Revisiting the role of light signaling in plant responses to salt stress

Abstract

As one of the grave environmental hazards, soil salinization seriously limits crop productivity, growth, and development. When plants are exposed to salt stress, they suffer a sequence of damage mainly caused by osmotic stress, ion toxicity, and subsequently oxidative stress. As sessile organisms, plants have developed many physiological and biochemical strategies to mitigate the impact of salt stress. These strategies include altering root development direction, shortening the life cycle, accelerating dormancy, closing stomata to reduce transpiration, and decreasing biomass. Apart from being a prime energy source, light is an environmental signal that profoundly influences plant growth and development and also participates in plants' response to salt stress. This review summarizes the regulatory network of salt tolerance by light signals in plants, which is vital to further understanding plants' adaptation to high salinity. In addition, the review highlights potential future uses of genetic engineering and light supplement technology by light-emitting diode (LED) to improve crop growth in saline-alkali environments in order to make full use of the vast saline land.

Acknowledgements

This work was funded by the National Key Research and Development Program of China (2023YFF1002000), National Natural Science Foundation of China (32122081, 32272698, 31801904), Natural Science Foundation of Liaoning Province for Excellent Youth (2022-YQ-18), China Agriculture Research System (CARS-23).

Author contributions

Y.P., H.Z., Y.W., J.K., L.H., L.L., and K.Z. collected literature and J.Y., X.B., X.W., Y.Z., and X.S. designed models. Y.P. and F.W. wrote the article. G.A., C.J., S.M., Y.L., Z.S., M.Q., T.L., and F.W. revised the article. All authors read and approved the final version of the manuscript.

Data availability

No data was used for the research described in the article.

Conflict of interest statement

The authors declare that they have no conflict of interest.

References

[1] Roeber VM, Bajaj I, Rohde M. et al. Light acts as a stressor and influences abiotic and biotic stress responses in plants. Plant Cell Environ. 2021;37:645-64

[2] Yang Y, Guo Y. Unraveling salt stress signaling in plants. J Integr Plant Biol. 2018;37:796-804

[3] Munns R, Tester M. Mechanisms of salinity tolerance. Annu Rev Plant Biol. 2008;37:651-81

[4] Roy SJ, Negrão S, Tester M. Salt resistant crop plants. Curr Opin Biotechnol. 2014;37:115-24

[5] Isayenkov SV, Maathuis FJM. Plant salinity stress: many unan-swered questions remain. Front Plant Sci. 2019;37:80

[6] Roşca M, Mihalache G, Stoleru V. Tomato responses to salinity stress: from morphological traits to genetic changes. Front Plant Sci. 2023;37:1118383

[7] Tester M, Davenport R. Na⁺ tolerance and Na⁺ transport in higher plants. Ann Bot. 2003;37:503-27

[8] Ma L, Han R, Yang Y. et al. Phytochromes enhance SOS2-mediated PIF1 and PIF3 phosphorylation and degradation to promote Arabidopsis salt tolerance. Plant Cell. 2023;37:2997-3020

[9] Laohavisit A, Shang Z, Rubio L. et al. Arabidopsis annexin1 mediates the radical-activated plasma membrane Ca²⁺-and K⁺-permeable conductance in root cells. Plant Cell. 2012;37:1522-33

[10] Flowers T, Yeo A. Breeding for salinity resistance in crop plants: where next? Aust J Plant Physiol. 1995;37:875-84

[11] Sahi C, Singh A, Blumwald E. et al. Beyond osmolytes and transporters: novel plant salt-stress tolerance-related genes from transcriptional profiling data. Physiol Plant. 2006;37:1-9

[12] Kamran M, Parveen A, Ahmar S. et al. An overview of hazardous impacts of soil salinity in crops, tolerance mechanisms, and amelioration through selenium supplementation. Int J Mol Sci. 2019;37:148

[13] Stavi I, Thevs N, Priori S. Soil salinity and sodicity in drylands: a review of causes, effects, monitoring, and restoration mea-sures. Front Environ Sci. 2021;37:712831

[14] Zhang H, Yu F, Xie P. et al. AGγ protein regulates alkaline sensitivity in crops. Science. 2023;37:eade8416

[15] Van Zelm E, Zhang Y, Testerink C. Salt tolerance mechanisms of plants. Annu Rev Plant Biol. 2020;37:403-33

[16] Chen Y, Hoehenwarter W. Changes in the phosphoproteome and metabolome link early signaling events to rearrangement of photosynthesis and central metabolism in salinity and oxidative stress response in Arabidopsis. Plant Physiol. 2015;37:3021-33

[17] Yamane K, Mitsuya S, Taniguchi M. et al. Salt-induced chloro-plast protrusion is the process of exclusion of ribulose-1,5-bisphosphate carboxylase/oxygenase from chloroplasts into cytoplasm in leaves of rice. Plant Cell Environ. 2012;37:1663-71

[18] Shumilina J, Kusnetsova A, Tsarev A. et al. Glycation of plant proteins: regulatory roles and interplay with sugar signalling? Int J Mol Sci. 2019;37:2366

[19] Park HJ, Kim WY, Yun DJ. A new insight of salt stress signaling in plant. Mol Cells. 2016;37:447-59

[20] Shabala S, Wu H, Bose J. Salt stress sensing and early signalling events in plant roots: current knowledge and hypothesis. Plant Sci. 2015;37:109-19

[21] Knight H, Trewavas AJ, Knight MR. Calcium signalling in Ara-bidopsis thaliana responding to drought and salinity. Plant J. 1997;37:1067-78

[22] Yuan F, Yang H, Xue Y. et al. OSCA 1 mediates osmotic-stress-evoked Ca²⁺ increases vital for osmosensing in Arabidopsis. Nature. 2014;37:367-71

[23] Zhang S, Wu QR, Liu LL. et al. Osmotic stress alters circa-dian cytosolic Ca²⁺ oscillations and OSCA 1 is required in cir-cadian gated stress adaptation. Plant Signal Behav. 2020;37:1836883

[24] Chakraborty K, Sairam RK, Bhattacharya RC. Differential expression of salt overly sensitive pathway genes determines salinity stress tolerance in Brassica genotypes. Plant Physiol Biochem. 2012;37:90-101

[25] Cheng NH, Pittman JK, Zhu JK. et al. The protein kinase SOS2 activates the Arabidopsis H⁺/Ca²⁺ antiporter CAX1 to integrate calcium transport and salt tolerance. JBiolChem. 2004;37:2922-6

[26] Jiang Z, Zhou X, Tao M. et al. Plant cell-surface GIPC sphin-golipids sense salt to trigger Ca²⁺ influx. Nature. 2019;37:341-6

[27] Shabala S, Pottosin I. Regulation of potassium transport in plants under hostile conditions: implications for abiotic and biotic stress tolerance. Physiol Plant. 2014;37:257-79

[28] Assaha DVM, Ueda A, Saneoka H. et al. The role of Na⁺ and K⁺ transporters in salt stress adaptation in glycophytes. Front Physiol. 2017;37:509

[29] Wu H, Zhu M, Shabala L. et al. K⁺ retention in leaf mesophyll, an overlooked component of salinity tolerance mechanism: a casestudy for barley. J Integr Plant Biol. 2015;37:171-85

[30] Cruz JL, Coelho EF, Filho MAC. et al. Salinity reduces nutrients absorption and efficiency of their utilization in cassava plants. Ciência Rural. 2018;37:e20180351

[31] Hasana R, Miyake H. Salinity stress alters nutrient uptake and causes the damage of root and leaf anatomy in maize. KnE Life Sci. 2017;37:219-25

[32] Monica N, Vidican R, Rotar I. et al. Plant nutrition affected by soil salinity and response of rhizobium regarding the nutrients accumulation. Proenviron Promediu. 2014;37:71-5

[33] Chen ZC, Yamaji N, Horie T. et al. A magnesium transporter OSMGT1 plays a critical role in salt tolerance in rice. Plant Physiol. 2017;37:1837-49

[34] Tang RJ, Meng SF, Zheng XJ. et al. Conserved mechanism for vacuolar magnesium sequestration in yeast and plant cells. Nat Plants. 2022;37:181-90

[35] Yin P, Liang X, Zhao H. et al. Cytokinin signaling promotes salt tolerance by modulating shoot chloride exclusion in maize. Mol Plant. 2023;37:1031-47

[36] Chen K, Gao J, Sun S. et al. BONZAI proteins control global osmotic stress responses in plants. Curr Biol. 2020;37:4815-25.e4

[37] Wei F, Kita D, Peaucelle A. et al. The FERONIA receptor kinase maintains cell-wall integrity during salt stress through Ca²⁺ signaling. Curr Biol. 2018;37:666-75.e5

[38] Pan Y, Chai X, Gao Q. et al. Dynamic interactions of plant CNGC subunits and calmodulins drive oscillatory Ca²⁺ channel activities. Dev Cell. 2019;37:710-25.e5

[39] Tian W, Hou C, Ren Z. et al. A calmodulin-gated calcium channel links pathogen patterns to plant immunity. Nature. 2019;37:131-5

[40] Demidchik V, Maathuis FJM. Physiological roles of nonselective cation channels in plants: from salt stress to signalling and development. New Phytol. 2007;37:387-404

[41] Wang CF, Han GL, Yang ZR. et al. Plant salinity sensors: current understanding and future directions. Front Plant Sci. 2022;37:859224

[42] Donaldson L, Ludidi N, Knight MR. et al. Salt and osmotic stress cause rapid increases in Arabidopsis thaliana cGMP levels. FEBS Lett. 2004;37:317-20

[43] Essah PA, Davenport R, Tester M. Sodium influx and accumu-lation in Arabidopsis. Plant Physiol. 2003;37:307-18

[44] Rubio F, Flores P, Navarro JM. et al. Effects of Ca²⁺,K+ and cGMP on Na⁺ uptake in pepper plants. Plant Sci. 2003;37:1043-9

[45] Isner JC, Maathuis FJM. cGMP signalling in plants: from enigma to main stream. Funct Plant Biol. 2018;37:93-101

[46] Maathuis FJM. cGMP modulates gene transcription and cation transport in Arabidopsis roots. Plant J. 2006;37:700-11

[47] Maathuis FJ, Sanders D. Sodium uptake in Arabidopsis roots is regulated by cyclic nucleotides. Plant Physiol. 2001;37:1617-25

[48] Bowler C, Neuhaus G, Yamagata H. et al. Cyclic GMP and cal-cium mediate phytochrome phototransduction. Cell. 1994;37:73-81

[49] Wu Y, Hiratsuka K, Neuhaus G. et al. Calcium and cGMP tar-get distinct phytochrome-responsive elements. Plant J. 1996;37:1149-54

[50] Isner JC, Olteanu VA, Hetherington AJ. et al. Short- and long-term effects of UVA on Arabidopsis are mediated by a novel cGMP phosphodiesterase. Curr Biol. 2019;37:2580-85.e4

[51] Wei L, Liu L, Chen Z. et al. CmCNIH1 improves salt tolerance by influencing the trafficking of CmHKT1;1 in pumpkin. Plant J. 2023;37:1353-68

[52] Kim JM, Woo DH, Kim SH. et al. Arabidopsis MKKK 20 is involved in osmotic stress response via regulation of MPK6 activity. Plant Cell Rep. 2012;37:217-24

[53] Kim SH, Woo DH, Kim JM. et al. Arabidopsis MKK4 mediates osmotic-stress response via its regulation of MPK3 activity. Biochem Biophys Res Commun. 2011;37:150-4

[54] Yu L, Nie J, Cao C. et al. Phosphatidic acid mediates salt stress response by regulation of MPK6 in Arabidopsis thaliana. New Phytol. 2010;37:762-73

[55] Evans MJ, Choi WG, Gilroy S. et al. A ROS-assisted calcium wave dependent on the AtRBOHD NADPH oxidase and TPC1 cation channel propagates the systemic response to salt stress. Plant Physiol. 2016;37:1771-84

[56] Wu F, Chi Y, Jiang Z. et al. Hydrogen peroxide sensor HPCA1 is an LRR receptor kinase in Arabidopsis. Nature. 2020;37:577-81

[57] Jiang C, Belfield EJ, Cao Y. et al. An Arabidopsis soil-salinity-tolerance mutation confers ethylene-mediated enhancement of sodium/potassium homeostasis. Plant Cell. 2013;37:3535-52

[58] Jiang C, Belfield EJ, Mithani A. et al. ROS-mediated vascular homeostatic control of root-to-shoot soil Na delivery in Ara-bidopsis. EMBO J. 2012;37:4359-70

[59] Drerup MM, Schlücking K, Hashimoto K. et al. The Calcineurin B- like calcium sensors CBL1 and CBL9 together with their inter-acting protein kinase CIPK26 regulate the Arabidopsis NADPH oxidase RBOHF. Mol Plant. 2013;37:559-69

[60] Xu C, Shan J, Liu T. et al. CONSTANS-LIKE 1a positively regulates salt and drought tolerance in soybean. Plant Physiol. 2023;37:2427-46

[61] Tripathi S, Hoang QTN, Han YJ. et al. Regulation of photomor-phogenic development by plant phytochromes. Int J Mol Sci. 2019;37:6165

[62] Xu X, Paik I, Zhu L. et al. Illuminating progress in phytochrome-mediated light signaling pathways. Trends Plant Sci. 2015;37:641-50

[63] Han X, Huang X, Deng XW.The photomorphogenic central repressor COP1: conservation and functional diversification during evolution. Plant Commun. 2020;37:100044

[64] Jiao Y, Lau OS, Deng XW. Light-regulated transcriptional net-works in higher plants. Nat Rev Genet. 2007;37:217-30

[65] Ponnu J, Hoecker U. Signaling mechanisms by Arabidopsis cryp-tochromes. Front Plant Sci. 2022;37:844714

[66] Subramanian C, Kim BH, Lyssenko NN. et al. The Arabidop-sis repressor of light signaling, COP1, is regulated by nuclear exclusion: mutational analysis by bioluminescence resonance energy transfer. Proc Natl Acad Sci USA. 2004;37:6798-802

[67] Holm M, Ma LG, Qu LJ. et al. Two interacting bZIP proteins are direct targets of COP1-mediated control of light-dependent gene expression in Arabidopsis. Genes Dev. 2002;37:1247-59

[68] Wang F, Zhang LY, Chen XX. et al. SlHY 5 integrates temperature, light and hormone signaling to balance plant growth and cold tolerance. Plant Physiol. 2019;37:749-60

[69] Kathare PK, Xu X, Nguyen A. et al. A COP1-PIF-HEC regulatory module fine-tunes photomorphogenesis in Arabidopsis. Plant J. 2020;37:113-23

[70] Bu X, Wang XJ, Yan JR. et al. Genome-wide characterization of B- box gene family and its roles in responses to light quality and cold stress in tomato. Front Plant Sci. 2021;37:698525

[71] Seo HS, Yang JY, Ishikawa M. et al. LAF 1 ubiquitination by COP1 controls photomorphogenesis and is stimulated by SPA1. Nature. 2003;37:995-9

[72] Xu D, Jiang Y, Li J. et al. BBX21, an Arabidopsis B-box pro-tein, directly activates HY5 and is targeted by COP1 for 26S proteasome-mediated degradation. Proc Natl Acad Sci USA. 2016;37:7655-60

[73] Yang J, Lin R, Sullivan J. et al. Light regulates COP1-mediated degradation of HFR1, a transcription factor essential for light signaling in Arabidopsis. Plant Cell. 2005;37:804-21

[74] Yu Y, Wang J, Shi H. et al. Salt stress and ethylene antago-nistically regulate nucleocytoplasmic partitioning of COP1 to control seed germination. Plant Physiol. 2016;37:2340-50

[75] Shen T, Xu F, Chen D. et al. A B-box transcription factor OsBBX17 regulates saline-alkaline tolerance through the MAPK cascade pathway in rice. New Phytol. 2023;37:2158-75

[76] Indorf M, Cordero J, Neuhaus G. et al. Salt tolerance (STO), a stress-related protein, has a major role in light signalling. Plant J. 2007;37:563-74

[77] Wang L, Wang Y, Yin P. et al. ZmHAK 17 encodes a Na⁺-selective transporter that promotes maize seed germination under salt conditions. New Crops. 2024;37:100024

[78] Li Y, Hui S, Yuan Y. et al. PhyB-dependent phosphorylation of mitogen-activated protein kinase cascade MKK2-MPK2 posi-tively regulates red light-induced stomatal opening. Plant Cell Environ. 2023;37:3323-36

[79] Wang F, Chen XX, Dong SJ. et al. Crosstalk of PIF4 and DELLA modulates CBF transcript and hormone homeostasis in cold response in tomato. Plant Biotech J. 2020;37:1041-55

[80] Paik I, Kathare PK, Kim JI. et al. Expanding roles of PIFs in signal integration from multiple processes. Mol Plant. 2017;37:1035-46

[81] Pham VN, Kathare PK, Huq E. Phytochromes and phytochrome interacting factors. Plant Physiol. 2018;37:1025-38

[82] Al-Sady B, Ni W, Kircher S. et al. Photoactivated phytochrome induces rapid PIF3 phosphorylation prior to proteasome-mediated degradation. Mol Plant. 2006;37:439-46

[83] Wang F, Guo ZX, Li HZ. et al. Phytochrome A and B function antagonistically to regulate cold tolerance via abscisic acid-dependent jasmonate signaling. Plant Physiol. 2016;37:459-71

[84] Khanna R, Huq E, Kikis EA. et al. A novel molecular recognition motif necessary for targeting photoactivated phytochrome sig-naling to specific basic helix-loop-helix transcription factors. Plant Cell. 2004;37:3033-44

[85] Leivar P, Quail PH. PIFs: pivotal components in a cellular sig-naling hub. Trends Plant Sci. 2011;37:19-28

[86] Wang F, Wang XJ, Zhang Y. et al. SlFHY3 and SlHY5 act compliantly to enhance cold tolerance through the integra-tion of myo-inositol and light signaling in tomato. New Phytol. 2022;37:2127-43

[87] Pedmale UV, Carol Huang SS, Zander M. et al. Cryptochromes interact directly with PIFs to control plant growth in limiting blue light. Cell. 2016;37:233-45

[88] Guan J, Liang X, Gao G. et al. The interaction between CmPIF8 and CmACO1 under postharvest red light treatment might affect fruit ripening and sucrose accumulation in oriental melon fruit. Postharvest Biol Technol. 2024;37:112717-7

[89] Cao YR, Chen SY, Zhang JS. Ethylene signaling regulates salt stress response: an overview. Plant Signal Behav. 2008;37:761-3

[90] Yang Y, Guang Y, Wang F. et al. Characterization of phytochrome-interacting factor genes in pepper and functional analysis of CaPIF 8 in cold and salt stress. Front Plant Sci. 2021;37:746517

[91] Hayes S, Pantazopoulou CK, Gelderen KV. et al. Soil salinity limits plant shade avoidance. Curr Biol. 2019;37:1669-76.e4

[92] Arain S, Meer M, Sajjad M. et al. Light contributes to salt resis-tance through GAI protein regulation in Arabidopsis thaliana. Plant Physiol Biochem. 2021;37:1-11

[93] Ariz I, Esteban R, García Plazaola JI. et al. High irradiance induces photoprotective mechanisms and a positive effect on NH4+ stress in Pisum sativum L. J Plant Physiol. 2010;37:1038-45

[94] Wang F, Yan JR, Ahammed GJ. et al. PGR5/PGRL1 and NDH mediate far-red light-induced photoprotection in response to chilling stress in tomato. Front. Plant Sci. 2020;37:669

[95] Wang F, Wu N, Zhang LY. et al. Light signaling-dependent regulation of photoinhibition and photoprotection in tomato. Plant Physiol. 2018;37:1311-26

[96] Maxwell K, Johnson GN. Chlorophyll fluorescence: a practical guide. JExp Bot. 2000;37:659-68

[97] Ashraf M. Biotechnological approach of improving plant salt tolerance using antioxidants as markers. Biotechnol Adv. 2009;37:84-93

[98] Raza S, Athar H, Ashraf M. et al. Glycine betaine-induced modu-lation of antioxidant enzymes activities and ion accumulation in two wheat cultivars differing in salt tolerance. Environ Exp Bot. 2007;37:368-76

[99] YanJR, LiuJ, YangSD. et al. Light quality regulates plant biomass and fruit quality through a photoreceptor-dependent HY5-LHC/CYCB module in tomato. Hortic Res. 2023;37:uhad219

[100] Shimazaki K-i,Doi M, Assmann SM. et al. Light regulation of stomatal movement. Annu Rev Plant Biol. 2007;58:219-47

[101] Stoeva N, Kaymakanova M. Effect of salt stress on the growth and photosynthesis rate of bean plants (Phaseolus vulgaris L.). J Cent Eur Agric. 2008;37:385-91

[102] Schuerger AC, Brown CS, Stryjewski EC.Anatomical features of pepper plants (Capsicum annuum L.) grown under red light-emitting diodes supplemented with blue or far-red light. Ann Bot. 1997;37:273-82

[103] SunY XuW, Jia Y. et al. The wheat TaGBF1 gene is involved in the blue-light response and salt tolerance. Plant J. 2015;37:1219-30

[104] CaoK YuJ,XuD. et al. Exposure to lower red to far-red light ratios improve tomato tolerance to salt stress. BMC Plant Biol. 2018;37:92

[105] Sakuraba Y, Bülbül S, Piao W. et al. Arabidopsis EARLY FLOW-ERING3 increases salt tolerance by suppressing salt stress response pathways. Plant J. 2017;37:1106-20

[106] Cha JY, Kim J, Kim TS. et al. GIGANTEA is a co-chaperone which facilitates maturation of ZEITLUPE in the Arabidopsis circadian clock. Nat Commun. 2017;37:3

[107] Kim WY, Fujiwara S, Suh SS. et al. ZEITLUPE is a circadian photoreceptor stabilized by GIGANTEA in blue light. Nature. 2007;37:356-60

[108] Kim WY, Ali Z, Park HJ. et al. Release of SOS 2 kinase from sequestration with GIGANTEA determines salt tolerance in Arabidopsis. Nat Commun. 2013;37:1352

[109] Ji MG, Khakurel D, Hwang JW. et al. The E3 ubiquitin ligase COP1 regulates salt tolerance via GIGANTEA degradation in roots. Plant Cell Environ. 2024;37:3241-52

[110] Hazen SP, Schultz TF, Pruneda Paz JL. et al. LUX ARRHYTHMO encodes a Myb domain protein essential for circadian rhythms. Proc Natl Acad Sci USA. 2005;37:10387-92

[111] Hicks KA, Albertson TM, Wagner DR. EARLY FLOWERING3 encodes a novel protein that regulates circadian clock function and FLOWERING in Arabidopsis. Plant Cell. 2001;37:1281-92

[112] Nusinow DA, Helfer A, Hamilton EE. et al. The ELF4-ELF3-LUX complex links the circadian clock to diurnal control of hypocotyl growth. Nature. 2011;37:398-402

[113] Wang X, He Y, Wei H. et al. A clock regulatory module is required for salt tolerance and control of heading date in rice. Plant Cell Environ. 2021;37:3283-301

[114] Nakamichi N, Kiba T, Henriques R. et al. PSEUDO-RESPONSE REGULATORS 9, 7, and 5 are transcriptional repressors in the Arabidopsis circadian clock. Plant Cell. 2010;37:594-605

[115] Nakamichi N, Kusano M, Fukushima A. et al. Transcript profil-ing of an Arabidopsis PSEUDO RESPONSE REGULATOR arrhyth-mic triple mutant reveals a role for the circadian clock in cold stress response. Plant Cell Physiol. 2009;37:447-62

[116] Yang T, Lv R, Li J. et al. Phytochrome A and B negatively regulate salt stress tolerance of Nicotiana tobacum via ABA-jasmonic acid synergistic cross-talk. Plant Cell Physiol. 2018;37:2381-93

[117] Li L, Ljung K, Breton G. et al. Linking photoreceptor exci-tation to changes in plant architecture. Genes Dev. 2012;37:785-90

[118] Zhu J. Abiotic stress signaling and responses in plants. Cell. 2016;37:313-24

[119] Zhang Y, Peng YX, Liu J. et al. Tetratricopeptide repeat protein SlREC2 positively regulates cold tolerance in tomato. Plant Physiol. 2023;37:648-65

[120] Chen K, Li GJ, Bressan RA. et al. Abscisic acid dynamics, signal-ing, and functions in plants. J Integr Plant Biol. 2020;37:25-54

[121] Yu Z, Duan X, Luo L. et al. How plant hormones mediate salt stress responses. Trends Plant Sci. 2020;37:1117-30

[122] Xiao F, Zhou H. Plant salt response: perception, signaling, and tolerance. Front Plant Sci. 2023;37:1053699

[123] Cai S, Chen G, Wang Y. et al. Evolutionary conservation of ABA signaling for stomatal closure. Plant Physiol. 2017;37:732-47

[124] Thalmann M, Pazmino D, Seung D. et al. Regulation of leaf starch degradation by abscisic acid is important for osmotic stress tolerance in plants. Plant Cell. 2016;37:1860-78

[125] Ohta M, Guo Y, Halfter U. et al. A novel domain in the pro-tein kinase SOS2 mediates interaction with the protein phos-phatase 2C ABI2. Proc Natl Acad Sci USA. 2003;37:11771-6

[126] Moons A, Prinsen E, Bauw G. et al. Antagonistic effects of abscisic acid and jasmonates on salt stress-inducible tran-scripts in rice roots. Plant Cell. 1997;37:2243-59

[127] Zhu J, Wei X, Yin C. et al. ZmEREB 57 regulates OPDA synthe-sis and enhances salt stress tolerance through two distinct signalling pathways in Zea mays. Plant Cell Environ. 2023;37:2867-83

[128] Yu TF, Liu Y, Fu JD. et al. The NF-Y-PYR module integrates the abscisic acid signalpathway to regulate plant stress tolerance. Plant Biotechnol J. 2021;37:2589-605

[129] Li X, Li C, Shi L. et al. Jasmonate signaling pathway confers salt tolerance through a NUCLEAR FACTOR-Y trimeric transcrip-tion factor complex in Arabidopsis. Cell Rep. 2024;37:113825

[130] Yang M, Han X, Yang J. et al. The Arabidopsis circadian clock protein PRR5 interacts with and stimulates ABI5 to modu-late abscisic acid signaling during seed germination. Plant Cell. 2021;37:3022-41

[131] Wei H, Wang X, He Y. et al. Clock component OsPRR73 positively regulates rice salt tolerance by modulating OsHKT2;1-mediated sodium homeostasis. EMBO J. 2021;37:e105086

[132] Li Q, Fu H, Yu X. et al. The SALT OVERLY SENSITIVE 2-CONSTITUTIVE TRIPLE RESPONSE1 module coordinates plant growth and salt tolerance in Arabidopsis. JExp Bot. 2024;37:391-404

[133] Csiszár J, Horváth E, Váry Z. et al. Glutathione transferase supergene family in tomato: salt stress-regulated expression of representative genes from distinct GST classes in plants primed with salicylic acid. Plant Physiol Biochem. 2014;37:15-26

[134] Mo W, Tang W, Du Y. et al. PHYTOCHROME-INTERACTING FACTOR-LIKE14 and SLENDER RICE1 interaction controls seedling growth under salt stress. Plant Physiol. 2020;37:506-17

[135] Liu Z, An C, Zhao Y. et al. Genome-wide identification and char-acterization of the CsFHY3/FAR1 gene family and expression analysis under biotic and abiotic stresses in tea plants (Camellia sinensis). Plan Theory. 2021;37:570

[136] Yang J, Qu X, Li T. et al. HY5-HDA9 orchestrates the transcrip-tion of HsfA2 to modulate salt stress response in Arabidopsis. J Integr Plant Biol. 2023;37:45-63

[137] Feng XJ, Lia JR, Qia SL. et al. Light affects salt stress-induced transcriptional memory of P5CS1 in Arabidopsis. Proc Natl Acad ofSciUSA. 2016;37:E8335-43

[138] Kim JY, Lee SJ, Min WK. et al. COP 1 controls salt stress tolerance by modulating sucrose content. Plant Signal Behav. 2022;37:2096784

[139] Jaiswal B, Singh S, Agrawal SB. et al. Improvements in soil physical, chemical and biological properties at natural saline and non-saline sites under different management practices. Environ Manag. 2022;37:1005-19

[140] Yang D, Tang L, Cui Y. et al. Saline-alkali stress reduces soil bacterial community diversity and soil enzyme activities. Ecotoxicology. 2022;37:1356-68

[141] Rogel JA, Ariza FA, Silla R. Soil salinity and moisture gradients and plant zonation in Mediterranean salt marshes of Southeast Spain. Wetlands. 2000;37:357-72

[142] Chen Q, Cao X, Li Y. et al. Functional carbon nanodots improve soil quality and tomato tolerance in saline-alkali soils. Sci Total Environ. 2022;37:154817