Functions of SlWRKY45 in Response to Low-temperature and Drought Stress in Tomato

doi:10.11983/CBB24101

Abstract

Abstract: INTRODUCTION Tomato (Solanum lycopersicum), a significant warm-season and water-dependent vegetable crop, is extensively cultivated worldwide. Whether grown in open fields or protected environments, tomatoes frequently encounter various environmental stresses, including drought and low temperatures, which significantly impact their yield and quality. Transcription factors play a pivotal role in plant stress responses by modulating the expression of specific target genes, thereby transmitting perceived stress signals downstream. WRKY transcription factors in tomatoes are known to regulate responses to multiple abiotic stresses. However, the specific role of the tomato SlWRKY45 in abiotic stress responses remains unclear. RATIONALEStudies have demonstrated that WRKY transcription factors play a crucial regulatory role in plant responses to abiotic stress. As an important economic vegetable crop, tomato is susceptible to various environmental stresses during its growth and development. By genetically overexpressing SlWRKY45 in tomato and investigating its function under low-temperature and drought stress conditions, the findings can provide a theoretical foundation for understanding the complex regulatory mechanisms of WRKY transcription factors. Additionally, this research offers valuable candidate genes for breeding stress-resistant tomato varieties. RESULTSExpression analysis revealed that low-temperature, drought, and abscisic acid (ABA) treatments significantly induced the expression of SlWRKY45. Overexpression of SlWRKY45 enhanced the resistance of tomato plants to drought and low-temperature stresses. Under drought and low-temperature conditions, the photosynthetic indices, antioxidant enzyme activities, and proline (Pro) contents in SlWRKY45 overexpression lines were significantly higher than those in wild-type (WT) plants. Conversely, the accumulation of reactive oxygen species (ROS) and malondialdehyde (MDA) levels in SlWRKY45-OE plants was significantly lower than in WT plants under the same stress conditions. Transcriptome data analysis indicated that SlWRKY45 regulates tomato's response to low-temperature stress primarily by influencing antioxidant enzyme activities and stress response pathways. Dual-luciferase assays demonstrated that SlWRKY45 could directly activate the expression of SlPOD1. Furthermore, the interaction between SlWRKY45 and SlWRKY46 was confirmed through yeast two-hybrid (Y2H) and bimolecular fluorescence complementation (BiFC) assays. CONCLUSIONOur findings demonstrate that SlWRKY45 positively regulates drought resistance and low-temperature tolerance in tomato. Additionally, SlWRKY45 can interact with SlWRKY46 and directly activate the expression of SlPOD1. These results offer valuable insights for further research into the regulatory mechanisms underlying abiotic stress responses and provide potential gene resources for genetic improvement through molecular breeding. Phenotypes of SlWRKY45-overexpressing and wild-type plants under drought and low-temperature treatments

Key words: tomato, SlWRKY45, drought stress, low-temperature stress, antioxidase

CLC Number:

S641.2

Bei Fan, Min Ren, Yanfeng Wang, Fengfeng Dang, Guoliang Chen, Guoting Cheng, Jinyu Yang, Huiru Sun. Functions of SlWRKY45 in Response to Low-temperature and Drought Stress in Tomato[J]. Chinese Bulletin of Botany, 2025, 60(2): 186-203.

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URL: https://www.chinbullbotany.com/EN/10.11983/CBB24101

https://www.chinbullbotany.com/EN/Y2025/V60/I2/186

Figures/Tables 7

Figure 1 Heat map of SlWRKYs under low-temperature treatment (A) and SlWRKY45 expression under 4°C (B), drought (C), and abscisic acid (ABA) (D) treatments and in different tissues (E) (B), (C), (D) The relative expression levels of SlWRKY45 at 0 h under treatments of 4°C, ABA and PEG6000 were utilized as controls, respectively; (E) The relative expression of SlWRKY45 in stems served as the control. ** represents significant differences at P<0.01.

Figure 2 Transcriptional self-activation of SlWRKY45 pGBKT7-53+pGADT7-T and the empty vector pGBKT7 represent the positive control and negative control, respectively.

Figure 3 Phenotype (A), photosynthesis indicators (B), DAB, and NBT staining (C) and physiological indicators (D) of SlWRKY45-OE transgenic and WT tomato plants under drought treatment (B), (D) ** represent significant differences at P<0.01. DAB: Diaminobenzidine; NBT: Nitrotetrazolium blue chloride; WT: Wild type; SOD: Superoxide; POD: Peroxidase; MDA: Malondialdehyde; Pro: Proline. #1 and #2 are different transgenic lines. Bars=1 cm

Figure 4 Phenotype (A), photosynthesis indicators (B), DAB, and NBT staining (C), and physiological indicators (D), (E) of SlWRKY45-OE transgenic and WT tomato plants under 4°C treatment (B)-(E) * and ** represent significant differences at P<0.05 and P<0.01, respectively. WT: Wild type. DAB, NBT, SOD, POD, MDA, and Pro are the same as shown in Figure 3. #1 and #2 are different transgenic lines. Bars=1 cm

Figure 5 RNA-seq analysis of SlWRKY45-OE transgenic tomato and WT plants before and after low-temperature treatment (A) Volcano diagram of DEGs (red, blue and gray represent upregulated, downregulated and undifferentially expressed genes, respectively); (B) GO cluster (the red asterisks and lines indicate significantly enriched pathways); (C) KEGG enrichment (the red asterisks and lines indicate significantly enriched pathways); (D) The heatmap of 6 selected DEGs; (E) The relative expression of the 6 selected DEGs (the relative expression of WT under normal temperature was used as control. ** represent significant differences at P<0.01. WT: Wild type. #1 and #2 are different transgenic lines)

Figure 6 Dual-luciferase reporter assays analysis of SlWRKY45 on SlPOD1 and SlPOD promoters activation (A) Sequence analysis of the SlPOD1 and SlPOD promoters (the bold underlines represent W-box-like and W-box elements); (B) Schematic representation of reporter and effector vectors; (C) LUC/REN analysis (62SK+SlPOD1 pro and 62SK+SlPOD pro are the control groups. ** represents significant differences at P<0.01)

Figure 7 Interaction of SlWRKY45 and SlWRKY46 via yeast two-hybrid (Y2H) (A) and bimolecular fluorescence complementation (BiFC) (B) (A) pGBKT7-53+pGADT7-T represents a positive control, pGBKT7+pGADT7-SlWRKY45 and pGBKT7-SlWRKY46+pGADT7 represent negative controls; (B) From left to right, the yellow fluorescent signal, bright field image and merge image are shown, nYFP-SlWRKY45+cYFP and nYFP+SlWRKY46-cYFP represent negative controls. Bars=20 µm

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