P < 0.001水平上差异显著。下同

Fig. 7 Identification of stress resistance of VvDGK1 transgenic Arabidopsis WT：Wild type of Arabidopsis Columbia. #1-#8：VvDGK1 transgenic lines. * indicates that the transgenic strain is significantly different from the wild-type with the same treatment at 0.05 level. ** indicates that the transgenic strain is significantly different from the wild-type with the same treatment at 0.01 level. *** indicates that the transgenic strain is significantly different from the wild-type with the same treatment at 0.001 level. The same below

图8 盐胁迫下VvDGK1转基因拟南芥H2O2（A）、MDA（B）含量测定及ROS组织化学染色（C，D）

Fig. 8 Determination of H2O2（A）and MDA（B）contents and ROS histochemical staining（C、D）of VvDGK1 transgenic Arabidopsis under salt stress

图9 盐胁迫下VvDGK1转基因拟南芥的抗氧化酶活性

Fig. 9 Antioxidant enzyme activities of VvDGK1 transgenic Arabidopsis under salt stress

图10 酵母双杂交验证与VvDGK1互作的蛋白

Fig. 10 Yeast two-hybridization to verify proteins interacting with VvDGK1

图11 BiFC验证VvDGK1与VvFd的相互作用

Fig. 11 Analysis of the interaction between VvDGK1 with VvFd in plants by BiFC

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