Research Progress on Calmodulin-mediated Transcriptional Regulation Mechanisms in Plant Abiotic Stress Responses

doi:10.13560/j.cnki.biotech.bull.1985.2026-0123

Abstract

Abstract:

Plants are continuously subjected to multiple abiotic stresses throughout their whole life cycle, including drought, salinity, extreme low/high temperature, and metal toxicity, which severely constrain crop growth, development and yield formation. To adapt to variable adverse environments, plants have evolved sophisticated calcium signal transduction networks that enhance stress adaptability by dynamically regulating the expression of stress-responsive genes. Calmodulin (CaM) and calmodulin-like proteins (CMLs) serve as the core calcium signal decoders in plants. They play central regulatory roles in abiotic stress responses by binding to downstream target proteins and modulating their activities. Stress stimuli trigger specific cytosolic Ca²⁺ signatures; after perceiving these Ca²⁺ signals, CaM/CML proteins interact with diverse transcription factors such as CAMTA, CBP60, Group IId WRKY, MYB, NAC and bZIP. This interaction mediates genome-wide transcriptional reprogramming, further regulating pathways involving antioxidant systems, ion homeostasis and hormone signaling, and ultimately conferring stress tolerance in plants. This review systematically summarizes the structural and functional characteristics of CaM/CML-binding transcription factors, with an emphasis on the molecular mechanisms of CaM/CML-mediated transcriptional regulation in general stress responses, as well as under cold, heat, salt, drought, and metal ion stress conditions. We also summarize the conservation and species-specificity of CaM/CML signaling pathways under different stresses, and dissect the crosstalk regulatory networks between CaM/CML signaling and phytohormone signals including abscisic acid, salicylic acid, and jasmonic acid. In addition, we propose key scientific challenges in this research field, covering the binding specificity of CaM/CML with transcription factors, the function of liquid-liquid phase separation, and the crosstalk response to multiple combined stresses. We further explore the application potential of single-cell sequencing, multi-omics integration and CRISPR/Cas9 gene editing technology in deciphering regulatory networks and improving crop stress resistance. In-depth clarification of CaM/CML-mediated transcriptional regulatory mechanisms can provide a theoretical basis and core gene targets for molecular breeding of stress-tolerant crops, and is of great scientific value and practical significance for breeding climate-smart crops and advancing green sustainable agricultural development.

Key words: Calcium signaling, Calmodulin, Calmodulin-like proteins, stress response, transcription factor, transcriptional regulation

CHANG Yu-qing, DU Jia, GUO Hang, XU Jing, WANG Yi-fan, WANG Shang, DU Li-qun, ZENG Hou-qing. Research Progress on Calmodulin-mediated Transcriptional Regulation Mechanisms in Plant Abiotic Stress Responses[J]. Biotechnology Bulletin, 2026, 42(11): 11-23.

Figures/Tables 1

References 57

[58]	Dubos C, Stracke R, Grotewold E, et al. MYB transcription factors in Arabidopsis [J]. Trends Plant Sci, 2010, 15(10): 573-581.
[59]	Yoo JH, Park CY, Kim JC, et al. Direct interaction of a divergent CaM isoform and the transcription factor, MYB2, enhances salt tolerance in Arabidopsis [J]. J Biol Chem, 2005, 280(5): 3697-3706.
[60]	Zhang YY, Ma CY, Li XQ, et al. Wheat tae-MIR1118 constitutes a functional module with calmodulin TaCaM2-1 and MYB member TaMYB44 to modulate plant low-N stress response [J]. Plant Cell Environ, 2025, 48(3): 2178-2199.
[61]	Kim HS, Park BO, Yoo JH, et al. Identification of a calmodulin-binding NAC protein as a transcriptional repressor in Arabidopsis [J]. J Biol Chem, 2007, 282(50): 36292-36302.
[62]	Duan Y, Xu ZS, Liu H, et al. Genome-wide identification of the TGA gene family and expression analysis under drought stress in Brassica napus L. [J]. Int J Mol Sci, 2024, 25(12): 6355.
[63]	Szymanski DB, Liao BR, Zielinski RE. Calmodulin isoforms differentially enhance the binding of cauliflower nuclear proteins and recombinant TGA3 to a region derived from the Arabidopsis Cam-3 promoter [J]. Plant Cell, 1996, 8(6): 1069.
[64]	Fang HH, Liu ZQ, Long YP, et al. The Ca²⁺/calmodulin2-binding transcription factor TGA3 elevates LCD expression and H₂S production to bolster Cr⁶⁺ tolerance in Arabidopsis [J]. Plant J, 2017, 91(6): 1038-1050.
[65]	Popescu SC, Popescu GV, Bachan S, et al. Differential binding of calmodulin-related proteins to their targets revealed through high-density Arabidopsis protein microarrays [J]. Proc Natl Acad Sci U S A, 2007, 104(11): 4730-4735.
[66]	Bjornson M, Dandekar A, Dehesh K. Determinants of timing and amplitude in the plant general stress response [J]. J Integr Plant Biol, 2016, 58(2): 119-126.
[67]	Walley JW, Coughlan S, Hudson ME, et al. Mechanical stress induces biotic and abiotic stress responses via a novel cis-element [J]. PLoS Genet, 2007, 3(10): e172.
[68]	Benn G, Wang CQ, Hicks DR, et al. A key general stress response motif is regulated non-uniformly by CAMTA transcription factors [J]. Plant J, 2014, 80(1): 82-92.
[69]	Bjornson M, Benn G, Song XS, et al. Distinct roles for mitogen-activated protein kinase signaling and CALMODULIN-BINDING TRANSCRIPTIONAL ACTIVATOR3 in regulating the peak time and amplitude of the plant general stress response [J]. Plant Physiol, 2014, 166(2): 988-996.
[70]	Benn G, Bjornson M, Ke HY, et al. Plastidial metabolite MEcPP induces a transcriptionally centered stress-response hub via the transcription factor CAMTA3 [J]. Proc Natl Acad Sci U S A, 2016, 113(31): 8855-8860.
[71]	张晓燕, 杨淑华, 丁杨林. 植物感知和传递低温信号的分子机制 [J]. 生物技术通报, 2023, 39(11): 28-35.
	Zhang XY, Yang SH, Ding YL. Molecular mechanism of cold signal perception and transduction in plants [J]. Biotechnol Bull, 2023, 39(11): 28-35.
[72]	Doherty CJ, Van Buskirk HA, Myers SJ, et al. Roles for Arabidopsis CAMTA transcription factors in cold-regulated gene expression and freezing tolerance [J]. Plant Cell, 2009, 21(3): 972-984.
[73]	Kidokoro S, Yoneda K, Takasaki H, et al. Different cold-signaling pathways function in the responses to rapid and gradual decreases in temperature [J]. Plant Cell, 2017, 29(4): 760-774.
[74]	Cai P, Lan YH, Gong FY, et al. Identification and molecular characterization of the CAMTA gene family in Solanaceae with a focus on the expression analysis of eggplant genes under cold stress [J]. Int J Mol Sci, 2024, 25(4): 2064.
[75]	Xi J, Qiu YJ, Du LQ, et al. Plant-specific trihelix transcription factor AtGT2L interacts with calcium/calmodulin and responds to cold and salt stresses [J]. Plant Sci, 2012, 185: 274-280.
[76]	Chu MX, Li JJ, Zhang JY, et al. AtCaM4 interacts with a Sec14-like protein, PATL1, to regulate freezing tolerance in Arabidopsis in a CBF-independent manner [J]. J Exp Bot, 2018, 69(21): 5241-5253.
[77]	Yang TB, Chaudhuri S, Yang LH, et al. A calcium/calmodulin-regulated member of the receptor-like kinase family confers cold tolerance in plants [J]. J Biol Chem, 2010, 285(10): 7119-7126.
[78]	Li H, Ding YL, Shi YT, et al. MPK3- and MPK6-mediated ICE1 phosphorylation negatively regulates ICE1 stability and freezing tolerance in Arabidopsis [J]. Dev Cell, 2017, 43(5): 630-642.e4.
[79]	Lin R, Song JN, Tang MJ, et al. CALMODULIN6 negatively regulates cold tolerance by attenuating ICE1-dependent stress responses in tomato [J]. Plant Physiol, 2023, 193(3): 2105-2121.
[80]	Zhang W, Zhou RG, Gao YJ, et al. Molecular and genetic evidence for the key role of AtCaM3 in heat-shock signal transduction in Arabidopsis [J]. Plant Physiol, 2009, 149(4): 1773-1784.
[81]	Xuan Y, Zhou S, Wang L, et al. Nitric oxide functions as a signal and acts upstream of AtCaM3 in thermotolerance in Arabidopsis seedlings [J]. Plant Physiol, 2010, 153(4): 1895-1906.
[82]	Wu HC, Luo DL, Vignols F, et al. Heat shock-induced biphasic Ca²⁺ signature and OsCaM1-1 nuclear localization mediate downstream signalling in acquisition of thermotolerance in rice (Oryza sativa L.) [J]. Plant Cell Environ, 2012, 35(9): 1543-1557.
[83]	Kutuzov MA, Bennett N, Andreeva AV. Interaction of plant protein Ser/thr phosphatase PP7 with calmodulin [J]. Biochem Biophys Res Commun, 2001, 289(2): 634-640.
[84]	Liu HT, Li GL, Chang H, et al. Calmodulin-binding protein phosphatase PP7 is involved in thermotolerance in Arabidopsis [J]. Plant Cell Environ, 2007, 30(2): 156-164.
[85]	Liu HT, Gao F, Li GL, et al. The calmodulin-binding protein kinase 3 is part of heat-shock signal transduction in Arabidopsis thaliana [J]. Plant J, 2008, 55(5): 760-773.
[86]	Kan Y, Mu XR, Zhang H, et al. TT2 controls rice thermotolerance through SCT1-dependent alteration of wax biosynthesis [J]. Nat Plants, 2022, 8(1): 53-67.
[87]	Prasad KVSK, Abdel-Hameed AAE, Jiang QY, et al. DNA-binding activity of CAMTA3 is essential for its function: identification of critical amino acids for its transcriptional activity [J]. Cells, 2023, 12(15): 1986.
[88]	Abdel-Hameed AAE, Prasad KVSK, Jiang QY, et al. Salt-induced stability of SR1/CAMTA3 mRNA is mediated by reactive oxygen species and requires the 3' end of its open reading frame [J]. Plant Cell Physiol, 2020, 61(4): 748-760.
[89]	Shkolnik D, Finkler A, Pasmanik-Chor M, et al. Calmodulin-binding transcription activator 6: a key regulator of Na⁺ homeostasis during germination [J]. Plant Physiol, 2019, 180(2): 1101-1118.
[90]	Shen QF, Fu LB, Su TT, et al. Calmodulin HvCaM1 negatively regulates salt tolerance via modulation of HvHKT1s and HvCAMTA4 [J]. Plant Physiol, 2020, 183(4): 1650-1662.
[91]	Wang DZ, Wu X, Gao SQ, et al. Systematic analysis and identification of drought-responsive genes of the CAMTA gene family in wheat (Triticum aestivum L.) [J]. Int J Mol Sci, 2022, 23(9): 4542.
[92]	Pandey N, Ranjan A, Pant P, et al. CAMTA 1 regulates drought responses in Arabidopsis thaliana [J]. BMC Genom, 2013, 14: 216.
[93]	Zeng HQ, Wu HC, Wang GP, et al. Arabidopsis CAMTA3/SR1 is involved in drought stress tolerance and ABA signaling [J]. Plant Sci, 2022, 319: 111250.
[94]	Noman M, Jameel A, Qiang WD, et al. Overexpression of GmCAMTA12 enhanced drought tolerance in Arabidopsis and soybean [J]. Int J Mol Sci, 2019, 20(19): 4849.
[95]	Baek D, Cho HM, Cha YJ, et al. Soybean calmodulin-binding transcription activators, GmCAMTA2 and GmCAMTA8, coordinate the circadian regulation of developmental processes and drought stress responses [J]. Int J Mol Sci, 2023, 24(14): 11477.
[96]	Wang S, Ge SQ, Liu XF, et al. A regulatory network involving calmodulin controls phytosulfokine peptide processing during drought-induced flower abscission [J]. Plant Cell, 2024, 37(1): koaf013.
[97]	Wei M, Xu XM, Li CH. Identification and expression of CAMTA genes in Populus trichocarpa under biotic and abiotic stress [J]. Sci Rep, 2017, 7: 17910.
[98]	Yang LY, Zhao Y, Zhang GZ, et al. Identification of CAMTA gene family in Heimia myrtifolia and expression analysis under drought stress [J]. Plants, 2022, 11(22): 3031.
[99]	Zhu XL, Wang BQ, Wei XH, et al. Characterization of the CqCAMTA gene family reveals the role of CqCAMTA03 in drought tolerance [J]. BMC Plant Biol, 2022, 22: 428.
[100]	Yoo CY, Mano N, Finkler A, et al. A Ca²⁺/CaM-regulated transcriptional switch modulates stomatal development in response to water deficit [J]. Sci Rep, 2019, 9: 12282.
[101]	Yoo CY, Pence HE, Jin JB, et al. The Arabidopsis GTL1 transcription factor regulates water use efficiency and drought tolerance by modulating stomatal density via transrepression of SDD1 [J]. Plant Cell, 2011, 22(12): 4128-4141.
[102]	Weng H, Yoo CY, Gosney MJ, et al. Poplar GTL1 is a Ca²⁺/calmodulin-binding transcription factor that functions in plant water use efficiency and drought tolerance [J]. PLoS One, 2012, 7(3): e32925.
[103]	Wan DL, Li RL, Zou B, et al. Calmodulin-binding protein CBP60g is a positive regulator of both disease resistance and drought tolerance in Arabidopsis [J]. Plant Cell Rep, 2012, 31(7): 1269-1281.
[104]	Ali MA, Azeem F, Nawaz MA, et al. Transcription factors WRKY11 and WRKY17 are involved in abiotic stress responses in Arabidopsis [J]. J Plant Physiol, 2018, 226: 12-21.
[1]	Zhang HM, Zhu JH, Gong ZZ, et al. Abiotic stress responses in plants [J]. Nat Rev Genet, 2022, 23(2): 104-119.
[2]	Leisner CP, Potnis N, Sanz-Saez A. Crosstalk and trade-offs: Plant responses to climate change-associated abiotic and biotic stresses [J]. Plant Cell Environ, 2023, 46(10): 2946-2963.
[3]	Luan S, Wang C. Calcium signaling mechanisms across Kingdoms [J]. Annu Rev Cell Dev Biol, 2021, 37: 311-340.
[4]	Sanders D, Pelloux J, Brownlee C, et al. Calcium at the crossroads of signaling [J]. Plant Cell, 2002, 14(): S401-S417.
[5]	Zeng HQ, Zhu QQ, Yuan PG, et al. Calmodulin and calmodulin-like protein-mediated plant responses to biotic stresses [J]. Plant Cell Environ, 2023, 46(12): 3680-3703.
[6]	La Verde V, Dominici P, Astegno A. Towards understanding plant calcium signaling through calmodulin-like proteins: a biochemical and structural perspective [J]. Int J Mol Sci, 2018, 19(5): 1331.
[7]	Kim MC, Chung WS, Yun DJ, et al. Calcium and calmodulin-mediated regulation of gene expression in plants [J]. Mol Plant, 2009, 2(1): 13-21.
[8]	Zhu XY, Dunand C, Snedden W, et al. CaM and CML emergence in the green lineage [J]. Trends Plant Sci, 2015, 20(8): 483-489.
[9]	Bongkoj B, Teerapong B. Genome-wide identification and analyses of the rice calmodulin and related potential calcium sensor proteins [J]. BMC Plant Biol, 2007, 7(1): 4.
[10]	Munir S, Khan MRG, Song JW, et al. Genome-wide identification, characterization and expression analysis of calmodulin-like (CML) proteins in tomato (Solanum lycopersicum) [J]. Plant Physiol Biochem, 2016, 102: 167-179.
[11]	Gifford JL, Walsh MP, Vogel HJ. Structures and metal-ion-binding properties of the Ca²⁺-binding helix-loop-helix EF-hand motifs [J]. Biochem J, 2007, 405(2): 199-221.
[12]	Dobney S, Chiasson D, Lam P, et al. The calmodulin-related calcium sensor CML42 plays a role in trichome branching [J]. J Biol Chem, 2009, 284(46): 31647-31657.
[105]	Inostroza-Blancheteau C, Aquea F, Loyola R, et al. Molecular characterisation of a calmodulin gene, VcCaM1, that is differentially expressed under aluminium stress in highbush blueberry [J]. Plant Biol J, 2013, 15(6): 1013-1018.
[106]	Tokizawa M, Kobayashi Y, Saito T, et al. Sensitive to proton rhizotoxicity1, calmodulin binding transcription activator2, and other transcription factors are involved in ALUMINUM-activated malate TRANSPORTER1 expression [J]. Plant Physiol, 2015, 167(3): 991-1003.
[107]	Zhu X, Wang P, Bai ZM, et al. Calmodulin-like protein CML24 interacts with CAMTA2 and WRKY46 to regulate ALMT1-dependent Al resistance in Arabidopsis thaliana [J]. New Phytol, 2022, 233(6): 2471-2487.
[13]	曾后清, 张亚仙, 汪尚, 等. 植物钙/钙调素介导的信号转导系统 [J]. 植物学报, 2016, 51(5): 705-723.
	Zeng HQ, Zhang YX, Wang S, et al. Calcium/calmodulin-mediated signal transduction system in plants [J]. Chin Bull Bot, 2016, 51(5): 705-723.
[14]	Hau B, Symonds K, Teresinski H, et al. Arabidopsis calmodulin-like proteins CML13 and CML14 interact with calmodulin-binding transcriptional activators and function in salinity stress response [J]. Plant Cell Physiol, 2024, 65(2): 282-300.
[15]	Scholz SS, Vadassery J, Heyer M, et al. Mutation of the Arabidopsis calmodulin-like protein CML37 deregulates the jasmonate pathway and enhances susceptibility to herbivory [J]. Mol Plant, 2014, 7(12): 1712-1726.
[16]	Leba LJ, Cheval C, Ortiz-Martín I, et al. CML9, an Arabidopsis calmodulin-like protein, contributes to plant innate immunity through a flagellin-dependent signalling pathway [J]. Plant J, 2012, 71(6): 976-989.
[17]	Tsai YC, Koo Y, Delk NA, et al. Calmodulin-related CML24 interacts with ATG4b and affects autophagy progression in Arabidopsis [J]. Plant J, 2013, 73(2): 325-335.
[18]	Nakahara KS, Masuta C, Yamada S, et al. Tobacco calmodulin-like protein provides secondary defense by binding to and directing degradation of virus RNA silencing suppressors [J]. Proc Natl Acad Sci U S A, 2012, 109(25): 10113-10118.
[19]	Yamaguchi T, Aharon GS, Sottosanto JB, et al. Vacuolar Na⁺/H⁺ antiporter cation selectivity is regulated by calmodulin from within the vacuole in a Ca²⁺- and pH-dependent manner [J]. Proc Natl Acad Sci U S A, 2005, 102(44): 16107-16112.
[20]	Wu XM, Qiao Z, Liu HP, et al. CML20, an Arabidopsis Calmodulin-like protein, negatively regulates guard cell ABA signaling and drought stress tolerance [J]. Front Plant Sci, 2017, 8: 824.
[21]	Xu GY, Rocha PSCF, Wang ML, et al. A novel rice calmodulin-like gene, OsMSR2, enhances drought and salt tolerance and increases ABA sensitivity in Arabidopsis [J]. Planta, 2011, 234(1): 47-59.
[22]	Munir S, Liu H, Xing YL, et al. Overexpression of calmodulin-like (ShCML44) stress-responsive gene from Solanum habrochaites enhances tolerance to multiple abiotic stresses [J]. Sci Rep, 2016, 6: 31772.
[23]	Kalaipandian S, Xue GP, Rae AL, et al. Overexpression of TaCML20, a calmodulin-like gene, enhances water soluble carbohydrate accumulation and yield in wheat [J]. Physiol Plant, 2019, 165(4): 790-799.
[24]	Song XY, Li JF, Lyu ML, et al. CALMODULIN-LIKE-38 and PEP1 RECEPTOR 2 integrate nitrate and brassinosteroid signals to regulate root growth [J]. Plant Physiol, 2021, 187(3): 1779-1794.
[25]	Liu S, Zheng YY, Zhao LY, et al. CALMODULIN-LIKE16 and PIN-LIKES7a cooperatively regulate rice seedling primary root elongation under chilling [J]. Plant Physiol, 2024, 195(2): 1660-1680.
[26]	Zhu QN, Tan QQ, Gao QY, et al. Calmodulin-like protein CML15 interacts with PP2C46/65 to regulate Papaya fruit ripening via integrating calcium, ABA and ethylene signals [J]. Plant Biotechnol J, 2024, 22(6): 1703-1723.
[27]	胡雅丹, 伍国强, 刘晨, 等. MYB转录因子在调控植物响应逆境胁迫中的作用 [J]. 生物技术通报, 2024, 40(6): 5-22.
	Hu YD, Wu GQ, Liu C, et al. Roles of MYB transcription factor in regulating the responses of plants to stress [J]. Biotechnol Bull, 2024, 40(6): 5-22.
[28]	Kim J, Kim HY. Functional analysis of a calcium-binding transcription factor involved in plant salt stress signaling [J]. FEBS Lett, 2006, 580(22): 5251-5256.
[29]	Kushwaha R, Singh A, Chattopadhyay S. Calmodulin7 plays an important role as transcriptional regulator in Arabidopsis seedling development [J]. Plant Cell, 2008, 20(7): 1747-1759.
[30]	Abbas N, Maurya JP, Senapati D, et al. Arabidopsis CAM7 and HY5 physically interact and directly bind to the HY5 promoter to regulate its expression and thereby promote photomorphogenesis [J]. Plant Cell, 2014, 26(3): 1036-1052.
[31]	Kumar S, Mazumder M, Gupta N, et al. Crystal structure of Arabidopsis thaliana calmodulin7 and insight into its mode of DNA binding [J]. FEBS Lett, 2016, 590(17): 3029-3039.
[32]	Senapati D, Kushwaha R, Dutta S, et al. COP1 regulates the stability of CAM7 to promote photomorphogenic growth [J]. Plant Direct, 2019, 3(6): e00144.
[33]	Truman W, Sreekanta S, Lu Y, et al. The CALMODULIN-BINDING PROTEIN60 family includes both negative and positive regulators of plant immunity [J]. Plant Physiol, 2013, 163(4): 1741-1751.
[34]	Wang YM, Shen CB, Jiang QC, et al. Seed priming with calcium chloride enhances stress tolerance in rice seedlings [J]. Plant Sci, 2022, 323: 111381.
[35]	Yu Q, Liu YL, Sun GZ, et al. Genome-wide analysis of the soybean calmodulin-binding protein 60 family and identification of GmCBP60A-1 responses to drought and salt stresses [J]. Int J Mol Sci, 2021, 22(24): 13501.
[36]	Shen L, He J, Yang X. Genome-wide identification of calmodulin-binding protein 60 gene family and function of SmCBP60A1 in eggplant response to salt stress [J]. Sci Hortic, 2023, 322: 112448.
[37]	Zhang YX, Xu SH, Ding PT, et al. Control of salicylic acid synthesis and systemic acquired resistance by two members of a plant-specific family of transcription factors [J]. Proc Natl Acad Sci U S A, 2010, 107(42): 18220-18225.
[38]	Wang L, Tsuda K, Sato M, et al. Arabidopsis CaM binding protein CBP60g contributes to MAMP-induced SA accumulation and is involved in disease resistance against Pseudomonas syringae [J]. PLoS Pathog, 2009, 5(2): e1000301.
[39]	Takagi K, Tasaki K, Komori H, et al. Hypersensitivity-related genes HSR201 and HSR203J are regulated by calmodulin-binding protein 60-type transcription factors and required for pathogen signal-induced salicylic acid synthesis [J]. Plant Cell Physiol, 2022, 63(7): 1008-1022.
[40]	Yang TB, Poovaiah BW. A calmodulin-binding/CGCG box DNA-binding protein family involved in multiple signaling pathways in plants [J]. J Biol Chem, 2002, 277(47): 45049-45058.
[41]	Xiao PX, Feng JW, Zhu XT, et al. Evolution analyses of CAMTA transcription factor in plants and its enhancing effect on cold-tolerance [J]. Front Plant Sci, 2021, 12: 758187.
[42]	Choi MS, Kim MC, Yoo JH, et al. Isolation of a calmodulin-binding transcription factor from rice (Oryza sativa L.) [J]. J Biol Chem, 2005, 280(49): 40820-40831.
[43]	Kim Y, Park S, Gilmour SJ, et al. Roles of CAMTA transcription factors and salicylic acid in configuring the low-temperature transcriptome and freezing tolerance of Arabidopsis [J]. Plant J, 2013, 75(3): 364-376.
[44]	Li XH, Huang L, Zhang YF, et al. Tomato SR/CAMTA transcription factors SlSR1 and SlSR3L negatively regulate disease resistance response and SlSR1L positively modulates drought stress tolerance [J]. BMC Plant Biol, 2014, 14: 286.
[45]	Pant P, Iqbal Z, Pandey BK, et al. Genome-wide comparative and evolutionary analysis of Calmodulin-binding Transcription Activator (CAMTA) family in Gossypium species [J]. Sci Rep, 2018, 8: 5573.
[46]	Wang GP, Zeng HQ, Hu XY, et al. Identification and expression analyses of calmodulin-binding transcription activator genes in soybean [J]. Plant Soil, 2015, 386(1/2): 205-221.
[47]	Li J, Guo HF, Lou QJ, et al. Natural variation of indels in the CTB3 promoter confers cold tolerance in Japonica rice [J]. Nat Commun, 2025, 16: 1613.
[48]	Chao LM, Kim Y, Gilmour SJ, et al. Temperature modulation of CAMTA3 gene induction activity is mediated through the DNA binding domain [J]. Plant J, 2022, 112(1): 235-248.
[49]	Du LQ, Ali GS, Simons KA, et al. Ca²⁺/calmodulin regulates salicylic-acid-mediated plant immunity [J]. Nature, 2009, 457(7233): 1154-1158.
[50]	Rushton PJ, Somssich IE, Ringler P, et al. WRKY transcription factors [J]. Trends Plant Sci, 2010, 15(5): 247-258.
[51]	Park CY, Lee JH, Yoo JH, et al. WRKY group IId transcription factors interact with calmodulin [J]. FEBS Lett, 2005, 579(6): 1545-1550.
[52]	Ding LP, Wu Z, Teng RD, et al. LlWRKY39 is involved in thermotolerance by activating LlMBF1c and interacting with LlCaM3 in lily (Lilium longiflorum) [J]. Hortic Res, 2021, 8: 36.
[53]	Wang GW, Wang XW, Ma HP, et al. PcWRKY11, an II-d WRKY transcription factor from Polygonum cuspidatum, enhances salt tolerance in transgenic Arabidopsis thaliana [J]. Int J Mol Sci, 2022, 23(8): 4357.
[54]	Chen W, Hao WJ, Xu YX, et al. Isolation and characterization of CsWRKY7, a subgroup IId WRKY transcription factor from Camellia sinensis, linked to development in Arabidopsis [J]. Int J Mol Sci, 2019, 20(11): 2815.
[55]	Mo WP, Liu Z, Zhang XY, et al. Calmodulin-like protein condensates decode PAMP-induced nuclear calcium to activate plant immunity [J]. Cell Rep, 2026, 45(4): 117092.
[56]	Sun YD, Yu DQ. Activated expression of AtWRKY53 negatively regulates drought tolerance by mediating stomatal movement [J]. Plant Cell Rep, 2015, 34(8): 1295-1306.
[57]	Jiao CY, Li KX, Zuo YX, et al. CALMODULIN1 and WRKY53 function in plant defense by negatively regulating the jasmonic acid biosynthesis pathway in Arabidopsis [J]. Int J Mol Sci, 2022, 23(14): 7718.