1. Introduction
Plants in natural environments face numerous abiotic stresses such as drought, salinity, and low temperatures, which severely impact their growth and development. As immobile organisms, plants must rely on their own adaptive mechanisms to cope with these challenges. Transcription factors are key regulators of gene expression, and the WRKY transcription factor family plays an essential role in plants' adaptation to stress. By specifically binding to DNA, WRKY transcription factors regulate the expression of downstream genes, thereby influencing physiological processes and enhancing plants' ability to adapt.
With advances in molecular biology techniques, an increasing number of WRKY genes have been identified and cloned, and their roles in plant responses to abiotic stress have gradually been revealed. This paper aims to discuss the functions of the WRKY transcription factor family and review their roles and mechanisms in plants' response to abiotic stress, providing insights for future research.
2. Background
Plants are sessile organisms that may encounter many abiotic environmental conditions during their life process that are detrimental to survival and growth or even lead to injury, destruction, and death. These environmental conditions are known as abiotic stress and may include low temperatures, high temperatures, drought, salinity, ultraviolet radiation, nutrient deficiencies, etc. Plants have evolved various molecular mechanisms involving signal transduction and gene expression to cope with the effects of abiotic stress. Transcription factors are DNA-binding proteins that influence gene transcription through specific interactions with the gene cis-acting elements and are an important component of plant evolution. In highly variable environments, transcription factors can regulate the physiological adaptation of plants to the environment[1]. Many transcription factor families have been identified in plants, such as MYB, bHLH, and WRKY.
The WRKY transcription factor family is one of the largest in plants and plays an important regulatory role in plant growth and development. The structure of WRKY transcription factors consists of two parts: the N-terminus contains a conserved WRKYGQK heptapeptide, and the C-terminus contains a C2H2 (C-X4-5-C-X22-23-H-X-H) or C2HC (C-X7-C-X23-H-X-C) type zinc finger structure. In the vast majority of plants, the sequence of WRKY gene family members is conserved. Still, some plants have unique sequences where WRKYGQK can be replaced by WRKYGKK or WRKYGEK, and the "RK" residues can be replaced by "RR, SK, KR, VK, or KK," and the zinc finger structure may also vary[2]. In 1994, after Ishiguro and others cloned the world's first WRKY gene SPF1 from sweet potato[3], with the continuous development of technology, an increasing number of WRKY genes have been identified and cloned. Among them, 74 WRKY gene family members were found in Arabidopsis[4], 109 in rice[5], 45 in barley[6], 83 in maritime pine[7], 81 in tomato[8], 104 in poplar[9], and 136 in maize[10]. Extensive analysis of WRKY transcription factor family members suggests that WRKY transcription factors can modulate plant hormone signaling pathways and abiotic stress signal transduction pathways, inducing the expression of many stress-related genes, thereby enhancing the plant's ability to respond to abiotic stress[11]. AtWRKY40, AtWRKY18, and AtWRKY60 can affect Arabidopsis' response to osmotic and salt stress[12], MaVQ5 and MaWRKY26 are involved in regulating cold stress processes in bananas[13], and overexpression of the OsWRKY71 gene can improve rice's cold tolerance[14]. Currently, regarding the study of the WRKY transcription factor family and plant response to abiotic stress, this article aims to discuss the functions of the WRKY transcription factor family and review the current research status of their roles and mechanisms in plant responses to abiotic stress.
2.1. The WRKY gene family and abiotic stress response
Abiotic stress stimulates interaction between plants and protective proteins, sending signals within the cell through a series of phosphorylation reactions to regulate activation or inhibition of downstream target gene transcription, control the expression of WRKY transcription factors, trigger various physiological and biochemical responses, and thereby respond to stress[15].
2.2. Drought stress
Water is the most significant environmental factor related to plant growth and development, playing a crucial role in plant internal metabolic processes. Drought stress can cause irreversible physiological damage to plant growth, antioxidant systems, and photosynthetic systems. WRKY is an important regulatory factor for plants responding to drought stress.
Studies indicate that the WRKY transcription factor family can directly influence the expression of drought-resistant genes to regulate drought resistance. WRKYTFs can regulate stress-related genes. Transgenic Arabidopsis integrating Iris germanica IgWRKY50 and IgWRKY32 showed significantly higher expression levels of stress-related genes RD29A, DREB2A, PP2CA, and ABA2 than WT after drought treatment[16].
Improving the efficiency of ROS scavenging is of great significance for enhancing plant drought resistance. WRKY TFs can regulate the expression of genes related to antioxidant enzymes and use the ROS scavenging system to reduce the content of reactive oxygen species in plants. By constructing PtWRKY2 overexpressing Arabidopsis lines, it was found that PtWRKY2 can significantly improve the survival rate of Arabidopsis under high-temperature stress. By conducting a comprehensive transcriptome analysis, it was found that PtWRKY2 may reduce ROS accumulation by enhancing the expression of antioxidant enzyme-encoding genes and thus improve plant survival rates[17].
In addition, WRKY transcription factors (TFs) control gene expression via the abscisic acid signaling pathway, impacting plant drought resistance. Abscisic acid plays a key role in regulating plant drought resistance. When plants are suffering from drought stress, ABA accumulates in the leaves and enhances the drought tolerance of terrestrial plants by inhibiting stomatal opening, increasing protective enzyme activity, and other means. Transgenic Arabidopsis overexpressing GhWRKY70 exhibited greater sensitivity to ABA-induced stomatal closure and higher expression levels of ABA biosynthesis-related genes[18]. XsWRKY20 from Xanthoceras sorbifolium can enhance the ability to cope with drought stress by integrating ROS homeostasis and ABA signaling pathways to regulate ABA signaling and antioxidant enzyme-related genes[19]. The ABA-mediated stomatal closure and other signaling pathways in Arabidopsis are specifically regulated by AtWRKY63[20]. Moreover, WRKY transcription factors can negatively regulate plant tolerance to drought stress. OsWRKY114 downregulates the transcription levels of ABA signaling pathway genes OsPYL2 and OsPYL10, and their expression decreases under drought stress to improve drought resistance[21].
2.3. Salt stress
There are large areas of saline-alkali land in arid and semi-arid regions, where it is difficult to grow crops. Enhancing crop resistance to salt stress is crucial for improving crop productivity. High concentrations of ions cause osmotic pressure imbalance within the plants, leading to metabolic disturbances and inhibited growth and development. Additionally, salt stress can induce oxidative stress, generate reactive oxygen species, affect photosynthesis, and limit plant growth. WRKY transcription factors play a key regulatory role in plants' response to salt stress.
GmWRKY54 of Glycine max inhibits the expression of STZ by positively regulating the dreb2a-mediated pathway in response to salt stress[22]. FcWRKY70 promotes the expression of arginine decarboxylase, enhancing plant salt tolerance[23]. Different WRKY genes exhibit various functions in plants; some are positive regulators, such as IbWRKY47, which significantly enhances salt tolerance in sweet potato[24], and MdWRKY55 in apple, which regulates salt stress response by forming a complex with MdNHX1, and its overexpression significantly improves salt tolerance in apple[25]. Conversely, some are negative regulators, such as CmWRKY17, which makes chrysanthemums more sensitive to salt stress[26], and the expression of the CaWRKY27 gene is induced by salt stress, with its overexpression making Arabidopsis and tobacco more sensitive to salt stress[27].
WRKY transcription factors also help plants cope with salt stress by regulating ions' homeostasis and antioxidant enzyme activities in plants. Sorghum SbWRKY50 regulates ion balance in plants by controlling the expression of SOS1 and HKT1[28]. Additionally, WRKY genes are also involved in signaling pathways such as ABA and H2O2, further regulating plant salt tolerance. For instance, GmWRKY13 reduces salt stress tolerance by up-regulating the expression of ABI1, while ZmWRKY17 inhibits tolerance to high salinity through exogenous ABA treatment[29].
2.4. Low-temperature stress
The impact of low-temperature stress on plant growth is significant. Plants that have been exposed to low temperatures see modifications in their cell structure and osmotic regulating chemicals, which account for 58% of the changes. Additionally, the expression and response mechanisms of related cold-resistant genes also change. WRKY transcription factors participate in the regulation of plant response to low-temperature stress through different signal transduction pathways. For example, PlWRKY70 of peony is widely expressed in buds and rapidly reaches peak levels 4 or 8 hours after low-temperature stress treatment[30]. WRKY contributes in the plant response to low-temperature stress via many pathways. Four LcWRKYs in loofah are considerably increased under low-temperature stress, and they contain methyl jasmonate, salicylic acid, and abscisic acid-responsive elements, all of which play essential roles in low-temperature stress[31]. In cockscomb grass, VbWRKY32 expression is upregulated after low-temperature stress, which upregulates the transcription levels of cold-responsive genes, increases antioxidant activity, maintains membrane stability, enhances osmotic regulation capacity, and increases plant low-temperature resistance[32]. In rice under low-temperature stress, the MADS-Box TF OsMADS57 and its interacting protein OsTB1 synergistically activate the transcriptional regulation of OsWRKY94, preventing tillering by inhibiting the transcription of the organ development gene D14[33].
3. Conclusion and Prospect
One of the biggest transcription factor families, the WRKY family, is essential to the regular functions of plant life. The intricate and varied roles of WRKY TF members have been thoroughly uncovered by growing amounts of study, and it is now evident how important WRKY transcription factors are to plants' reactions to abiotic stress. However, there are many members of WRKY transcription factors, and the regulatory pathways are very complex, so further research is needed on the molecular mechanisms and functions in regulating important biological functions. Grain is the lifeline of the national economy. Nowadays, the global climate is unstable, with frequent extreme disasters. Global crops will face more abiotic stress. Saline-alkaline disasters, droughts, and extremely high or low temperatures can all have an additional impact on crop development. In order to identify resistance genes, enhance planting techniques, and other aspects of crop science, we must investigate the roles and mechanisms of WRKY transcription factors in greater detail. This is a great benefit to improving crop yield and quality.
References
[1]. Pandey, S. P. , & Somssich, I. E. . (2009). The role of wrky transcription factors in plant immunity. Plant Physiology, 150(4), 1648-1655.
[2]. Wang, SY. , Wu GQ. & Wei M. .(2024).Functional mechanisms of WRKY transcription factors in regulating plant response to abiotic stresses .Chinese Journal of Biotechnology, (01), 35-52.
[3]. Ishiguro, S. , & Nakamura, K. . (1994). Characterization of a cdna encoding a novel dna-binding protein, spf1, that recognizes sp8 sequences in the 5' upstream regions of genes coding for sporamin and -amylase from sweet potato. MOLECULAR AND GENERAL GENETICS.
[4]. Ulker, B. , & Somssich, I. E. . (2004). Lker b, somssich ie. wrky transcription factors: from dna binding towards biological function. curr opin plant biol 7: 491-498. Current Opinion in Plant Biology, 7(5), 491-498.
[5]. Ross, C. A. , Liu, Y. , & Shen, Q. J. . (2010). The wrky gene family in rice (oryza sativa). Journal of Integrative Plant Biology, 49(6), 827-842.
[6]. Mangelsen Elke, Kilian Joachim, Berendzen Kenneth , Kolukisaoglu Üner , Harter Klaus, Jansson Christer & Wanke Dierk.(2008).Phylogenetic and comparative gene expression analysis of barley ( Hordeum vulgare ) WRKY transcription factor family reveals putatively retained functions between monocots and dicots.BMC Genomics(1), 194.
[7]. Donini, P. , Liu, J. J. , & Ekramoddoullah, A. K. M. . (2009). Identification and characterization of the wrky transcription factor family in pinus monticola. Genome, 52(1), 77-88.
[8]. Ling, J. , Jiang, W. , Zhang, Y. , Yu, H. , & Xie, B. . (2011). Genome-wide analysis of wrky gene family in cucumis sativus. Bmc Genomics, 12(1), 471.
[9]. He, H. , Dong, Q. , Shao, Y. , Jiang, H. , Zhu, S. , & Cheng, B. , et al. (2012). Genome-wide survey and characterization of the wrky gene family in populus trichocarpa. Plant Cell Reports, 31(7), 1199-1217.
[10]. Kai-Fa, W. , Juan, C. , Yan-Feng, C. , Ling-Juan, W. , & Dao-Xin, X. . (2012). Molecular phylogenetic and expression analysis of the complete wrky transcription factor family in maize. Dna Research An International Journal for Rapid Publication of Reports on Genes & Genomes(2), 153-164.
[11]. WU Zhen, ZHANG Ming-Ying, YAN Feng, LI Yi-min, GAO Jing, YAN Yong-Gang, ZHANG Gang.(2024).Identification and Analysis of WRKY Gene Family in Rheum palmatum L..Biotechnology Bulletin(01), 250-261.doi:10.13560/j.cnki.biotech.bull.1985.2023-0326.
[12]. Yu Heping, & Hou Hesheng. (2012). Plant WRKY transcription factors and their involvement in ABA signaling. Tianjin Agricultural Sciences, 18(6), 4.
[13]. YR, Chen, LG, Wang, HP, & Zhang, et al. (2013). Arabidopsis transcription factor wrky8 functions antagonistically with its interacting partner vq9 to modulate salinity stress tolerance. PLANT J, 2013, 74(5)(-), 730-745.
[14]. Ran, L. , Jin, Z. , Jiancai, L. , Guoxin, Z. , Qi, W. , & Wenbo, B. , et al. (2015). Prioritizing plant defence over growth through wrky regulation facilitates infestation by non-target herbivores. eLife Sciences, 4.
[15]. Xu, Z. S. , Chen, M. , Li, L. C. , & Ma, Y. Z. . (2008). Functions of the erf transcription factor family in plants. Botany-botanique, 86(9), 969-977.
[16]. Zhang Jingwei, Huang Dazhuang, Zhao Xiaojie, Zhang Man, Wang Qian, Hou Xueyan... & Sun Pai.(2022).Drought-responsive WRKY transcription factor genes IgWRKY50 and IgWRKY32 from Iris germanica enhance drought resistance in transgenic Arabidopsis#13;.Frontiers in Plant Science983600-983600.
[17]. Cui Wanning. WRKY gene mining and functional analysis in response to high temperature in mid-summer[D].Huaibei Normal University, 2023.DOI:10.27699/d.cnki.ghbmt.2023.000061.
[18]. Yuan Bo. Preliminary study on drought resistance mechanism of upland cotton transcription factor GhWRKY70 gene[D]. Zhejiang Sci-Tech University, 2023.DOI:10.27786/d.cnki.gzjlg.2023.000486.
[19]. Xiong, C. , Zhao, S. , Yu, X. , Sun, Y. , &Li, J. . (2020). Yellowhorn drought-induced transcription factor xswrky20 acts as a positive regulator in drought stress through ros homeostasis and aba signaling pathway. Plant Physiology and Biochemistry, 155.
[20]. Ren, X. , Chen, Z. , Liu, Y. , Zhang, H. , Zhang, M. , & Liu, Q. , et al. (2010). Abo3, a wrky transcription factor, mediates plant responses to abscisic acid and drought tolerance in arabidopsis. Plant Journal for Cell & Molecular Biology, 63(3), 417-429.
[21]. SONG G, SON S, LEE KS, PARK YJ, SUH EJ, LEE SI, PARK SR. (2022).OsWRKY114 negatively regulates drought tolerance by restricting stomatal closure in rice[J]. Plants, 11(15): 1938.
[22]. Zhou, Q. Y. , Tian, A. G. , Zou, H. F. , Xie, Z. M. , Lei, G. , & Huang, J. , et al. (2008). Soybean wrky-type transcription factor genes, gmwrky13, gmwrky21, and gmwrky54, confer differential tolerance to abiotic stresses in transgenic arabidopsis plants. Plant Biotechnol J, 6(5), 486-503.
[23]. Gong, X. , Zhang, J. , Hu, J. , Wang, W. , Wu, H. , & Zhang, Q. , et al. (2015). Fcwrky70, a wrky protein of fortunella crassifolia, functions in drought tolerance and modulates putrescine synthesis by regulating arginine decarboxylase gene. Plant.
[24]. Ma Yue, Xue Hao, Zhang Feng, Jiang Qiu, Yang Shuang, Yue Pengtao... & Zhang Zhihong.(2020).The miR156/SPL module regulates apple salt stress tolerance by activating MdWRKY100 expression..Plant biotechnology journal(2), 311-323.
[25]. Dong Q , Zheng W , Duan D , et al.(2020).MdWRKY30 , a group IIa WRKY gene from apple, confers tolerance to salinity and osmotic stresses in transgenic apple callus and Arabidopsis seedlings[J].Plant Science, 299110611-110611.
[26]. Peiling, Li, Aiping, Song, Chunyan, & Gao, et al. (2015). Chrysanthemum wrky gene cmwrky17 negatively regulates salt stress tolerance in transgenic chrysanthemum and arabidopsis plants. Plant Cell Reports.
[27]. Ling, H. , Yin-Huan, W. , Qian, Z. , Bei, W. , Qing-Lin, L. , & Lei, Z. . (2018). Chrysanthemum dgwrky2 gene enhances tolerance to salt stress in transgenic chrysanthemum. International Journal of Molecular Sciences, 19(7), 2062.
[28]. Yang, G. , Zhang, W. , Liu, Z. , Yi-Maer, A. Y. , & Xu, Z. . (2017). Both jrwrky2 and jrwrky7 of juglans regia mediate responses to abiotic stresses and abscisic acid through formation of homodimers and interaction. Plant Biol, 19(2), 268-278.
[29]. Cai, R. , Dai, W. , Zhang, C. , Wang, Y. , Wu, M. , & Zhao, Y. , et al. (2017). The maize wrky transcription factor zmwrky17 negatively regulates salt stress tolerance in transgenic arabidopsis plants. Planta, 246(6), 1215-1231.
[30]. Caiyun Han, Junjie Li, Yan Ma, Jing Guo, Xianfeng Guo & Jinguang Xu.(2019).PlWRKY70: a Paeonia lactiflora transcription factor that sensitively responds to low-temperature, salt, and waterlogging stresses.Canadian Journal of Plant Science(2), 146-155.
[31]. Kang Yumei, Xue Zhuzheng, Li Yongping, et al. Identification of WRKY gene family and its expression analysis under low temperature stress in loofah[J/OL].Southwest Journal of Agricultural Sciences, 1-23[2024-08-03].http://kns.cnki.net/kcms/detail/51.1213.S.20240223.0929.002.html.
[32]. Wang, M. Q. , Huang, Q. X. , Lin, P. , Zeng, Q. H. , & Zhang, F. . (2020). The overexpression of a transcription factor gene vbwrky32 enhances the cold tolerance in verbena bonariensis. Frontiers in Plant Science, 10.
[33]. Chen, Liping, Zhao, Yuan, Shujuan, & Zhang, et al. (2018). Osmads57 together with ostb1 coordinates transcription of its target oswrky94 and d14 to switch its organogenesis to defense for cold adaptation in rice. The New Phytologist.
Cite this article
Zhao,T. (2024). The review of WRKY transcription factor family functions and its abiotic stress responses in plants. Theoretical and Natural Science,67,195-200.
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The datasets used and/or analyzed during the current study will be available from the authors upon reasonable request.
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References
[1]. Pandey, S. P. , & Somssich, I. E. . (2009). The role of wrky transcription factors in plant immunity. Plant Physiology, 150(4), 1648-1655.
[2]. Wang, SY. , Wu GQ. & Wei M. .(2024).Functional mechanisms of WRKY transcription factors in regulating plant response to abiotic stresses .Chinese Journal of Biotechnology, (01), 35-52.
[3]. Ishiguro, S. , & Nakamura, K. . (1994). Characterization of a cdna encoding a novel dna-binding protein, spf1, that recognizes sp8 sequences in the 5' upstream regions of genes coding for sporamin and -amylase from sweet potato. MOLECULAR AND GENERAL GENETICS.
[4]. Ulker, B. , & Somssich, I. E. . (2004). Lker b, somssich ie. wrky transcription factors: from dna binding towards biological function. curr opin plant biol 7: 491-498. Current Opinion in Plant Biology, 7(5), 491-498.
[5]. Ross, C. A. , Liu, Y. , & Shen, Q. J. . (2010). The wrky gene family in rice (oryza sativa). Journal of Integrative Plant Biology, 49(6), 827-842.
[6]. Mangelsen Elke, Kilian Joachim, Berendzen Kenneth , Kolukisaoglu Üner , Harter Klaus, Jansson Christer & Wanke Dierk.(2008).Phylogenetic and comparative gene expression analysis of barley ( Hordeum vulgare ) WRKY transcription factor family reveals putatively retained functions between monocots and dicots.BMC Genomics(1), 194.
[7]. Donini, P. , Liu, J. J. , & Ekramoddoullah, A. K. M. . (2009). Identification and characterization of the wrky transcription factor family in pinus monticola. Genome, 52(1), 77-88.
[8]. Ling, J. , Jiang, W. , Zhang, Y. , Yu, H. , & Xie, B. . (2011). Genome-wide analysis of wrky gene family in cucumis sativus. Bmc Genomics, 12(1), 471.
[9]. He, H. , Dong, Q. , Shao, Y. , Jiang, H. , Zhu, S. , & Cheng, B. , et al. (2012). Genome-wide survey and characterization of the wrky gene family in populus trichocarpa. Plant Cell Reports, 31(7), 1199-1217.
[10]. Kai-Fa, W. , Juan, C. , Yan-Feng, C. , Ling-Juan, W. , & Dao-Xin, X. . (2012). Molecular phylogenetic and expression analysis of the complete wrky transcription factor family in maize. Dna Research An International Journal for Rapid Publication of Reports on Genes & Genomes(2), 153-164.
[11]. WU Zhen, ZHANG Ming-Ying, YAN Feng, LI Yi-min, GAO Jing, YAN Yong-Gang, ZHANG Gang.(2024).Identification and Analysis of WRKY Gene Family in Rheum palmatum L..Biotechnology Bulletin(01), 250-261.doi:10.13560/j.cnki.biotech.bull.1985.2023-0326.
[12]. Yu Heping, & Hou Hesheng. (2012). Plant WRKY transcription factors and their involvement in ABA signaling. Tianjin Agricultural Sciences, 18(6), 4.
[13]. YR, Chen, LG, Wang, HP, & Zhang, et al. (2013). Arabidopsis transcription factor wrky8 functions antagonistically with its interacting partner vq9 to modulate salinity stress tolerance. PLANT J, 2013, 74(5)(-), 730-745.
[14]. Ran, L. , Jin, Z. , Jiancai, L. , Guoxin, Z. , Qi, W. , & Wenbo, B. , et al. (2015). Prioritizing plant defence over growth through wrky regulation facilitates infestation by non-target herbivores. eLife Sciences, 4.
[15]. Xu, Z. S. , Chen, M. , Li, L. C. , & Ma, Y. Z. . (2008). Functions of the erf transcription factor family in plants. Botany-botanique, 86(9), 969-977.
[16]. Zhang Jingwei, Huang Dazhuang, Zhao Xiaojie, Zhang Man, Wang Qian, Hou Xueyan... & Sun Pai.(2022).Drought-responsive WRKY transcription factor genes IgWRKY50 and IgWRKY32 from Iris germanica enhance drought resistance in transgenic Arabidopsis#13;.Frontiers in Plant Science983600-983600.
[17]. Cui Wanning. WRKY gene mining and functional analysis in response to high temperature in mid-summer[D].Huaibei Normal University, 2023.DOI:10.27699/d.cnki.ghbmt.2023.000061.
[18]. Yuan Bo. Preliminary study on drought resistance mechanism of upland cotton transcription factor GhWRKY70 gene[D]. Zhejiang Sci-Tech University, 2023.DOI:10.27786/d.cnki.gzjlg.2023.000486.
[19]. Xiong, C. , Zhao, S. , Yu, X. , Sun, Y. , &Li, J. . (2020). Yellowhorn drought-induced transcription factor xswrky20 acts as a positive regulator in drought stress through ros homeostasis and aba signaling pathway. Plant Physiology and Biochemistry, 155.
[20]. Ren, X. , Chen, Z. , Liu, Y. , Zhang, H. , Zhang, M. , & Liu, Q. , et al. (2010). Abo3, a wrky transcription factor, mediates plant responses to abscisic acid and drought tolerance in arabidopsis. Plant Journal for Cell & Molecular Biology, 63(3), 417-429.
[21]. SONG G, SON S, LEE KS, PARK YJ, SUH EJ, LEE SI, PARK SR. (2022).OsWRKY114 negatively regulates drought tolerance by restricting stomatal closure in rice[J]. Plants, 11(15): 1938.
[22]. Zhou, Q. Y. , Tian, A. G. , Zou, H. F. , Xie, Z. M. , Lei, G. , & Huang, J. , et al. (2008). Soybean wrky-type transcription factor genes, gmwrky13, gmwrky21, and gmwrky54, confer differential tolerance to abiotic stresses in transgenic arabidopsis plants. Plant Biotechnol J, 6(5), 486-503.
[23]. Gong, X. , Zhang, J. , Hu, J. , Wang, W. , Wu, H. , & Zhang, Q. , et al. (2015). Fcwrky70, a wrky protein of fortunella crassifolia, functions in drought tolerance and modulates putrescine synthesis by regulating arginine decarboxylase gene. Plant.
[24]. Ma Yue, Xue Hao, Zhang Feng, Jiang Qiu, Yang Shuang, Yue Pengtao... & Zhang Zhihong.(2020).The miR156/SPL module regulates apple salt stress tolerance by activating MdWRKY100 expression..Plant biotechnology journal(2), 311-323.
[25]. Dong Q , Zheng W , Duan D , et al.(2020).MdWRKY30 , a group IIa WRKY gene from apple, confers tolerance to salinity and osmotic stresses in transgenic apple callus and Arabidopsis seedlings[J].Plant Science, 299110611-110611.
[26]. Peiling, Li, Aiping, Song, Chunyan, & Gao, et al. (2015). Chrysanthemum wrky gene cmwrky17 negatively regulates salt stress tolerance in transgenic chrysanthemum and arabidopsis plants. Plant Cell Reports.
[27]. Ling, H. , Yin-Huan, W. , Qian, Z. , Bei, W. , Qing-Lin, L. , & Lei, Z. . (2018). Chrysanthemum dgwrky2 gene enhances tolerance to salt stress in transgenic chrysanthemum. International Journal of Molecular Sciences, 19(7), 2062.
[28]. Yang, G. , Zhang, W. , Liu, Z. , Yi-Maer, A. Y. , & Xu, Z. . (2017). Both jrwrky2 and jrwrky7 of juglans regia mediate responses to abiotic stresses and abscisic acid through formation of homodimers and interaction. Plant Biol, 19(2), 268-278.
[29]. Cai, R. , Dai, W. , Zhang, C. , Wang, Y. , Wu, M. , & Zhao, Y. , et al. (2017). The maize wrky transcription factor zmwrky17 negatively regulates salt stress tolerance in transgenic arabidopsis plants. Planta, 246(6), 1215-1231.
[30]. Caiyun Han, Junjie Li, Yan Ma, Jing Guo, Xianfeng Guo & Jinguang Xu.(2019).PlWRKY70: a Paeonia lactiflora transcription factor that sensitively responds to low-temperature, salt, and waterlogging stresses.Canadian Journal of Plant Science(2), 146-155.
[31]. Kang Yumei, Xue Zhuzheng, Li Yongping, et al. Identification of WRKY gene family and its expression analysis under low temperature stress in loofah[J/OL].Southwest Journal of Agricultural Sciences, 1-23[2024-08-03].http://kns.cnki.net/kcms/detail/51.1213.S.20240223.0929.002.html.
[32]. Wang, M. Q. , Huang, Q. X. , Lin, P. , Zeng, Q. H. , & Zhang, F. . (2020). The overexpression of a transcription factor gene vbwrky32 enhances the cold tolerance in verbena bonariensis. Frontiers in Plant Science, 10.
[33]. Chen, Liping, Zhao, Yuan, Shujuan, & Zhang, et al. (2018). Osmads57 together with ostb1 coordinates transcription of its target oswrky94 and d14 to switch its organogenesis to defense for cold adaptation in rice. The New Phytologist.