植物在长期进化过程中，为适应复杂多变的外界环境，已形成一套精密的分子机制来感知和响应各种生物与非生物胁迫。在非生物胁迫（如高温、低温、干旱、高盐等）条件下，植物激素脱落酸（ABA）的含量显著增加。ABA通过多重生理途径提升植物的抗逆性，包括调控种子休眠状态、降低蒸腾作用、延缓生长发育进程以及激活抗逆相关基因的表达，从而帮助植物更好地适应不良环境。在ABA介导的应激响应过程中，促分裂原活化蛋白激酶（MAPK）级联信号通路（MAPKKK-MAPKK-MAPK）已被揭示在调节植物非生物胁迫的反应中起着重要的作用。我们实验室先前的研究表明，ABA激活的MKK1-MPK1级联通路在水稻ABA响应和胁迫耐受性调控中具有重要功能，且MAPKKK28能够与MKK1发生相互作用并在体外对其进行磷酸化修饰。然而，在ABA信号转导过程中，MAPKKK28是否确实作用于MKK1-MPK1级联上游有待阐明。值得注意的是，在酵母和哺乳动物细胞中，STE20/MAP4Ks类激酶能够磷酸化并调控下游MAP3Ks，但在植物中尚未发现MAP4Ks直接调控下游MAPKKKs的报道。

本研究利用生物化学与分子生物学，细胞生物学，遗传学以及植物生理学的研究手段，首先探究了MAPKKK28在ABA介导的MKK1-MPK1级联活化以及ABA介导的生理反应中的作用，进而分析了MAP4K5在调控ABA介导的MAPKKK28活化以及ABA介导的生理反应中的作用。主要研究结果如下：

（1）为了探究MAPKKK28在ABA信号转导以及氧化胁迫和渗透胁迫中的功能，我们成功获得了纯合的T2代MAPKKK28基因敲除突变体材料（mapkkk28-KO）和MAPKKK28过表达材料（MAPKKK28-OE）。利用这些材料，我们首先探究了MAPKKK28在ABA调控中的生理反应中的作用。研究结果表明，MAPKKK28显著增强了水稻对ABA调控的生理反应（包括种子萌发，根系生长以及气孔关闭）的敏感性。我们也探究了MAPKKK28在水稻对氧化胁迫和渗透胁迫响应中的作用。研究结果显示，MAPKKK28能显著提高水稻在氧化胁迫（100 mM H₂O₂处理）与渗透胁迫（20%聚乙二醇（PEG）6000处理）下的存活率，降低活性氧（ROS，包括H₂O₂与O₂^.-）积累、丙二醛（MDA）含量以及电解质泄漏率。这些结果表明，MAPKKK28可提升水稻对ABA的响应能力，并增强幼苗在氧化胁迫和渗透胁迫下的抗逆性。

（2）为了确定MAPKKK28磷酸化MKK1属于经典的MAPK级联的磷酸化方式，利用体外磷酸化实验证明了MAPKKK28磷酸化MKK1活化环中的Ser-215和Thr-221位点。为了验证MKK1的Ser-215和Thr-221位点磷酸化状态是否会影响其与MAPKKK28的相互作用关系。利用双分子荧光互补（BiFC）、萤火虫荧光素酶互补成像技术（LCI）和酵母双杂（Y2H）实验结果表明，MKK1蛋白的Ser-215和Thr-221位点的磷酸化修饰对其与MAPKKK28的结合能力没有显著影响。

（3）为了阐明MAPKKK28与MKK1-MPK1在ABA信号通路中的级联关系及其功能，本研究通过多种实验方法进行了系统探究。首先，通过免疫共沉淀（Co-IP）和LCI实验证实，ABA处理显著增强了MAPKKK28与MKK1之间的相互作用。进一步的免疫沉淀凝胶激酶实验表明，在ABA作用下，MKK1的激酶活性依赖于MAPKKK28。这些结果表明MAPKKK28在ABA信号通路中位于MKK1的上游发挥调控作用。为在遗传层面证实MAPKKK28-MKK1级联的功能，本研究构建了mapkkk28-KO/mkk1-KO、MAPKKK28-OE/mkk1-KO和mapkkk28-KO/MKK1-OE等系列遗传材料。通过遗传学和生物化学分析，我们证实了MAPKKK28作用于MKK1-MPK1级联上游来调节水稻的ABA反应以及水稻对氧化胁迫与渗透胁迫的耐性。

（4）为了弄清楚MAP4K激酶是否属于MAP3K真正的上游调控因子，我们利用Y2H、GST-pull down、LCI、Co-IP揭示了一个水稻OsMAP4K成员—MAP4K5与MAPKKK28在体内外存在互作，且MAP4K5可以体外磷酸化MAPKKK28。为了更好的了解MAP4K5的具体特征和功能，我们进行了一系列的生化分析和遗传分析。结果表明，MAP4K5与拟南芥的SIK1同源性最高，具有典型的STE20（MAP4K）的不变的赖氨酸K、GxGxxG、HRD和DFG结构特点。MAP4K5定位于细胞质、细胞膜和细胞核，并且MAP4K5的表达水平可以被ABA、H₂O₂和PEG所诱导。为了研究MAP4K5是否在水稻中响应氧化胁迫和渗透胁迫，首先，我们利用ABA和20% PEG 6000处理WT水稻幼苗，发现MAP4K5的激酶活性可以被ABA和PEG所诱导。遗传表型分析结果显示，非生物胁迫后map4k5-KO突变体水稻存活率低于WT水稻，而MAP4K5-OE过表达水稻存活率高于WT植株。同时测定MDA和电解质泄漏率，发现非生物胁迫后map4k5-KO突变体水稻叶片损伤远远高于WT水稻，而MAP4K5-OE过表达水稻的情况则相反。这表明MAP4K5通过增强水稻细胞清除ROS的能力，从而减少水稻幼苗期细胞受到的氧化损伤。免疫沉淀凝胶激酶实验证明在ABA途径中，MAPKKK28的激酶活性依赖于MAP4K5。

综上所述，本研究揭示了一条新的ABA激活的完整MAPK级联通路—MAPKKK28-MKK1-MPK1。更重要的是发现了在ABA信号转导中，MAP4K5是MAPKKK28上游的直接激活子，从而揭示了植物细胞中一种调节MAPK级联活化的新机制。这一研究为利用分子模块设计育种，提高作物的耐逆性提供了重要的理论基础。

During the long-term evolutionary process, plants have developed a sophisticated molecular mechanism to perceive and respond to various biotic and abiotic stresses in order to adapt to complex and changing external environments. Under abiotic stress conditions such as high temperature, low temperature, drought, and high salinity, the level of the plant hormone abscisic acid (ABA) significantly increases. As a crucial "stress hormone", ABA enhances plant adaptation to stressful environments through multiple physiological mechanisms, including promoting dormancy, reducing water loss, inhibiting plant growth, and inducing the expression of stress-related genes. In ABA signal transduction, the mitogen-activated protein kinase MAPK cascade signaling pathway (MAPKKK-MAPKK-MAPK) has been shown to play a pivotal role in regulating plant responses to abiotic stress.

Previous research in our laboratory has demonstrated that the ABA-activated MKK1-MPK1 cascade pathway plays a significant role in ABA response and stress tolerance regulation in rice. Moreover, MAPKKK28 has been found to interact with MKK1 and phosphorylate it in vitro. However, whether MAPKKK28 indeed acts upstream of the MKK1-MPK1 cascade during ABA signal transduction remains to be elucidated. Notably, in yeast and mammalian cells, STE20/MAP4Ks-like kinases can phosphorylate and regulate downstream MAP3Ks, but there has been no report of MAP4Ks directly regulating downstream MAPKKKs in plants.

This study employed methodologies from biochemistry and molecular biology, cell biology, genetics, and plant physiology to explore the role of MAPKKK28 in the ABA-mediated activation of the MKK1-MPK1 cascade and ABA-mediated physiological responses. Subsequently, the study analyzed the role of MAP4K5 in regulating the ABA-mediated activation of MAPKKK28 and ABA-mediated physiological responses. The main findings are as follows:

(1) In order to explore the function of MAPKKK28 in ABA signal transduction, oxidative stress and osmotic stress, we successfully obtained homozygous T2 transgenic MAPKKK28 gene knockout mutant materials (mapkkk28-KO) and overexpression materials (MAPKKK28-OE). Using these materials, we first investigated the role of MAPKKK28 in physiological responses regulated by ABA. The results showed that MAPKKK28 significantly enhanced the physiological responses of rice to ABA regulation, including seed germination, root growth, and stomatal closure. We also investigated the role of MAPKKK28 in rice responses to oxidative and osmotic stress. The results showed that MAPKKK28 could significantly improve the survival rate of rice under oxidative stress (100 mM H₂O₂ treatment) and osmotic stress (20% polyethylene glycol (PEG) 6000 treatment), and reduce the accumulation of reactive oxygen species (ROS, including H₂O₂ and O₂^.-), malondialdehyde (MDA) content and electrolyte leakage rate. These results indicated that MAPKKK28 could positively regulate the sensitivity of rice to ABA and enhance the tolerance of rice seedlings to oxidative stress and osmotic stress.

(2) In order to confirm that phosphorylation of MKK1 by MAPKKK28 belongs to the classical phosphorylation mode of MAPK cascade, in vitro phosphorylation experiments were carried out. The result demonstrated that MAPKKK28 phosphorylated Ser-215 and Thr-221 sites in the activation loop of MKK1. To explore whether the phosphorylation status of Ser-215 and Thr-221 in MKK1 affects its interaction with MAPKKK28, the Bimolrcular fluorescence complementation (BiFC), the firefly luciferase complementation imaging (LCI) and Yeast two-hybrid (Y2H) experiments were employed. The results confirmed that the phosphorylation status of Ser-215 and Thr-221 in MKK1 does not influence its interaction with MAPKKK28.

(3) To elucidate the relationship between MAPKKK28 and MKK1-MPK1 cascade in ABA signaling pathway and their functions, this study systematically explored through a variety of experimental methods. First, we confirmed that ABA treatment significantly enhanced the interaction between MAPKKK28 and MKK1 by Co-IP (Co-immunoprecipitation) and LCI assays. Further immunoprecipitation gel kinase assays revealed that MKK1’s kinase activity was dependent on MAPKKK28 under ABA. These results suggested that MAPKKK28 plays a regulatory role upstream of MKK1 in the ABA signaling pathway. In order to confirm the function of the MAPKKK28-MKK1 cascade at the genetic level, a series of genetic materials such as mapkkk28-KO/mkk1-KO, MAPKKK28-OE/mkk1-KO and mapkkk28-KO/MKK1-OE were constructed. Through genetic and biochemical analysis, we demonstrated that MAPKKK28 acted upstream of the MKK1-MPK1 cascade to regulate ABA response and tolerance to oxidative stress and osmotic stress in rice.

(4) To clarify whether MAP4K kinase is the true upstream regulator of MAP3K, we used Y2H, GST-pull down, LCI, and Co-IP assays to reveal that MAP4K5, a member of rice OsMAP4K, can interact with MAPKKK28 in vivo and in vitro. Moreover, MAP4K5 can phosphorylate MAPKKK28 in vitro. In order to better understand the specific characteristics and functions of MAP4K5, we conducted a series of biochemical and genetic analyses. The results showed that MAP4K5 had the highest homology with SIK1 of Arabidopsis thaliana, with the typical STE20 (MAP4K) invariant lysine K, GxGxxG, HRD and DFG structure characteristics. MAP4K5 is localized in cytoplasm, cell membrane and nucleus, and the expression level of MAP4K5 can be induced by ABA, H₂O₂ and PEG. To explore whether MAP4K5 responds to oxidative and osmotic stress in rice, we first treated WT rice seedlings with ABA and 20% PEG 6000 and found that MAP4K5 kinase activity can be induced by ABA and PEG. Phenotypic analysis revealed that the survival rate of map4k5-KO mutant rice was lower than that of WT rice after abiotic stress, while the survival rate of MAP4K5-OE overexpressing rice was higher than that of WT plants. Simultaneously, measurements of MDA and electrolyte leakage rates showed that the leaf damage in map4k5-KO rice was significantly higher than that in WT rice after abiotic stress, whereas the situation was the opposite in MAP4K5-OE rice. The results indicated that the MAP4K5 enhances the ability of rice cells to clear ROS, thereby reducing oxidative damage to cells during the seedling stage. Immunoprecipitation kinase assay demonstrated that the kinase activity of MAPKKK28 is dependent on MAP4K5 in the ABA pathway,.

In summary, this study unveiled a novel ABA-activated complete MAPK cascade pathway—MAPKKK28-MKK1-MPK1. More importantly, it identified MAP4K5 as the direct upstream activator of MAPKKK28 in ABA signal transduction, revealing a new mechanism regulating MAPK cascade activation in plant cells. This research provides a significant theoretical foundation for utilizing molecular module design in breeding to enhance crop stress tolerance.

[1] 陈婧．水稻OsMKKK28参与ABA诱导的抗氧化防护且与OsMKK1/OsMKK6相互作用［D］．南京：南京农业大学，2020.

[2] 梁丹，张朝昕，王冕，等．花生MAPK13基因的克隆及表达分析研究［J］．花生学报，2019，48（2）：10－18．

[3] 马金姣，兰金苹，张彤，等．过表达OsMPK17激酶蛋白质增强了水稻的耐旱性［J］．作物学报，2020，46（1）：20－30．

[4] 任雯．玉米干旱胁迫和镉毒害响应蛋白鉴定与分析 [D] ．北京：中国农业科学院，2018．

[5] Adachi M, Fukuda M, Nishida E. Two co-existing mechanisms for nuclear import of MAP kinase: passive diffusion of a monomer and active transport of a dimer[J]. The EMBO Journal, 1999, 18(19):5347-5358.

[6] Addicott F T, Lyon J L, Ohkuma K, et al. Abscisic acid: A new name for abscisin II (dormin)[J]. Science, 1968, 159(3822):1493.

[7] Agurla S, Gahir S, Munemasa S, et al. Mechanism of stomatal closure in plants exposed to drought and cold stress[J]. Advances in Experimental Medicine and Biology, 2018, 1081:215-232.

[8] Ali F, Qanmber G, Li F, et al. Updated role of ABA in seed maturation, dormancy, and germination[J]. Journal of Advanced Research, 2021, 35:199-214.

[9] Alzwiy I A, Morris P C. A mutation in the Arabidopsis MAP kinase kinase 9 gene results in enhanced seedling stress tolerance[J]. Plant Science, 2007, 173(3):302-308.

[10] Ariel F, Diet A, Verdenaud M, et al. Environmental regulation of lateral root emergence in Medicago truncatula requires the HD-Zip I transcription factor HB1[J]. Plant Cell, 2010, 22(7):2171-2183.

[11] Asai T, Tena G, Plotnikova J, et al. MAP kinase signalling cascade in Arabidopsis innate immunity[J]. Nature, 2002, 415(6875):977-983.

[12] Baek D, Kim M C, Kumar D, et al. AtPR5K2, a PR5-like receptor kinase, modulates plant responses to drought stress by phosphorylating protein phosphatase 2Cs[J]. Frontiers in Plant Science, 2019, 10:1146.

[13] Banno H, Hirano K, Nakamura T, et al. NPK1, a tobacco gene that encodes a protein with a domain homologous to yeast BCK1, STE11, and Byr2 protein kinases[J]. Molecular and Cellular Biology, 1993, 13(8):4745-4752.

[14] Barrero J M, Piqueras P, González-Guzmán M, et al. A mutational analysis of the ABA1 gene of Arabidopsis thaliana highlights the involvement of ABA in vegetative development[J]. Journal of Experimental Botany, 2005, 56(418):2071-2083.

[15] Benhamman R, Bai F, Drory S B, et al. The Arabidopsis Mitogen-Activated Protein Kinase Kinase Kinase 20 (MKKK20) acts upstream of MKK3 and MPK18 in two separate signaling pathways involved in root microtubule functions[J]. Frontiers in Plant Science, 2017, 8:1352.

[16] Bhaskara G B, Wen T N, Nguyen T T, et al. Protein phosphatase 2Cs and microtubule-associated stress protein 1 control microtubule stability, plant growth, and drought response[J]. Plant Cell, 2017, 29(1):169-191.

[17] Bi G, Zhou Z, Wang W, et al. Receptor-like cytoplasmic kinases directly link diverse pattern recognition Receptors to the activation of mitogen-activated protein kinase cascades in Arabidopsis[J]. Plant Cell, 2018, 30(7):1543-1561.

[18] Brandt B, Brodsky D E, Xue S, et al. Reconstitution of abscisic acid activation of SLAC1 anion channel by CPK6 and OST1 kinases and branched ABI1 PP2C phosphatase action[J]. Proceedings of the National Academy of Sciences of the United States of America, 2012, 109(26):10593-10598.

[19] Brenner D, Brechmann M, Röhling S, et al. Phosphorylation of CARMA1 by HPK1 is critical for NF-kappaB activation in T cells[J]. Proceedings of the National Academy of Sciences of the United States of America, 2009, 106(34):14508-14513.

[20] Cai G, Wang G, Wang L, et al. A maize mitogen-activated protein kinase kinase, ZmMKK1, positively regulated the salt and drought tolerance in transgenic Arabidopsis[J]. Journal of Plant Physiology, 2014a, 171(12):1003-1016.

[21] Cai G, Wang G, Wang L, et al. ZmMKK1, a novel group a mitogen-activated protein kinase kinase gene in maize, conferred chilling stress tolerance and was involved in pathogen defense in transgenic tobacco[J]. Plant Science, 2014b, 214:57-73.

[22] Çakır B, Kılıçkaya O. Mitogen-activated protein kinase cascades in Vitis vinifera[J]. Frontiers in Plant Science, 2015, 6:556.

[23] Carrera A C, Alexandrov K, Roberts T M. The conserved lysine of the catalytic domain of protein kinases is actively involved in the phosphotransfer reaction and not required for anchoring ATP[J]. Proceedings of the National Academy of Sciences of the United States of America, 1993, 90(2):442-446.

[24] Castells E, Puigdomènech P, Casacuberta J M. Regulation of the kinase activity of the MIK GCK-like MAP4K by alternative splicing[J]. Plant Molecular Biology, 2006, (4-5):747-756.

[25] Chae M J, Lee J S, Nam M H, et al. A rice dehydration-inducible SNF1-related protein kinase 2 phosphorylates an abscisic acid responsive element-binding factor and associates with ABA signaling[J]. Plant Molecular Biology, 2007, 63(2):151-169.

[26] Chaiwongsar S, Strohm A K, Su S H, et al. Genetic analysis of the Arabidopsis protein kinases MAP3Kε1 and MAP3Kε2 indicates roles in cell expansion and embryo development[J]. Frontiers in Plant Science, 2012, 3:228.

[27] Champion A, Picaud A, Henry Y. Reassessing the MAP3K and MAP4K relationships[J]. Trends in Plant Science, 2004, 9(3):123-129.

[28] Chang Y, Nguyen B H, Xie Y, et al. Co-overexpression of the constitutively active form of OsbZIP46 and ABA-activated protein kinase SAPK6 improves drought and temperature stress resistance in rice[J]. Frontiers in Plant Science, 2017, 8:1102.

[29] Chardin C, Schenk S T, Hirt H, et al. Mitogen-Activated Protein Kinases in nutritional signaling in Arabidopsis[J]. Plant Science, 2017, 260:101-108.

[30] Chen J, Wang L, Yang Z, et al. The rice Raf-like MAPKKK OsILA1 confers broad-spectrum resistance to bacterial blight by suppressing the OsMAPKK4-OsMAPK6 cascade[J]. Journal of Integrative Plant Biology, 2021a, 63(10):1815-1842.

[31] Chen J, Wang L, Yuan M. Update on the roles of rice MAPK cascades[J]. International Journal of Molecular Sciences, 2021b, 22(4):1679.

[32] Chen K, Li G J, Bressan R A, et al. Abscisic acid dynamics, signaling, and functions in plants[J]. Journal of Integrative Plant Biology, 2020a, 62(1):25-54.

[33] Chen L, Sun H, Wang F, et al. Genome-wide identification of MAPK cascade genes reveals the GhMAP3K14-GhMKK11-GhMPK31 pathway is involved in the drought response in cotton[J]. Plant Molecular Biology, 2020b, (1-2):211-223.

[34] Chen L, Zhang B, Xia L, et al. The GhMAP3K62-GhMKK16-GhMPK32 kinase cascade regulates drought tolerance by activating GhEDT1-mediated ABA accumulation in cotton[J]. Journal of Advanced Research, 2023, 51:13-25.

[35] Chen M, Ni L, Chen J, et al. Rice calcium/calmodulin-dependent protein kinase directly phosphorylates a mitogen-activated protein kinase kinase to regulate abscisic acid responses[J]. Plant Cell, 2021c, 33(5):1790-1812.

[36] Chen Q, Hu T, Li X, et al. Phosphorylation of SWEET sucrose transporters regulates plant root:shoot ratio under drought[J]. Nature Plants, 2022, 8(1):68-77.

[37] Chen X, Wang J, Zhu M, et al. A cotton Raf-like MAP3K gene, GhMAP3K40, mediates reduced tolerance to biotic and abiotic stress in Nicotiana benthamiana by negatively regulating growth and development[J]. Plant Science, 2015, 240:10-24.

[38] Cheng Z, Li J F, Niu Y, et al. Pathogen-secreted proteases activate a novel plant immune pathway[J]. Nature, 2015, 521(7551):213-216.

[39] Chinchilla D, Zipfel C, Robatzek S, et al. A flagellin-induced complex of the receptor FLS2 and BAK1 initiates plant defence[J]. Nature, 2007, 448(7152):497-500.

[40] Chmielecki J, Peifer M, Jia P, et al. Targeted next-generation sequencing of DNA regions proximal to a conserved GXGXXG signaling motif enables systematic discovery of tyrosine kinase fusions in cancer[J]. Nucleic Acids Research, 2010, (20):6985-6996.

[41] Cho S K, Larue C T, Chevalier D, et al. Regulation of floral organ abscission in Arabidopsis thaliana[J]. Proceedings of the National Academy of Sciences of the United States of America, 2008, 105(40):15629-15634.

[42] Chuang H C, Wang X, Tan T H. MAP4K family kinases in immunity and inflammation[J]. Advances in Immunology, 2016, 129:277-314.

[43] Colcombet J, Hirt H. Arabidopsis MAPKs: a complex signalling network involved in multiple biological processes[J]. The Biochemical Journal, 2008, 413(2):217-226.

[44] Cornforth J W, Milborrow B V, Ryback G, et al. Chemistry and physiology of ‘Dormins’ in sycamore: Identity of sycamore ‘Dormin’ with abscisin II[J]. Nature, 1965, 205:1269-1270.

[45] Cui L, Song Y, Zhao Y, et al. Nei 6 You 7075, a hybrid rice cultivar, exhibits enhanced disease resistance and drought tolerance traits[J]. BMC Plant Biology, 2024, 24(1):1252.

[46] Dan I, Watanabe N M, Kusumi A. The Ste20 group kinases as regulators of MAP kinase cascades[J]. Trends in Cell Biology, 2001, 11(5):220-230.

[47] Dangol S, Nguyen N K, Singh R, et al. Mitogen-activated protein kinase OsMEK2 and OsMPK1 signaling is required for ferroptotic cell death in Rice-Magnaporthe oryzae interactions[J]. Frontiers in Plant Science, 2021, 12:710794.

[48] Danquah A, De Zelicourt A, Boudsocq M, et al. Identification and characterization of an ABA-activated MAP kinase cascade in Arabidopsis thaliana[J]. Plant Journal, 2015, 82(2):232-244.

[49] Danquah A, de Zelicourt A, Colcombet J, et al. The role of ABA and MAPK signaling pathways in plant abiotic stress responses[J]. Biotechnology Advances, 2014, 32(1):40-52.

[50] de Zelicourt A, Colcombet J, Hirt H. The role of MAPK modules and ABA during abiotic stress signaling[J]. Trends in Plant Science, 2016, 21(8):677-685.

[51] del Pozo O, Pedley K F, Martin G B. MAPKKK alpha is a positive regulator of cell death associated with both plant immunity and disease[J]. The EMBO Journal, 2004, 23(15):3072-3082.

[52] Demir F, Horntrich C, Blachutzik J O, et al. Arabidopsis nanodomain-delimited ABA signaling pathway regulates the anion channel SLAH3[J]. Proceedings of the National Academy of Sciences of the United States of America, 2013, 110(20):8296-8301.

[53] Ding Y, Cao J, Ni L, et al. ZmCPK11 is involved in abscisic acid-induced antioxidant defence and functions upstream of ZmMPK5 in abscisic acid signalling in maize[J]. Journal of Experimental Botany, 2013, 64(4):871-884.

[54] Ding Y, Lv J, Shi Y, et al. EGR2 phosphatase regulates OST1 kinase activityand freezing tolerance in Arabidopsis[J]. EMBO Journal, 2019, 38(1):e99819.

[55] Du X, Jin Z, Liu Z, et al. H2S persulfidated and increased kinase activity of MPK4 to response cold stress in Arabidopsis[J]. Frontiers in Molecular Biosciences, 2021, 8:635470.

[56] Duan H, Ma Y C, Liu R, et al. Effect of combined waterlogging and salinity stresses on euhalophyte Suaeda glauca[J]. Plant Physiology and Biochemistry, 2018, 127:231-237.

[57] Duan L, Dietrich D, Ng C H, et al. Endodermal ABA signaling promotes lateral root quiescence during salt stress in Arabidopsis seedlings[J]. Plant Cell, 2013, 25(1):324-341.

[58] Eagles C F, Wareing P F. Dormancy regulators in woody plants: Experimental induction of dormancy in Betula pubescens[J]. Nature, 1963, 199:874-875.

[59] Ehonen S, Yarmolinsky D, Kollist H, et al. Reactive oxygen species, photosynthesis, and environment in the regulation of stomata[J]. Antioxidants and Redox Signaling, 2019, 30(9):1220-1237.

[60] Engelman J A, Lisanti M P, Scherer P E. Specific inhibitors of p38 mitogen-activated protein kinase block 3T3-L1 adipogenesis[J]. Journal of Biological Chemistry, 1998, 273(48):32111-32120.

[61] Farooq A, Zhou M M. Structure and regulation of MAPK phosphatases[J]. Cellular Signalling, 2004, 16(7):769-779.

[62] Fedoroff N V, Battisti D S, Beachy R N, et al. Radically rethinking agriculture for the 21st century[J]. Science, 2010, 327(5967):833-834.

[63] Finkelstein R R, Lynch T J. The Arabidopsis abscisic acid response gene ABI5 encodes a basic leucine zipper transcription factor[J]. Plant Cell, 2000, 12(4):599-609.

[64] Franz S, Ehlert B, Liese A, et al. Calcium-dependent protein kinase CPK21 functions in abiotic stress response in Arabidopsis thaliana[J]. Molecular Plant, 2011, 4(1):83-96.

[65] Frey A, Godin B, Bonnet M, et al. Maternal synthesis of abscisic acid controls seed development and yield in Nicotiana plumbaginifolia[J]. Planta, 2004, 218(6):958-964.

[66] Fu S F, Chou W C, Huang D D, et al. Transcriptional regulation of a rice mitogen-activated protein kinase gene, OsMAPK4, in response to environmental stresses[J]. Plant Cell and Physiology. 2002, 43(8):958-963.

[67] Fujii H, Chinnusamy V, Rodrigues A, et al. In vitro reconstitution of an abscisic acid signalling pathway[J]. Nature, 2009, 462(7273):660-664.

[68] Fujii H, Verslues P E, Zhu J K. Identification of two protein kinases required for abscisic acid regulation of seed germination, root growth, and gene expression in Arabidopsis[J]. Plant Cell, 2007, 19(2):485-494.

[69] Fujii H, Zhu J K. Arabidopsis mutant deficient in 3 abscisic acid-activated protein kinases reveals critical roles in growth, reproduction, and stress[J]. Proceedings of the National Academy of Sciences of the United States of America, 2009, 106(20):8380-8385.

[70] Furuya T, Matsuoka D, Nanmori T. Membrane rigidification functions upstream of the MEKK1-MKK2-MPK4 cascade during cold acclimation in Arabidopsis thaliana[J]. FEBS Letters, 2014, 588(11):2025-2030.

[71] Gao Z, Chen Y F, Randlett M D, et al. Localization of the Raf-like kinase CTR1 to the endoplasmic reticulum of Arabidopsis through participation in ethylene receptor signaling complexes[J]. Journal of Biological Chemistry, 2003, 278(36):34725-34732.

[72] Geiger D, Maierhofer T, Al-Rasheid K A, et al. Stomatal closure by fast abscisic acid signaling is mediated by the guard cell anion channel SLAH3 and the receptor RCAR1[J]. Science Signaling, 2011, 4(173):ra32.

[73] Ghassemian M, Nambara E, Cutler S, et al. Regulation of abscisic acid signaling by the ethylene response pathway in Arabidopsis[J]. Plant Cell, 2000, 12(7):1117-1126.

[74] Gu L, Liu Y, Zong X, et al. Overexpression of maize mitogen-activated protein kinase gene, ZmSIMK1 in Arabidopsis increases tolerance to salt stress[J]. Molecular Biology Reports, 2010, 37(8):4067-4073.

[75] Gupta A, Rico-Medina A, Caño-Delgado A I. The physiology of plant responses to drought[J]. Science, 2020, 368(6488):266-269.

[76] Haak D C, Fukao T, Grene R, et al. Multilevel regulation of abiotic stress responses in plants[J]. Frontiers in Plant Science, 2017, 8:1564.

[77] Hamel L P, Nicole M C, Sritubtim S, et al. Ancient signals: Comparative genomics of plant MAPK and MAPKK gene families[J]. Trends in Plant Science, 2006, 11(4):192-198.

[78] Han W, Rong H, Zhang H, et al. Abscisic acid is a negative regulator of root gravitropism in Arabidopsis thaliana[J]. Biochemical and Biophysical Research Communications, 2009, 378(4):695-700.

[79] Hanks S K, Hunter T. The eukaryotic protein kinase superfamily: kinase (catalytic) domain structure and classification 1[J]. The FASEB Journal,1995, (9):576-596.

[80] Hanks S K, Quinn A M, Hunter T. The protein kinase family: conserved features and deduced phylogeny of the catalytic domains[J]. Science, 1988, 241(4861):42-52.

[81] Hanks S K, Quinn A M. Protein kinase catalytic domain sequence database: identification of conserved features of primary structure and classification of family members[J]. Methods in Enzymology, 1991, 200:38-62.

[82] Hao Q, Yin P, Li W, et al. The molecular basis of ABA-independent inhibition of PP2Cs by a subclass of PYL proteins[J]. Molecules and Cells, 2011, 42(5):662-672.

[83] Hardin S C, Wolniak S M. Expression of the mitogen-activated protein kinase kinase ZmMEK1 in the primary root of maize[J]. Planta, 2001, 213(6):916-926.

[84] He Y, Hao Q, Li W, et al. Identification and characterization of ABA receptors in Oryza sativa[J]. PLoS One, 2014, 9(4):e95246.

[85] Hemberg T. Growth-inhibiting substances in terminal buds of fraxinus[J]. Plant Physiology, 1949a, 2:37-44.

[86] Hemberg T. Significance of growth-inhibiting substances and auxins for the rest-period of the potato tuber[J]. Physiologia Plantarum, 1949b, 2:24-36.

[87] Hemmer W, McGlone M, Tsigelny I, et al. Role of the glycine triad in the ATP-binding site of cAMP-dependent protein kinase[J]. Journal of Biological Chemistry, 1997, 272(27):16946-16954.

[88] Hirayama T, Shinozaki K. Perception and transduction ofabscisic acid signals: Keys to the function of the versatile plant hor-mone ABA[J]. Trends in Plant Science, 2007, 12(8):343-435.

[89] Hoang M H, Nguyen X C, Lee K, et al. Phosphorylation by AtMPK6 is required for the biological function of AtMYB41 in Arabidopsis[J]. Biochemical and Biophysical Research Communications, 2012, 422(1):181-186.

[90] Hosotani S, Yamauchi S, Kobayashi H, et al. A BLUS1 kinase signal and a decrease in intercellular CO2 concentration are necessary for stomatal opening in response to blue light[J]. Plant Cell, 2021, 33(5):1813-1827.

[91] Huai J, Wang M, He J, et al. Cloning and characterization of the SnRK2 gene family from Zea mays[J]. Plant Cell Reports, 2008, 27(12):1861-1868.

[92] Huang R, Zheng R, He J, et al. Noncanonical auxin signaling regulates cell division pattern during lateral root development[J]. Proceedings of the National Academy of Sciences of the United States of America, 2019, 116(42):21285-21290.

[93] Hwang J U, Yim S, Do T H T, et al. Arabidopsis thaliana Raf22 protein kinase maintains growth capacity during postgerminative growth arrest under stress[J]. Plant Cell and Environment, 2018, 41(7):1565-1578.

[94] Iftikhar H, Naveed N, Virk N,et al. In silico analysis reveals widespread presence of three gene families, MAPK, MAPKK and MAPKKK, of the MAPK cascade from crop plants of Solanaceae in comparison to the distantly-related syntenic species from Rubiaceae, coffee[J]. PeerJ, 2017, 5:e3255.

[95] Jagodzik P, Tajdel-Zielinska M, Ciesla A, et al. Mitogen-activated protein kinase cascades in plant hormone signaling[J]. Frontiers in Plant Science, 2018, 9:1387.

[96] Jeong S, Lim C W, Kim M. et al. Modulation of phosphorylation status of MAP3 kinases under abiotic stress responses[J]. Physiologia Plantarum, 2024, 176:e14146.

[97] Jeong S, Lim C W, Lee S C. Pepper SnRK2.6-activated MEKK protein CaMEKK23 is directly and indirectly modulated by clade A PP2Cs in response to drought stress[J]. New Phytologist, 2023, 238(1):237-251.

[98] Jia H, Hao L, Guo X, et al. A Raf-like MAPKKK gene, GhRaf19, negatively regulates tolerance to drought and salt and positively regulates resistance to cold stress by modulating reactive oxygen species in cotton[J]. Plant Science, 2016a, 252:267-281.

[99] Jia W, Li B, Li S, et al. Mitogen-activated protein ki nase cascade MKK7-MPK6 plays important roles in plant development and regulates shoot branching by phosphor ylating PIN1 in Arabidopsis[J]. PLOS Biol, 2016b,14(9):e1002550.

[100] Jiang Y, Han B, Zhang H, et al. MAP4K4 associates with BIK1 to regulate plant innate immunity [J]. EMBO Reports, 2019, 20(11): e47965.

[101] Jiao Q, Chen T, Niu G, et al. N-glycosylation is involved in stomatal development by modulating the release of active abscisic acid and auxin in Arabidopsis[J]. Journal of Experimental Botany, 2020, 71(19):5865-5879.

[102] Jin H, Axtell M J, Dahlbeck D, et al. NPK1, an MEKK1-like mitogen-activated protein kinase kinase kinase, regulates innate immunity and development in plants[J]. Developmental Cell, 2002, 3(2):291-297.

[103] Jonak C, Okrész L, Bögre L, et al. Complexity, cross talk and integration of plant MAP kinase signalling[J]. Current Opinion in Plant Biology, 2002, 5(5):415-424.

[104] Jouannic S, Hamal A, Leprince A S, et al. Characterisation of novel plant genes encoding MEKK/STE11 and RAF-related protein kinases[J]. Gene, 1999, 229(1-2):171-181.

[105] Kang J, Peng Y, Xu W. Crop root responses to drought stress: Molecular mechanisms, nutrient regulations, and interactions with microorganisms in the rhizosphere[J]. International Journal of Molecular Sciences, 2022 , 23(16):9310.

[106] Kim J A, Cho K, Singh R, et al. Rice OsACDR1 (Oryza sativa accelerated cell death and resistance 1) is a potential positive regulator of fungal disease resistance[J]. Molecular Cells, 2009, 28(5):431-439.

[107] Kim J M, Woo D H, Kim S H, et al. Arabidopsis MKKK20 is involved in osmotic stress response via regulation of MPK6 activity[J]. Plant Cell Reports, 2012, 31(1):217-224.

[108] Kim S H, Woo D H, Kim J M, et al. Arabidopsis MKK4 mediates osmotic-stress response via its regulation of MPK3 activity[J]. Biochemical and Biophysical Research Communications, 2011, 412(1):150-154.

[109] Kim T H, Böhmer M, Hu H H, et al. Guard cell signal transduction network: advances in understanding abscisic acid, CO2, and Ca2+ signaling[J]. Annual Review of Plant Biology, 2010, 61:561-591.

[110] Kobayashi Y, Yamamoto S, Minami H, et al. Differential activation of the rice sucrose nonfermenting1-related protein kinase2 family by hyperosmotic stress and abscisic acid[J]. Plant Cell, 2004, 16(5):1163-1177.

[111] Komatsu K, Suzuki N, Kuwamura M, et al. Group A PP2Cs evolved in landplants as key regulators of intrinsic desiccation tolerance[J]. Nature Communications, 2013, 4:2219.

[112] Kong Q, Qu N, Gao M, et al. The MEKK1-MKK1/MKK2-MPK4 kinase cascade negatively regulates immunity mediated by a mitogen-activated protein kinase kinase kinase in Arabidopsis[J]. Plant Cell, 2012, 24(5):2225-2236.

[113] Kong X, Lv W, Zhang D, et al. Genome-wide identification and analysis of expression profiles of maize mitogen-activated protein kinase kinase kinase[J]. PLoS One, 2013, 8(2):e57714.

[114] Kong X, Pan J, Zhang M, et al. ZmMKK4, a novel group C mitogen-activated protein kinase kinase in maize (Zea mays), confers salt and cold tolerance in transgenic Arabidopsis[J]. Plant Cell and Environment, 2011, 34(8):1291-1303.

[115] Krysan P J, Colcombet J. Cellular complexity in MAPK signaling in plants: Questions and emerging tools to answer them[J]. Frontiers in Plant Science, 2018, 9:1674.

[116] Kumar K, Raina S K, Sultan S M. Arabidopsis MAPK signaling pathways and their cross talks in abiotic stress response[J]. Journal of Plant Biochemistry and Biotechnology, 2020, 29:700-714.

[117] Kumar K, Rao K P, Sharma P, et al. Differential regulation of rice mitogen activated protein kinase kinase (MKK) by abiotic stress[J]. Plant Physiology and Biochemistry, 2008, 46(10):891-897.

[118] Lampard G R, Wengier D L, Bergmann D C. Manipulation of mitogen-activated protein kinase kinase signaling in the Arabidopsis stomatal lineage reveals motifs that contribute to protein localization and signaling specificity[J]. Plant Cell, 2014, 26(8):3358-3371.

[119] Leberer E, Dignard D, Harcus D, et al. The protein kinase homologue Ste20p is required to link the yeast pheromone response G-protein beta gamma subunits to downstream signalling components[J]. EMBO Journal, 1992, 11(13):4815-4824.

[120] Leberer E, Thomas D Y, Whiteway M. Pheromone signalling and polarized morphogenesis in yeast[J]. Current Opinion in Genetics and Development, 1997, 7(1):59-66.

[121] Lee D, Redfern O, Orengo C. Predicting protein function from sequence and structure[J]. Nature Reviews Molecular Cell Biology, 2007, 8(12):995-1005.

[122] Lee S J, Lee M H, Kim J I, et al. Arabidopsis putative MAP kinase kinase kinases Raf10 and Raf11 are positive regulators of seed dormancy and ABA response[J]. Plant and Cell Physiology, 2015, 56(1):84-97.

[123] Lee S K, Kim B G, Kwon T R, et al. Overexpression of the mitogen-activated protein kinase gene OsMAPK33 enhances sensitivity to salt stress in rice (Oryza sativa L.)[J]. The Journal of Bioscience and Bioengineering, 2011, 36(1):139-151.

[124] Lefebvre V, North H, Frey A, et al. Functional analysis of Arabidopsis NCED6 and NCED9 genes indicates that ABA synthesized in the endosperm is involved in the induction of seed dormancy[J]. Plant Journal, 2006, 45(3):309-319.

[125] Leprince A, Jouannic S, Hamal A, et al. Molecular characterisation of plant cDNAs BnMAP4Kalpha1 and BnMAP4Kalpha2 belonging to the GCK/SPS1 subfamily of MAP kinase kinase kinase kinase[J]. Biochimica et Biophysica Acta-General Subjects, 1999, 1444(1):1-13.

[126] Li C H, Wang G, Zhao J L, et al. The receptor-like kinase SIT1 mediates salt sensitivity by activating MAPK3/6 and regulating ethylene homeostasis in rice[J]. Plant Cell, 2014, 26(6):2538-2553.

[127] Li K, Yang F, Zhang G, et al. AIK1, A mitogen-activated protein kinase, modulates abscisic acid responses through the MKK5-MPK6 kinase cascade[J]. Plant Physiology, 2017a, 173(2):1391-1408.

[128] Li S P, Xiang X Q, Diao Z J, et al. The OsBSK1-2-MAPK module regulates blast resistance in rice[J]. The Crop Journal, 2024a, 12:110-120.

[129] Li W, Wang L, Sheng X, et al. Molecular basis for the selective and ABA-independent inhibition of PP2CA by PYL13[J]. Cell Research, 2013, 23(12):1369-1379.

[130] Li X P, Xu Q Q, Liao W B, et al. Hydrogen peroxide is involved in abscisic acid-induced adventitious rooting in cucumber (Cucumis sativus L.) under drought stress[J]. Journal of Plant Biology, 2016a, 59:536-548.

[131] Li X, Chen L, Forde B G, et al. The biphasic root growth response to abscisic acid in Arabidopsis involves interaction with ethylene and auxin signalling pathways[J]. Frontiers in Plant Science, 2017b, 8:1493.

[132] Li X, Yu B, Wu Q, et al. OsMADS23 phosphorylated by SAPK9 confers drought and salt tolerance by regulating ABA biosynthesis in rice[J]. PLoS Genetics, 2021, 17(8):e1009699.

[133] Li Y, Cai H, Liu P, et al. Arabidopsis MAPKKK18 positively regulates drought stress resistance via downstream MAPKK3[J]. Biochemical and Biophysical Research Communications, 2017c, 484(2):292-297.

[134] Li Y, Chang Y, Zhao C, et al. Expression of the inactive ZmMEK1 induces salicylic acid accumulation and salicylic acid-dependent leaf senescence[J]. Journal of Integrative Plant Biology, 2016b, 58(8):724-736.

[135] Li Y, Li Y, Zou X, et al. Bioinformatic identification and expression analyses of the MAPK-MAP4K gene family reveal a putative functional MAP4K10-MAP3K7/8-MAP2K1/11-MAPK3/6 cascade in wheat (Triticum aestivum L.)[J]. Plants (Basel), 2024b, 13(7):941.

[136] Li Y, Wang Y, Tan S, et al. Root growth adaptation is mediated by PYLs ABA receptor-PP2A protein phosphatase complex[J]. Advanced Science, 2019, 7(3):1901455.

[137] Lim C W, Baek W, Jung J, et al. Function of ABA in stomatal defense against biotic and drought stresses[J]. International Journal of Molecular Sciences, 2015, 16(7):15251-15270.

[138] Lin Z, Li Y, Wang Y, et al. Initiation and amplification of SnRK2 activation in abscisic acid signaling[J]. Nature Communications, 2021, 12(1):2456.

[139] Lin Z, Li Y, Zhang Z, et al. A RAF-SnRK2 kinase cascade mediates early osmotic stress signaling in higher plants[J]. Nature Communications, 2020, 11(1):613.

[140] Liu J, Sun X, Liao W, et al. Involvement of OsGF14b adaptation in the drought resistance of rice plants[J]. Rice (N Y), 2019, 12(1):82.

[141] Liu Y, Leary E, Saffaf O, et al. Overlapping functions of YDA and MAPKKK3/MAPKKK5 upstream of MPK3/MPK6 in plant immunity and growth/development[J]. Journal of Integrative Plant Biology, 2022, 64(8):1531-1542.

[142] Liu Y, Zhang S. Phosphorylation of 1-aminocyclopropane-1-carboxylic acid synthase by MPK6, a stress-responsive mitogen-activated protein kinase, induces ethylene biosynthesis in Arabidopsis[J]. Plant Cell, 2004, 16(12):3386-3399.

[143] Liu Y, Zhou J. MAPping kinase regulation of ICE1 in freezing tolerance[J]. Trends in Plant Science, 2018, 23(2):91-93.

[144] Liu Y. Roles of mitogen-activated protein kinase cascades in ABA signaling[J]. Plant Cell Reports, 2012, 31(1):1-12.

[145] Liu Y; Zhu Y; Xu X; et al. OstMAPKKK5, a truncated mitogen-activated protein kinase kinase kinase 5, positively regulates plant height and yield in rice[J].Crop Journal, 2019,7(5):707-714.

[146] Llompart B, Castells E, Río A, et al. The direct activation of MIK, a germinal center kinase (GCK)-like kinase, by MARK, a maize atypical receptor kinase, suggests a new mechanism for signaling through kinase-dead receptors[J]. Journal of Biological Chemistry, 2003, 278(48):48105-48111.

[147] Lu C, Chen M X, Liu R, et al. Abscisic acid regulates auxin distribution to mediate maize lateral root development under salt stress[J]. Frontiers in Plant Science, 2019, 10:716.

[148] Ma H, Chen J, Zhang Z, et al. MAPK kinase 10.2 promotes disease resistance and drought tolerance by activating different MAPKs in rice[J]. Plant Journal, 2017, 92(4):557-570.

[149] Ma Y, Szostkiewicz I, Korte A, et al. Regulators of PP2C phosphatase activity function as abscisic acid sensors[J]. Science, 2009, 324(5930):1064-1068.

[150] Machida N, Umikawa M, Takei K, et al. Mitogen-activated protein kinase kinase kinase kinase 4 as a putative effector of Rap2 to activate the c-Jun N-terminal kinase[J]. Journal of Biological Chemistry, 2004, 279(16):15711-15714.

[151] Major G, Daigle C, Stafford-Richard T, et al. Characterization of ScMAP4K1, a MAP kinase kinase kinase kinase involved in ovule, seed and fruit development in Solanum chacoense bitt[J]. Plant Biology, 2009, 10:27-46.

[152] Manna M, Rengasamy B, Sinha A K. Revisiting the role of MAPK signalling pathway in plants and its manipulation for crop improvement[J]. Plant Cell and Environment, 2023, 46(8):2277-2295.

[153] Mao X, Zhang J, Liu W, et al. The MKKK62-MKK3-MAPK7/14 module negatively regulates seed dormancy in rice[J]. Rice, 2019, 12(1):2.

[154] MAPK Group. Mitogen-activated protein kinase cascades in plants: a new nomenclature[J]. Trends in Plant Science, 2002, 7(7):301-308.

[155] Matsuoka D, Nanmori T, Sato K, et al. Activation of AtMEK1, an Arabidopsis mitogen-activated protein kinase kinase, in vitro and in vivo: analysis of active mutants expressed in E. coli and generation of the active form in stress response in seedlings[J]. Plant Journal, 2002, 29(5):637-647.

[156] Matsuoka D, Soga K, Yasufuku T, et al. Control of plant growth and development by overexpressing MAP3K17, an ABA-inducible MAP3K, in Arabidopsis[J]. Plant Biotechnology (Tokyo), 2018, 35(2):171-176.

[157] Mega R, Abe F, Kim J S, et al. Tuning water-use efficiency and drought tolerance in wheat using abscisic acid receptors[J]. Nature Plants, 2019, 5(2):153-159.

[158] Miao C, Xiao L, Hua K, et al. Mutations in a subfamily of abscisic acid receptor genes promote rice growth and productivity[J]. Proceedings of the National Academy of Sciences of the United States of America, 2018, 115(23):6058-6063.

[159] Miao Y, Laun T M, Smykowski A, et al. Arabidopsis MEKK1 can take a short cut: it can directly interact with senescence-related WRKY53 transcription factor on the protein level and can bind to its promoter[J]. Plant Molecular Biology, 2007, 65(1-2):63-76.

[160] Mitula F, Tajdel M, Cieśla A, et al. Arabidopsis ABA-activated kinase MAPKKK18 is regulated by protein phosphatase 2C ABI1 and the ubiquitin-proteasome pathway[J]. Plant and Cell Physiology, 2015, 56(12):2351-2367.

[161] Modi V, Dunbrack R L Jr. Defining a new nomenclature for the structures of active and inactive kinases[J]. PANS, 2019, 116(14):6818-6827.

[162] Montillet J L, Leonhardt N, Mondy S, et al. An abscisic acid-independent oxylipin pathway controls stomatal closure and immune defense in Arabidopsis[J]. PLoS Biology, 2013, 11(3):e1001513.

[163] Mori I C, Murata Y, Yang Y, et al. CDPKs CPK6 and CPK3 function in ABA regulation of guard cell S-type anion- and Ca2+-permeable channels and stomatal closure[J]. Public Library of Science Biology, 2006, 4(10):e327.

[164] Moustafa K, AbuQamar S, Jarrar M, et al. MAPK cascades and major abiotic stresses[J]. Plant Cell Reports, 2014, 33(8):1217-1225.

[165] Moustafa K, Lefebvre-De Vos D, Leprince A S, et al. Analysis of the Arabidopsis mitogen-activated protein kinase families: Organ specificity and transcriptional regulation upon water stresses[J]. Scholarly Research Exchange, 2008, 2008:1-12．

[166] Murata Y, Pei Z M, Mori I C, et al. Abscisic acid activation of plasma membrane Ca2+ channels in guard cells requires cytosolic NAD(P)H and is differentially disrupted upstream and downstream of reactive oxygen species production in abi1-1 and abi2-1 protein phosphatase 2C mutants[J]. Plant Cell, 2001, 13(11):2513-2523.

[167] Mustilli A C, Merlot S, Vavasseur A, et al. Arabidopsis OST1 protein kinase mediates the regulation of stomatal aperture by abscisic acid and acts upstream of reactive oxygen species production[J]. Plant Cell, 2002, 14(12):3089-3099.

[168] Na Y J, Choi H K, Park M Y, et al. OsMAPKKK63 is involved in salt stress response and seed dormancy control[J]. Plant Signaling and Behavior, 2019, 14(3):e1578633.

[169] Nagy S K, Darula Z, Kállai B M, et al. Activation of AtMPK9 through autophosphorylation that makes it independent of the canonical MAPK cascades[J]. The Biochemical Journal, 2015, 467(1):167-175.

[170] Nakagami H, Kiegerl S, Hirt H. OMTK1, a novel MAPKKK, channels oxidative stress signaling through direct MAPK interaction[J]. Journal of Biological Chemistry, 2004, 279(26):26959-26966.

[171] Nakagami H, Soukupova H, Schikora A, et al. A mitogen-activated protein kinase kinase kinase mediates reactive oxygen species homeostasis in Arabidopsis[J]. Journal of Biological Chemistry, 2006, 281(50):38697-38704.

[172] Nakashima K, Fujita Y, Kanamori N, et al. Three Arabidopsis SnRK2 protein kinases, SRK2D/SnRK2.2, SRK2E/SnRK2.6/OST1 and SRK2I/SnRK2.3, involved in ABA signaling are essential for the control of seed development and dormancy[J]. Plant and Cell Physiology, 2009, 50(7):1345-1363.

[173] Nambara E, Marion-Poll A. Abscisic acid biosynthesis and catabolism[J]. Annual Review of Plant Biology, 2005, 56:165-185.

[174] Nemoto K, Kagawa M, Nozawa A, et al. Identification of new abscisic acid receptor agonists using a wheat cell-free based drug screening system[J]. Scientific Reports, 2018, 8(1):4268.

[175] Nguyen Q T C, Lee S J, Choi S W, et al. Arabidopsis Raf-like kinase Raf10 is a regulatory component of core ABA signaling[J]. Molecules and Cells, 2019, 42(9):646-660.

[176] Ni L, Fu X, Zhang H, et al. Abscisic acid inhibits rice protein phosphatase PP45 via H2O2 and relieves repression of the Ca2+/CaM-dependent protein kinase DMI3[J]. Plant Cell, 2019, 31(1):128-152.

[177] Ning J, Li X, Hicks L M, et al. A Raf-like MAPKKK gene DSM1 mediates drought resistance through reactive oxygen species scavenging in rice[J]. Plant Physiology, 2010, 152(2):876-890.

[178] Nishimura N, Sarkeshik A, Nito K, et al. PYR/PYL/RCAR family members are major in-vivo ABI1 protein phosphatase 2C-interacting proteins in Arabidopsis[J]. Plant Journal, 2010, 61(2):290-299.

[179] Ohkuma K, Lyon J L, Addicott F T, et al. Abscisin II, an abscission-accelerating substance from young cotton fruit[J]. Science, 1963, 142:1592-1593.

[180] Pan J, Zhang M, Kong X, et al. ZmMPK17, a novel maize group D MAP kinase gene, is involved in multiple stress responses[J]. Planta, 2012, 235(4):661-676.

[181] Pan L, De Smet I. Expanding the mitogen-activated protein kinase (MAPK) universe: An update on MAP4Ks[J]. Frontiers in Plant Science, 2020, 11:1220.

[182] Pan L, Fonseca De Lima C F, Vu L D, et al. A comprehensive phylogenetic analysis of the MAP4K family in the green lineage[J]. Frontiers in Plant Science, 2021, 12:650171.

[183] Park S Y, Fung P, Nishimura N, et al. Abscisic acid inhibits type 2C protein phosphatases via the PYR/PYL family of START proteins[J]. Science, 2009, 324(5930):1068-1071.

[184] Paul A, Srinivasan N. Genome-wide and structural analyses of pseudokinases encoded in the genome of Arabidopsis thaliana provide functional insights[J]. Proteins, 2020, 88(12):1620-1638.

[185] Pawela A, Banasiak J, Biała W, et al. MtABCG20 is an ABA exporter influencing root morphology and seed germination of Medicago truncatula[J]. Plant Journal, 2019, 98(3):511-523.

[186] Pei Z M, Murata Y, Benning G, et al. Calcium channels activated by hydrogen peroxide mediate abscisic acid signalling in guard cells[J]. Nature, 2000, 406(6797):731-734.

[187] Pitzschke A, Djamei A, Bitton F, et al. A major role of the MEKK1-MKK1/2-MPK4 pathway in ROS signalling[J]. Molecular Plant, 2009, 2(1):120-137.

[188] Postiglione A E, Muday G K. The role of ROS homeostasis in ABA-induced guard cell signaling[J]. Frontiers in Plant Science, 2020, 11:968.

[189] Prince S J, Murphy M, Mutava R N, et al. Root xylem plasticity to improve water use and yield in water-stressed soybean[J]. The Journal of Experimental Botany, 2017, 68(8):2027-2036.

[190] Qi M, Elion E. A. MAP kinase pathways[J]. Journal of Cell Science, 2005, 118(Pt 16):3569-3572.

[191] Rao K P, Richa T, Kumar K, et al. In silico analysis reveals 75 members of mitogen-activated protein kinase kinase kinase gene family in rice[J]. DNA Research, 2010, 17(3):139-153.

[192] Raz V, Bergervoet J H, Koornneef M. Sequential steps for developmental arrest in Arabidopsis seeds[J]. Development, 2001, 128(2):243-252.

[193] Rentel M C, Lecourieux D, Ouaked F, et al. OXI1 kinase is necessary for oxidative burst-mediated signalling in Arabidopsis[J]. Nature, 2004, 427(6977):858-861.

[194] Ronzier E, Corratgé-Faillie C, Sanchez F, et al. CPK13, a noncanonical Ca2+-dependent protein kinase, specifically inhibits KAT2 and KAT1 shaker K+ channels and reduces stomatal opening[J]. Plant Physiology, 2014, 166(1):314-326.

[195] Rubio S, Rodrigues A, Saez A, et al. Triple loss of function of protein phosphatases type 2C leads to partial constitutive response to endogenousabscisic acid[J]. Plant Physiology, 2009, 150(3):1345-1355.

[196] Saha J, Chatterjee C, Sengupta A, et al. Genome-wide analysis and evolutionary study of sucrose non-fermenting 1-related protein kinase 2 (SnRK2) gene family members in Arabidopsis and Oryza[J]. Computational Biology and Chemistry, 2014, 49:59-70.

[197] Samakovli D, Tichá T, Vavrdová T, et al. YODA-HSP90 module regulates phosphorylation-dependent inactivation of SPEECHLESS to control stomatal development under acute heat stress in Arabidopsis[J]. Molecular Plant, 2020, 13(4):612-633.

[198] Santiago J, Rodrigues A, Saez A, et al. Modulation of drought resistance by the abscisic acid receptor PYL5 through inhibition of clade A PP2Cs[J]. Plant Journal, 2009, 60(4):575-588.

[199] Saruhashi M, Kumar Ghosh T, Arai K, et al. Plant Raf-like kinase integrates abscisic acid and hyperosmotic stress signaling upstream of SNF1-related protein kinase2[J]. Proceedings of the National Academy of Sciences of the United States of America, 2015, 112(46):E6388-E6396.

[200] Sato A, Sato Y, Fukao Y, et al. Threonine at position 306 of the KAT1 potassium channel is essential for channel activity and is a target site for ABA-activated SnRK2/OST1/SnRK2.6 protein kinase[J]. The Biochemical Journal, 2009, 424(3):439-448.

[201] Saucedo-García M, González-Córdova C D, Ponce-Pineda I G, et al. Effects of MPK3 and MPK6 kinases on the chloroplast architecture and function induced by cold acclimation in Arabidopsis[J]. Photosynth Research, 2021, 149(1-2):201-212.

[202] Schnabel J, Hombach P, Waksman T, et al. A chemical genetic approach to engineer phototropin kinases for substrate labeling[J]. Journal of Biological Chemistry, 2018, 293(15):5613-5623.

[203] Shen H, Liu C, Zhang Y, et al. OsWRKY30 is activated by MAP kinases to confer drought tolerance in rice[J]. Plant Molecular Biology, 2012, 80(3):241-253.

[204] Shen X, Liu H, Yuan B, et al. OsEDR1 negatively regulates rice bacterial resistance via activation of ethylene biosynthesis[J]. Plant Cell and Environment, 2011, 34(2):179-191.

[205] Shi B, Ni L, Liu Y, et al. OsDMI3-mediated activation of OsMPK1 regulates the activities of antioxidant enzymes in abscisic acid signalling in rice[J]. Plant Cell and Environment, 2014, 37(2):341-352.

[206] Shi J, Zhang L, An H, et al. GhMPK16, a novel stress-responsive group D MAPK gene from cotton, is involved in disease resistance and drought sensitivity[J]. BMC Molecular Biology, 2011, 12:22.

[207] Shi Z, Zhao B, Song W, et al. Genome-wide identification and characterization of the MAPKKK, MKK, and MPK families in Chinese elite maize inbred line Huangzaosi[J]. Plant Genome, 2022, 15(3):e20216.

[208] Shkolnik-Inbar D, Bar-Zvi D. ABI4 mediates abscisic acid and cytokinin inhibition of lateral root formation by reducing polar auxin transport in Arabidopsis[J]. Plant Cell, 2010, 22(11):3560-3573.

[209] Shukla V, Mattoo A K. Sucrose non-fermenting 1-related protein kinase 2 (SnRK2): a family of protein kinases involved in hyperosmotic stress signaling[J]. Physiology and Molecular Biology of Plants, 2008, 14(1-2):91-100.

[210] Sies H， Belousov V V， Chandel N S，et al. Defining roles of specific reactive oxygen species (ROS) in cell biology and physiology[J]. Nature Reviews Molecular Cell Biology, 2022, 23(7):499-515.

[211] Sies H, Jones D P. Reactive oxygen species (ROS) as pleiotropic physiological signalling agents[J]. Nature Reviews Molecular Cell Biology, 2020, 21(7):363-383.

[212] Singh A, Pandey A, Srivastava A K, et al. Plant protein phosphatases 2C: from genomic diversity to functional multiplicity and importance in stress management[J]. Critical Reviews in Biotechnology, 2016, 36(6):1023-1035.

[213] Singh R, Lee M O, Lee J E, et al. Rice mitogen-activated protein kinase interactome analysis using the yeast two-hybrid system[J]. Plant Physiology, 2012, 160(1):477-487.

[214] Siodmak A, Shahul Hameed U F, Rayapuram N, et al. Essential role of the CD docking motif of MPK4 in plant immunity, growth, and development[J]. New Phytologist, 2023, 239(3):1112-1126.

[215] Soma F, Takahashi F, Suzuki T, et al. Plant Raf-like kinases regulate the mRNA population upstream of ABA-unresponsive SnRK2 kinases under drought stress[J]. Nature Communications, 2020, 11(1):1373.

[216] Song A, Hu Y, Ding L, et al. Comprehensive analysis of mitogen-activated protein kinase cascades in chrysanthemum[J]. PeerJ, 2018, 6:e5037.

[217] Sopeña-Torres S, Jordá L, Sánchez-Rodríguez C, et al. YODA MAP3K kinase regulates plant immune responses conferring broad-spectrum disease resistance[J]. New Phytologist, 2018, 218(2):661-680.

[218] Steinberg S F. Post-translational modifications at the ATP-positioning G-loop that regulate protein kinase activity[J]. Pharmacological Research, 2018, 135:181-187.

[219] Sun T, Zhang Y. MAP kinase cascades in plant development and immune signaling[J]. EMBO Reports, 2022, 23(2):e53817.

[220] Sussmilch F C, McAdam S A M. Surviving a dry future: Abscisic acid (ABA)-mediated plant mechanisms for conserving water under low humidity[J]. Plants (Basel), 2017, 6(4):54.

[221] Takahashi Y, Soyano T, Kosetsu K, et al. HINKEL kinesin, ANP MAPKKKs and MKK6/ANQ MAPKK, which phosphorylates and activates MPK4 MAPK, constitute a pathway that is required for cytokinesis in Arabidopsis thaliana[J]. Plant and Cell Physiology. 2010, 51(10):1766-1776.

[222] Takahashi Y, Zhang J, Hsu P K, et al. MAP3Kinase-dependent SnRK2-kinase activation is required for abscisic acid signal transduction and rapid osmotic stress response[J]. Nature Communications, 2020, 11(1):12.

[223] Takemiya A, Yamauchi S, Yano T, et al. Identification of a regulatory subunit of protein phosphatase 1 which mediates blue light signaling for stomatal opening[J]. Plant and Cell Physiology, 2013, 54(1):24-35.

[224] Tang N, Zhang H, Li X, et al. Constitutive activation of transcription factor OsbZIP46 improves drought tolerance in rice[J]. Plant Physiology, 2012, 158(4):1755-1768.

[225] Tanoue T, Adachi M, Moriguchi T, et al. A conserved docking motif in MAP kinases common to substrates, activators and regulators[J]. Nature Cell Biology, 2000, 2(2):110-116.

[226] Taylor S S, Kornev A P. Protein kinases: evolution of dynamic regulatory proteins[J]. Trends in Biochemical Sciences, 2011, 36(2):65-77.

[227] Teige M, Scheikl E, Eulgem T, et al. The MKK2 pathway mediates cold and salt stress signaling in Arabidopsis[J]. Molecular Cell, 2004, 15(1):141-152.

[228] Tena G, Boudsocq M, Sheen J. Protein kinase signaling networks in plant innate immunity[J]. Current Opinion in Plant Biology, 2011, 14(5):519-529.

[229] Teng Z N, Lyu J H, Chen Y K, et al. Effects of stress-induced ABA on root architecture development: Positive and negative actions[J]. The Crop Journal, 2023, 11(4):1072-1079.

[230] Thulasi Devendrakumar K, Li X, Zhang Y. MAP kinase signalling: interplays between plant PAMP- and effector-triggered immunity[J]. Cellular And Molecular Life Sciences, 2018, 75(16):2981-2989.

[231] Topp C N. Hope in change: The role of root plasticity in crop yield stability[J]. Plant Physiology, 2016, 172(1):5-6.

[232] Ueno Y, Yoshida R, Kishi-Kaboshi M, et al. Abiotic stresses antagonize the rice defence pathway through the tyrosine-dephosphorylation of OsMPK6[J]. Public Library of Science Pathogens, 2015, 11(10):e1005231.

[233] Umezawa T, Sugiyama N, Mizoguchi M, et al. Type 2C protein phosphatases directlyregulate abscisic acid-activated protein kinases in Arabidopsis[J]. Proceedings of the National Academy of Sciences of the United States of America, 2009, 106(41):17588-17593.

[234] van Kleeff P J M, Gao J, Mol S, et al. The Arabidopsis GORK K+-channel is phosphorylated by calcium-dependent protein kinase 21 (CPK21), which in turn is activated by 14-3-3 proteins[J]. Plant Physiology and Biochemistry, 2018, 125:219-231.

[235] Virk N, Li D, Tian L, et al. Arabidopsis Raf-like mitogen-activated protein kinase kinase kinase gene Raf43 is required for tolerance to multiple abiotic stresses[J]. Public Library of Science ONE, 2015, 10(7):e0133975.

[236] Vu L D, Xu X, Zhu T, et al. The membrane-localized protein kinase MAP4K4/TOT3 regulates thermomorphogenesis[J]. Nature Communications, 2021, 12(1):2842.

[237] Wang C, Lu W, He X, et al. The cotton Mitogen-Activated Protein Kinase Kinase 3 functions in drought tolerance by regulating stomatal responses and root growth[J]. Plant and Cell Physiology, 2016, 57(8):1629-1642.

[238] Wang C, Wang G, Zhang C, et al. OsCERK1-mediated chitin perception and immune signaling requires receptor-like cytoplasmic kinase 185 to activate an MAPK cascade in rice[J]. Molecular Plant, 2017a, 10(4):619-633.

[239] Wang F, Jing W, Zhang W. The mitogen-activated protein kinase cascade MKK1-MPK4 mediates salt signaling in rice[J]. Plant Science, 2014, 227:181-189.

[240] Wang H, Ngwenyama N, Liu Y, et al. Stomatal development and patterning are regulated by environmentally responsive mitogen-activated protein kinases in Arabidopsis[J]. The Plant Cell, 2007, 19(1):63-73.

[241] Wang J, Pan C, Wang Y, et al. Genome-wide identification of MAPK, MAPKK, and MAPKKK gene families and transcriptional profiling analysis during development and stress response in cucumber[J]. BMC Genomics, 2015, 16(1):386.

[242] Wang P, Qi S, Wang X, et al. The OPEN STOMATA1-SPIRAL1 module regulates microtubule stability during abscisic acid-induced stomatal closure in Arabidopsis[J]. Plant Cell, 2023, 35(1):260-278.

[243] Wang R S, Pandey S, Li S, et al. Common and unique elements of the ABA-regulated transcriptome of Arabidopsis guard cells[J]. BMC Genomics, 2011, 12:216.

[244] Wang S H, Lim J H, Kim S S, et al. Mutation of SPOTTED LEAF3 (SPL3) impairs abscisic acid-responsive signalling and delays leaf senescence in rice[J]. Journal of Experimental Botany, 2015, 66(22):7045-7059.

[245] Wang W, Feng B, Zhou J M, et al. Plant immune signaling: Advancing on two frontiers[J]. Journal of Integrative Plant Biology, 2020, 62(1):2-24.

[246] Wang Y, Wu Y, Zhang H, et al. Arabidopsis MAPKK kinases YODA, MAPKKK3, and MAPKKK5 are functionally redundant in development and immunity[J]. Plant Physiology, 2022, 190(1):206-210.

[247] Wen Y, Li X, Guo C, et al. Characterization and expression analysis of mitogen-activated protein kinase cascade genes in wheat subjected to phosphorus and nitrogen deprivation, high salinity, and drought[J]. Journal of Plant Biochemistry and Biotechnology, 2015, 24:184-196.

[248] Wrzaczek M, Hirt H. Plant MAP kinase pathways: how many and what for?[J]. Biology of the cell, 2001, 93(1-2):81-87.

[249] Wu C, Whiteway M, Thomas D Y, et al. Molecular characterization of Ste20p, a potential mitogen-activated protein or extracellular signal-regulated kinase kinase (MEK) kinase kinase from Saccharomyces cerevisiae[J]. Journal of Biological Chemistry, 1995, 270(27):15984-15992.

[250] Wu J, Ichihashi Y, Suzuki T, et al. Abscisic acid-dependent histone demethylation during postgermination growth arrest in Arabidopsis[J]. Plant Cell and Environment. 2019, 42(7):2198-2214.

[251] Wu J, Wang J, Pan C, et al. Genome-wide identification of MAPKK and MAPKKK gene families in tomato and transcriptional profiling analysis during development and stress response[J]. Public Library of Science ONE, 2014, 9(7):e103032.

[252] Wu L, Zu X, Zhang H, et al. Overexpression of ZmMAPK1 enhances drought and heat stress in transgenic Arabidopsis thaliana[J]. Plant Molecular Biology, 2015, 88(4-5):429-443.

[253] Wu Q, Liu Y, Xie Z, et al. OsNAC016 regulates plant architecture and drought tolerance by interacting with the kinases GSK2 and SAPK8[J]. Plant Physiology, 2022, 189(3):1296-1313.

[254] Xie C, Yang L, Gai Y. MAPKKKs in Plants: Multidimensional regulators of plant growth and stress responses[J]. International Journal of Molecular Sciences, 2023, 24(4):4117.

[255] Xie G, Kato H, Imai R. Biochemical identification of the OsMKK6-OsMPK3 signalling pathway for chilling stress tolerance in rice[J]. The Biochemical Journal, 2012, 443(1):95-102.

[256] Xie Y, Ding M, Zhang B, et al. Genome-wide characterization and expression profiling of MAPK cascade genes in Salvia miltiorrhiza reveals the function of SmMAPK3 and SmMAPK1 in secondary metabolism[J]. BMC Genomics, 2020, 21(1):630.

[257] Xin J, Li C, Ning K, et al. AtPFA-DSP3, an atypical dual-specificity protein tyrosine phosphatase, affects salt stress response by modulating MPK3 and MPK6 activity[J]. Plant Cell and Environment, 2021, 44(5):1534-1548.

[258] Xing Y, Jia W, Zhang J. AtMKK1 mediates ABA-induced CAT1 expression and H2O2 production via AtMPK6-coupled signaling in Arabidopsis[J]. Plant Journal, 2008, 54(3):440-451.

[259] Xiong J, Cui X, Yuan X, et al. The Hippo/STE20 homolog SIK1 interacts with MOB1 to regulate cell proliferation and cell expansion in Arabidopsis[J]. The Journal of Experimental Botany, 2016, 67(5):1461-1475.

[260] Xu J, Li Y, Wang Y, et al. Activation of MAPK kinase 9 induces ethylene and camalexin biosynthesis and enhances sensitivity to salt stress in Arabidopsis[J]. Journal of Biological Chemistry, 2008, 283(40):26996-27006.

[261] Xu J, Zhang S. Mitogen-activated protein kinase cascades in signaling plant growth and development[J]. Trends in Plant Science, 2015, 20(1):56-64.

[262] Xu R, Duan P, Yu H, et al. Control of grain size and weight by the OsMKKK10-OsMKK4-OsMAPK6 signal ing pathway in rice[J]. Molecular Plant, 2018, 11(6):860-873.

[263] Xu W, Jia L, Shi W, et al. Abscisic acid accumulation modulates auxin transport in the root tip to enhance proton secretion for maintaining root growth under moderate water stress[J]. New Phytologist, 2013, 197(1):139-150.

[264] Xu X, Liu H, Praat M, et al. Stomatal opening under high temperatures is controlled by the OST1-regulated TOT3-AHA1 module[J]. Nature Plants, 2025, 11(1):105-117.

[265] Yamada K, Yamaguchi K, Shirakawa T, et al. The Arabidopsis CERK1-associated kinase PBL27 connects chitin perception to MAPK activation[J]. EMBO Journal, 2016, 35(22):2468-2483.

[266] Yamada K, Yamaguchi K, Yoshimura S, et al. Conservation of chitin-induced MAPK signaling pathways in rice and Arabidopsis[J]. Plant and Cell Physiology, 2017, 58(6):993-1002.

[267] Yan H, Zhao Y, Shi H, et al. BRASSINOSTEROID-SIGNALING KINASE1 Phosphorylates MAPKKK5 to regulate immunity in Arabidopsis[J]. Plant Physiology, 2018, 176(4):2991-3002.

[268] Yan Z, Wang J, Wang F, et al. MPK3/6-induced degradation of ARR1/10/12 promotes salt tolerance in Arabidopsis[J]. EMBO Reports, 2021, 22(10):e52457.

[269] Ye W, Adachi Y, Munemasa S, et al. Open stomata 1 kinase is essential for yeast elicitor-induced stomatal closure in Arabidopsis[J]. Plant and Cell Physiology, 2015, 56(6):1239-1248.

[270] Yoo S J, Kim S H, Kim M J, et al. Involvement of the OsMKK4-OsMPK1 cascade and its downstream transcription factor OsWRKY53 in the wounding response in rice[J]. Plant Pathology Journal, 2014, 30(2):168-177.

[271] Yoshida R, Hobo T, Ichimura K, et al. ABA-activated SnRK2 protein kinase is required for dehydration stress signaling in Arabidopsis[J]. Plant and Cell Physiology, 2002, 43(12):1473-1483.

[272] Yoshida R, Umezawa T, Mizoguchi T, et al. The regulatory domain of SRK2E/OST1/SnRK2.6 interacts with ABI1 and integrates abscisic acid (ABA) and osmotic stress signals controlling stomatal closure in Arabidopsis[J]. Journal of Biological Chemistry, 2006, 281(8):5310-5318.

[273] Yoshida T, Christmann A, Yamaguchi-Shinozaki K, et al. Revisiting the basal role of ABA-roles outside of stress[J]. Trends in Plant Science, 2019, 24(7):625-635.

[274] Yu J, Kang L, Li Y, et al. RING finger protein RGLG1 and RGLG2 negatively modulate MAPKKK18 mediated drought stress tolerance in Arabidopsis[J]. Journal of Integrative Plant Biology, 2021, 63(3):484-493.

[275] Yu L, Nie J, Cao C, et al. Phosphatidic acid mediates salt stress response by regulation of MPK6 in Arabidopsis thaliana[J]. New Phytologist, 2010, 188(3):762-773.

[276] Yu M H, Zhao Z Z, He J X. Brassinosteroid signaling in plant⁻microbe interactions[J]. International Journal of Molecular Sciences, 2018, 19(12):4091.

[277] Yu X, Han J, Wang E, et al. Genome-wide identification and homoeologous expression analysis of PP2C genes in wheat (Triticum aestivum L.)[J]. Frontiers in Genetics, 2019, 10:561.

[278] Zeng Q, Peng F, Wang J, et al. Identification of the MAP4K gene family reveals GhMAP4K13 regulates drought and salt stress tolerance in cotton[J]. Physiologia Plantarum, 2025, 177(1):e70031.

[279] Zhai N, Jia H, Liu D, et al. GhMAP3K65, a Cotton Raf-like MAP3K gene, enhances susceptibility to pathogen infection and heat stress by negatively modulating growth and development in transgenic Nicotiana benthamiana[J]. International Journal of Molecular Sciences, 2017, 18(11):2462.

[280] Zhan H, Yue H, Zhao X, et al. Genome-wide identification and analysis of MAPK and MAPKK gene families in bread wheat (Triticum aestivum L.)[J]. Genes (Basel), 2017, 8(10):284.

[281] Zhang H, Han W, De Smet I, et al. ABA promotes quiescence of the quiescent centre and suppresses stem cell differentiation in the Arabidopsis primary root meristem[J]. Plant Journal, 2010, 64(5):764-774.

[282] Zhang L, Li Y, Lu W, et al. Cotton GhMKK5 affects disease resistance, induces HR-like cell death, and reduces the tolerance to salt and drought stress in transgenic Nicotiana benthamiana[J]. The Journal of Experimental Botany, 2012, 63(10):3935-3951.

[283] Zhang L, Xi D, Li S, et al. A cotton group C MAP kinase gene, GhMPK2, positively regulates salt and drought tolerance in tobacco[J]. Plant Molecular Biology, 2011, 77(1-2):17-31.

[284] Zhang M M, Wu H J, Su J B, et al. Maternal control of embryogenesis by MPK6 and its upstream MKK4/MKK5 in Arabidopsis[J]. Plant Journal, 2017a, 92(6):1005-1019.

[285] Zhang M, Chiang Y H, Toruño T Y, et al. The MAP4 kinase SIK1 ensures robust extracellular ROS burst and antibacterial immunity in plants[J]. Cell Host Microbe, 2018, 24(3):379-391.

[286] Zhang M, Zhang S. Mitogen-activated protein kinase cascades in plant signaling[J]. Journal of Integrative Plant Biology, 2022, 64(2):301-341.

[287] Zhang X, Xu X, Yu Y, et al. Integration analysis of MKK and MAPK family members highlights potential MAPK signaling modules in cotton[J]. Scientific Reports, 2016, 6:29781.

[288] Zhang Y, Wang X P, Luo Y Z, et al. OsABA8ox2, an ABA catabolic gene, suppresses root elongation of rice seedlings and contributes to drought response[J]. The Crop Journal, 2022, 8(3):480-491.

[289] Zhang Z, Li J, Li F, et al. OsMAPK3 Phosphorylates OsbHLH002/OsICE1 and inhibits its ubiquitination to activate OsTPP1 and enhances rice chilling tolerance[J]. Developmental Cell, 2017b, 43(6):731-743.e5.

[290] Zhao C, Nie H, Shen Q, et al. EDR1 physically interacts with MKK4/MKK5 and negatively regulates a MAP kinase cascade to modulate plant innate immunity[J]. PLoS Genetics, 2014a, 10(5):e1004389.

[291] Zhao C, Wang P, Si T, et al. MAP kinase cascades regulate the cold response by modulating ICE1 protein stability[J]. Developmental Cell, 2017, 43(5):618-629.

[292] Zhao Y, Chan Z, Xing L, et al. The unique mode of action of a divergent member of the ABA-receptor protein family in ABA and stress signaling[J]. Cell Research, 2013, 23(12):1380-1395.

[293] Zhao Y, Xing L, Wang X, et al. The ABA receptor PYL8 promotes lateral root growth by enhancing MYB77-dependent transcription of auxin-responsive genes[J]. Science Signaling, 2014b ,7(328):ra53.

[294] Zhao Y, Zhang Z, Gao J, et al. Arabidopsis duodecuple mutant of PYL ABA receptors reveals PYL repression of ABA-independent SnRK2 activity[J]. Cell Reports, 2018, 23(11):3340-3351.e5.

[295] Zhou C, Cai Z, Guo Y, et al. An Arabidopsis mitogen-activated protein kinase cascade, MKK9-MPK6, plays a role in leaf senescence[J]. Plant Physiology, 2009, 150(1):167-177.

[296] Zhu D, Chang Y, Pei T, et al. MAPK-like protein 1 positively regulates maize seedling drought sensitivity by suppressing ABA biosynthesis[J]. Plant Journal, 2020, 102(4):747-760.

[297] Zhu J K. Abiotic stress signaling and responses in plants[J]. Cell, 2016, 167(2):313-324.

[298] Zhu Q K, Shao Y M, Ge S T, et al. A MAPK cascade downstream of IDA-HAE/HSL2 ligand-receptor pair in lateral root emergence[J]. Nature Plants, 2019a, 5(4):414-423.

[299] Zhu W, Tan W, Li Q, et al. Genome-wide characterization and expression profiling of the MAPKKK genes in Gossypium arboreum L[J]. Genome, 2019b, 62(9):609-622.

[300] Zhu X, Zhang N, Liu X, et al. Mitogen-activated protein kinase 11 (MAPK11) maintains growth and photosynthesis of potato plant under drought condition[J]. Plant Cell Reports, 2021, 40(3):491-506.

[301] Zong W, Tang N, Yang J, et al. Feedback regulation of ABA signaling and biosynthesis by a bZIP transcription factor targets drought-resistance-related genes[J]. Plant Physiology, 2016, 171(4):2810-2825.

[302] Zou J J, Li X D, Ratnasekera D, et al. Arabidopsis CALCIUM-DEPENDENT PROTEIN KINASE8 and CATALASE3 function in abscisic acid-mediated signaling and H2O2 homeostasis in stomatal guard cells under drought stress[J]. Plant Cell, 2015, 27(5):1445-1460.

[303] Zou J J, Wei F J, Wang C, et al. Arabidopsis calcium-dependent protein kinase CPK10 functions in abscisic acid- and Ca2+-mediated stomatal regulation in response to drought stress[J]. Plant Physiology, 2010, 154(3):1232-1243.