SlMAPKKK43调控番茄对灰霉病的抗性

doi:10.16420/j.issn.0513-353x.2023-0757

园艺学报 ›› 2024, Vol. 51 ›› Issue (2): 309-320.doi: 10.16420/j.issn.0513-353x.2023-0757

• 遗传育种·种质资源·分子生物学 • 上一篇下一篇

SlMAPKKK43调控番茄对灰霉病的抗性

董晓南¹, 吕红梅¹, 赵立群³, 何秉青⁴, 张姣姣¹, 赵冰¹, 郭仰东¹^,²^,^*(), 张娜¹^,²^,^*()

¹ 中国农业大学园艺学院蔬菜系，北京市设施蔬菜生长发育调控重点实验室，北京 100193
² 中国农业大学三亚研究院，海南三亚 572025
³ 北京市农业技术推广站，北京 100029
⁴ 北京市昌平区农业技术推广站，北京 102200

收稿日期:2023-10-07 修回日期:2024-01-07 出版日期:2024-02-25 发布日期:2024-02-26
通讯作者:
*E-mail：zhangna@cau.edu.cn，
yaguo@cau.edu.cn
基金资助:
国家自然科学基金项目(32002053); 国家自然科学基金项目(32172598); 国家自然科学基金项目(32172599); 北京市设施蔬菜创新团队项目(BAIC01-2024)

SlMAPKKK43 Regulates Tomato Resistance to Gray Mold

DONG Xiaonan¹, LÜ Hongmei¹, ZHAO Liqun³, HE Bingqing⁴, ZHANG Jiaojiao¹, ZHAO Bing¹, GUO Yangdong¹^,²^,^*(), ZHANG Na¹^,²^,^*()

¹ Beijing Key Laboratory of Growth and Development Regulation of Protected Vegetables，Department of Vegetable，College of Horticulture，China Agricultural University，Beijing 100193，China
² Sanya Institute of China Agricultural University，Sanya，Hainan 572025，China
³ Beijing Agricultural Technology Extension Station，Beijing 100029，China
⁴ Beijing Changping District Agricultural Technology Extension Station，Beijing 102200，China

Received:2023-10-07 Revised:2024-01-07 Published:2024-02-25 Online:2024-02-26

摘要/Abstract

摘要：

促丝裂原活化蛋白激酶（Mitogen-activated protein kinase，MAPK）级联途径在植物的多种发育和生理过程中发挥重要作用，并应对各种生物和非生物胁迫。本研究中利用农杆菌介导的遗传转化方法获得了SlMAPKKK43过表达植株和CRISPR-Cas9介导的SlMAPKKK43敲除突变体，用灰葡萄孢接种转基因株系的离体叶片和果实，分析发现SlMAPKKK43正调控番茄对灰霉病的抗性。为进一步解析SlMAPKKK43调控番茄灰霉病抗性的分子机制，利用Pull down-MS技术筛选到已知的灰霉病调控因子SlMKK2和SlMKK4可能作为SlMAPKKK43的底物，利用体外磷酸化试验初步验证SlMAPKKK43可以磷酸化SlMKK2和SlMKK4。

关键词: 番茄, 灰葡萄孢, MAPK级联, SlMAPKKK43

Abstract:

The mitogen-activated protein kinase（MAPK）cascade plays an important role in multiple developmental and physiological processes in plants and in response to various biotic and abiotic stresses. In this study，SlMAPKKK43 overexpressed plants and CRISPR-Cas9-mediated SlMAPKKK43 knockout mutant using Agrobacterium infection genetic transformation were obtained. Through the inoculation of Botrytis cinerea with transgenic lines，we found that SlMAPKKK43 positively regulated the tolerance of tomato to Botrytis cinerea. In order to further elucidate the molecular mechanism of SlMAPKKK43 regulating the resistance of gray mold，the known gray mold regulators SlMKK2 and SlMKK4 were screened by pull down-MS technology，and in vitro phosphorylation assay was used to preliminarily verify that SlMAPKKK43 could phosphorylate SlMKK2 and SlMKK4.

Key words: tomato, Botrytis cinerea, MAPK cascade, SlMAPKKK43

董晓南, 吕红梅, 赵立群, 何秉青, 张姣姣, 赵冰, 郭仰东, 张娜. SlMAPKKK43调控番茄对灰霉病的抗性[J]. 园艺学报, 2024, 51(2): 309-320.

DONG Xiaonan, LÜ Hongmei, ZHAO Liqun, HE Bingqing, ZHANG Jiaojiao, ZHAO Bing, GUO Yangdong, ZHANG Na. SlMAPKKK43 Regulates Tomato Resistance to Gray Mold[J]. Acta Horticulturae Sinica, 2024, 51(2): 309-320.

图/表 9

表1 本研究中所用的引物

Table 1 The primers used in this study

用途 Function	引物名称 Primer name	序列（5′-3′） Sequence
扩增基因全长 Full length amplification	SlMAPKKK43	F：ATGGATAGTCCCACAATGACCAA R：TTTGACATTCATTGCCGCCG
构建过表达载体 Construct overexpression vector	1305-SlMAPKKK43	F：AGCTCGGTACCCGGGGATCCATGGATAGTCCCACAATGACCAA R：ATGGTCTTTGTAGTCAAGCTTTTTGACATTCATTGCCGCCG
构建基因编辑载体 Construct gene editing vector	PB-SlMAPKKK43	BsF：ATATATGGTCTCGATTGAAGTATCAGCAGCCTGACGGTT BsR：ATTATTGGTCTCGAAACGAAGAGTACAGTCGTCTTCCAA F0：TGAAGTATCAGCAGCCTGACGGTTTTAGAGCTAGAAATAGC R0：AACGAAGAGTACAGTCGTCTTCCAATCTCTTAGTCGACTCTAC
qRT-PCR	qSlMAPKKK43	F：CACATTCGAACTGGACGGGA R：CATGGGCTGATTTCCCTGGT
	qSlUbi3	F：TCCATCTCGTGCTCCGTCT R：CTGAACCTTTCCAGTGTCATCAA
	qSlEFα	F：GGAACTTGAGAAGGAGCCTAAG R：CAACACCAACAGCAACAGTCT
过表达植株鉴定 Identification of overexpressing lines	OE- SlMAPKKK43	F：GATGACGCACAATCCCACTATCC R：CCATCACCAGAATCCAACTTATCG
基因编辑植株鉴定 Identification of gene edited lines	CR- SlMAPKKK43	F1：ATGGATAGTCCCACAATGACCAA R1：TCCACATACCTTCTCTCATTTTCCC F2：ACTCTTGATGGTTGTCTCCACAA R2：TTTCCTGTATCGAGAAGGGAATGC

图1 番茄MAPKKK的进化树分析

Fig. 1 Phylogenetic analysis of MAPKKKs in tomato

图2 SlMAPKKK43在番茄不同组织中表达量的qRT-PCR分析（A）及在线（http://bar.utoronto.ca/eplant_tomato/）查询结果（B）不同的字母表示差异显著（P < 0.05）。

Fig. 2 qRT-PCR analysis of the expression of SlMAPKKK43 in different tomato tissues（A）and online （http://bar.utoronto.ca/eplant_tomato/）query results（B） Different letters above the columns indicated statistically significant differences（P < 0.05）.

图3 逆境胁迫下番茄SlMAPKKK43的转录应答显著性分析

Fig. 3 Analysis of transcriptional response of SlMAPKKK43 under stresses

图4 番茄SlMAPKKK43过表达植株的PCR（A）及其相对表达量（B）鉴定

Fig. 4 PCR identification of SlMAPKKK43 overexpressing plants（A）and relative expression level（B）

图5 SlMAPKKK43基因编辑植株的鉴定 A：SlMAPKKK43编辑靶位点示意图；B：两种基因编辑类型序列及测序结果；C：两种基因编辑类型编码氨基酸序列预测结果。

Fig. 5 Identification of SlMAPKKK43 gene-edited plants A：Schematic diagram of SlMAPKKK43 gene editing target site；B：Sequence diagrams and sequencing results of two types of gene editing；C：Prediction results of amino acid sequences encoded by two gene editing types.

图6 番茄野生型（WT）、SlMAPKKK43转基因（OE）和基因编辑（CR）株系离体叶片和果实接种灰葡萄孢后的表型及病斑面积

Fig. 6 Phenotype and lesion area of tomato wild-type（WT），SlMAPKKK43 transgenic（OE）and gene-edited（CR）lines after inoculation with Botrytis cinerea

表2 SlMAPKKK43互作蛋白筛选结果列表（部分）

Table 2 List of interaction proteins screened with SlMAPKKK43（Partial）

基因ID Gene ID	基因注释 Description
Solyc04g071615	脱落酸应激成熟蛋白 Abscicic acid stress ripening protein 4
Solyc04g074410	油菜素类固醇调节基因 Protein EXORDIUM-like
Solyc03g123800	SlMKK2 Mitogen-activated protein kinase kinase 4/5
Solyc02g080670	L-抗坏血酸氧化酶同源物 L-ascorbate oxidase homolog
Solyc06g062370	磷酸酶 Acid phosphatase 1
Solyc03g097920	SlMKK4 SF344-mitogen-activated kinase kinase kinase
Solyc02g086880	脱氢酶 Formate dehydrogenase
Solyc01g091330	谷胱甘肽转移酶 Glutathione S-transferase zeta class-like isoform X1
Solyc01g007260	F-box 蛋白 MORE AXILLARY GROWTH2 Protein SMAX1-LIKE 3

图7 SlMAPKKK43-MBP在体外直接磷酸化SlMKK2-MBP-His和SlMKK4-MBP-His α-pSer/Thr为通用型磷酸化丝氨酸/苏氨酸抗体，α-MBP为麦芽糖结合蛋白抗体，α-His为多组氨酸标签蛋白抗体，ALP为碱性磷酸酶，能对底物去磷酸化。

Fig. 7 SlMAPKKK43-MBP directly phosphorylates SlMKK2-MBP-His and SlMKK4-MBP-His in vitro α-pSer/Thr is a phosphoserine/threonine antibody，α-MBP is a maltose-binding protein antibody，α-His is His tag antibody，and ALP is an alkaline phosphatase that dephosphorylates substrates.

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SlMAPKKK43调控番茄对灰霉病的抗性

SlMAPKKK43 Regulates Tomato Resistance to Gray Mold

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SlMAPKKK43调控番茄对灰霉病的抗性

SlMAPKKK43 Regulates Tomato Resistance to Gray Mold

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