Regulation of Reactive Oxygen Species during Salt Stress in Plants and Their Crosstalk with Other Signaling Molecules—Current Perspectives and Future Directions

Plants - Tập 12 Số 4 - Trang 864
Mahipal Singh Kesawat1, Neela Satheesh2, Bhagwat Singh Kherawat3, Ajay Kumar4, Hyun Uk Kim5, Sang-Min Chung6, Manu Kumar6
1Department of Genetics and Plant Breeding, Faculty of Agriculture, Sri Sri University, Cuttack 754006, India
2Department of Food Nutrition and Dietetics, Faculty of Agriculture, Sri Sri University, Cuttack 754006, India
3Krishi Vigyan Kendra, Bikaner II, Swami Keshwanand Rajasthan Agricultural University, Bikaner 334603, India
4Centre of Advanced Study in Botany, Banaras Hindu University, Varanasi-221005, India
5Department of Bioindustry and Bioresource Engineering, Plant Engineering Research Institute, Sejong University, Seoul, 05006, Republic of Korea
6Department of Life Science, College of Life Science and Biotechnology, Dongguk University, Goyang 10326, Republic of Korea

Tóm tắt

Salt stress is a severe type of environmental stress. It adversely affects agricultural production worldwide. The overproduction of reactive oxygen species (ROS) is the most frequent phenomenon during salt stress. ROS are extremely reactive and, in high amounts, noxious, leading to destructive processes and causing cellular damage. However, at lower concentrations, ROS function as secondary messengers, playing a critical role as signaling molecules, ensuring regulation of growth and adjustment to multifactorial stresses. Plants contain several enzymatic and non-enzymatic antioxidants that can detoxify ROS. The production of ROS and their scavenging are important aspects of the plant’s normal response to adverse conditions. Recently, this field has attracted immense attention from plant scientists; however, ROS-induced signaling pathways during salt stress remain largely unknown. In this review, we will discuss the critical role of different antioxidants in salt stress tolerance. We also summarize the recent advances on the detrimental effects of ROS, on the antioxidant machinery scavenging ROS under salt stress, and on the crosstalk between ROS and other various signaling molecules, including nitric oxide, hydrogen sulfide, calcium, and phytohormones. Moreover, the utilization of “-omic” approaches to improve the ROS-regulating antioxidant system during the adaptation process to salt stress is also described.

Từ khóa


Tài liệu tham khảo

Kumar, 2013, Insights into genomics of salt stress response in rice, Rice, 6, 1, 10.1186/1939-8433-6-27

Hirayama, 2010, Research on plant abiotic stress responses in the post-genome era: Past, present and future, Plant J., 61, 1041, 10.1111/j.1365-313X.2010.04124.x

Kesawat, M.S., Kherawat, B.S., Singh, A., Dey, P., Kabi, M., Debnath, D., Saha, D., Khandual, A., Rout, S., and Ali, A. (2021). Genome-wide identification and characterization of the brassinazole-resistant (BZR) gene family and its expression in the various developmental stage and stress conditions in wheat (Triticum aestivum L.). Int. J. Mol. Sci., 22.

Kesawat, M.S., Kherawat, B.S., Singh, A., Dey, P., Routray, S., Mohapatra, C., Saha, D., Ram, C., Siddique, K.H., and Kumar, A. (2022). Genome-Wide Analysis and Characterization of the Proline-Rich Extensin-like Receptor Kinases (PERKs) Gene Family Reveals Their Role in Different Developmental Stages and Stress Conditions in Wheat (Triticum aestivum L.). Plants, 11.

Kumar, M., Kesawat, M.S., Ali, A., Lee, S.-C., Gill, S.S., and Kim, H.U. (2019). Integration of abscisic acid signaling with other signaling pathways in plant stress responses and development. Plants, 8.

Munns, 2008, Mechanisms of salinity tolerance, Annu. Rev. Plant Biol., 59, 651, 10.1146/annurev.arplant.59.032607.092911

Volkov, 2017, Salinity Tolerance in Plants: Mechanisms and Regulation of Ion Transport, Front. Plant Sci., 8, 1795, 10.3389/fpls.2017.01795

Ashraf, 2009, Biotechnological approach of improving plant salt tolerance using antioxidants as markers, Biotechnol. Adv., 27, 84, 10.1016/j.biotechadv.2008.09.003

Hasanuzzaman, M., Bhuyan, M.B., Zulfiqar, F., Raza, A., Mohsin, S.M., Mahmud, J.A., Fujita, M., and Fotopoulos, V. (2020). Reactive oxygen species and antioxidant defense in plants under abiotic stress: Revisiting the crucial role of a universal defense regulator. Antioxidants, 9.

Sachdev, S., Ansari, S.A., Ansari, M.I., Fujita, M., and Hasanuzzaman, M. (2021). Abiotic stress and reactive oxygen species: Generation, signaling, and defense mechanisms. Antioxidants, 10.

Singhal, 2021, Crucial cell signaling compounds crosstalk and integrative multi-omics techniques for salinity stress tolerance in plants, Front. Plant Sci., 12, 670369, 10.3389/fpls.2021.670369

Mehla, N., Sindhi, V., Josula, D., Bisht, P., and Wani, S.H. (2017). Reactive Oxygen Species and Antioxidant Systems in Plants: Role and Regulation under Abiotic Stress, Springer.

Hasanuzzaman, M., Bhuyan, M.B., Anee, T.I., Parvin, K., Nahar, K., Mahmud, J.A., and Fujita, M. (2019). Regulation of ascorbate-glutathione pathway in mitigating oxidative damage in plants under abiotic stress. Antioxidants, 8.

Gill, 2010, Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants, Plant Physiol. Biochem., 48, 909, 10.1016/j.plaphy.2010.08.016

Singh, 2019, Reactive oxygen species-mediated signaling during abiotic stress, Plant Gene, 18, 100173, 10.1016/j.plgene.2019.100173

Hasanuzzaman, M., Hossain, M.A., Silva, J.A., and Fujita, M. (2012). Crop Stress and Its Management: Perspectives and Strategies, Springer.

Raja, 2017, Abiotic stress: Interplay between ROS, hormones and MAPKs, Environ. Exp. Bot., 137, 142, 10.1016/j.envexpbot.2017.02.010

Kaur, 2019, Effect of heat stress on antioxidative defense system and its amelioration by heat acclimation and salicylic acid pre-treatments in three pigeonpea genotypes, Indian J. Agric. Biochem., 32, 106, 10.5958/0974-4479.2019.00014.5

Mittler, 2017, ROS are good, Trends Plant Sci., 22, 11, 10.1016/j.tplants.2016.08.002

Antoniou, 2016, Unravelling chemical priming machinery in plants: The role of reactive oxygen–nitrogen–sulfur species in abiotic stress tolerance enhancement, Curr. Opin. Plant Biol., 33, 101, 10.1016/j.pbi.2016.06.020

Noctor, 2016, Intracellular redox compartmentation and ROS-related communication in regulation and signaling, Plant Physiol., 171, 1581, 10.1104/pp.16.00346

Kohli, S.K., Khanna, K., Bhardwaj, R., Abd_Allah, E.F., Ahmad, P., and Corpas, F.J. (2019). Assessment of subcellular ROS and NO metabolism in higher plants: Multifunctional signaling molecules. Antioxidants, 8.

Dietz, 2016, Thiol-based peroxidases and ascorbate peroxidases: Why plants rely on multiple peroxidase systems in the photosynthesizing chloroplast?, Mol. Cells, 39, 20, 10.14348/molcells.2016.2324

Dogra, 2020, Singlet oxygen metabolism: From genesis to signaling, Front. Plant Sci., 10, 1640, 10.3389/fpls.2019.01640

Muller, 2001, Non-photochemical quenching. A response to excess light energy, Plant Physiol., 125, 1558, 10.1104/pp.125.4.1558

Li, 2009, Sensing and responding to excess light, Annu. Rev. Plant Biol., 60, 239, 10.1146/annurev.arplant.58.032806.103844

2005, Singlet oxygen production in photosynthesis, J. Exp. Bot., 56, 337

Fufezan, 2008, Singlet oxygen production in photosystem II and related protection mechanism, Photosynth. Res., 98, 551, 10.1007/s11120-008-9349-3

Flors, 2006, Imaging the production of singlet oxygen in vivo using a new fluorescent sensor, Singlet Oxygen Sensor Green®, J. Exp. Bot., 57, 1725, 10.1093/jxb/erj181

Havaux, 2009, Singlet oxygen in plants: Production, detoxification and signaling, Trends Plant Sci., 14, 219, 10.1016/j.tplants.2009.01.008

Bose, 2014, ROS homeostasis in halophytes in the context of salinity stress tolerance, J. Exp. Bot., 65, 1241, 10.1093/jxb/ert430

Kerchev, 2016, Lack of GLYCOLATE OXIDASE1, but not GLYCOLATE OXIDASE2, attenuates the photorespiratory phenotype of CATALASE2-deficient Arabidopsis, Plant Physiol., 171, 1704, 10.1104/pp.16.00359

2016, ROS generation in peroxisomes and its role in cell signaling, Plant Cell Physiol., 57, 1364

Reumann, 2016, Characterization, prediction and evolution of plant peroxisomal targeting signals type 1 (PTS1s), Biochim. Biophys. Acta BBA-Mol. Cell Res., 1863, 790, 10.1016/j.bbamcr.2016.01.001

Corpas, 2019, Plant peroxisomes at the crossroad of NO and H2O2 metabolism, J. Integr. Plant Biol., 61, 803, 10.1111/jipb.12772

Gilroy, 2016, ROS, calcium, and electric signals: Key mediators of rapid systemic signaling in plants, Plant Physiol., 171, 1606, 10.1104/pp.16.00434

Corpas, 2020, Plant peroxisomes: A factory of reactive species, Front. Plant Sci., 11, 853, 10.3389/fpls.2020.00853

Lisenbee, 2005, Arabidopsis peroxisomes possess functionally redundant membrane and matrix isoforms of monodehydroascorbate reductase, Plant J., 43, 900, 10.1111/j.1365-313X.2005.02503.x

Leterrier, 2005, Peroxisomal monodehydroascorbate reductase. Genomic clone characterization and functional analysis under environmental stress conditions, Plant Physiol., 138, 2111, 10.1104/pp.105.066225

Choudhary, 2020, ROS and oxidative burst: Roots in plant development, Plant Divers., 42, 33, 10.1016/j.pld.2019.10.002

Sharma, 2012, Reactive oxygen species, oxidative damage, and antioxidative defense mechanism in plants under stressful conditions, J. Bot., 2012, 217037

Heyno, 2011, Oxygen activation at the plasma membrane: Relation between superoxide and hydroxyl radical production by isolated membranes, Planta, 234, 35, 10.1007/s00425-011-1379-y

Banerjee, 2015, Seed birth to death: Dual functions of reactive oxygen species in seed physiology, Ann. Bot., 116, 663, 10.1093/aob/mcv098

Janků, M., Luhová, L., and Petřivalský, M. (2019). On the origin and fate of reactive oxygen species in plant cell compartments. Antioxidants, 8.

Maurya, A.K. (2020). Agronomic Crops, Springer.

Camejo, 2016, Reactive oxygen species, essential molecules, during plant–pathogen interactions, Plant Physiol. Biochem., 103, 10, 10.1016/j.plaphy.2016.02.035

Dumont, 2019, Consequences of oxidative stress on plant glycolytic and respiratory metabolism, Front. Plant Sci., 10, 166, 10.3389/fpls.2019.00166

Martinez, 2016, Accumulation of flavonols over hydroxycinnamic acids favors oxidative damage protection under abiotic stress, Front. Plant Sci., 7, 838, 10.3389/fpls.2016.00838

Karuppanapandian, 2013, Cobalt-induced oxidative stress causes growth inhibition associated with enhanced lipid peroxidation and activates antioxidant responses in Indian mustard (Brassica juncea L.) leaves, Acta Physiol. Plant, 35, 2429, 10.1007/s11738-013-1277-y

Bowler, 1992, Superoxide dismutase and stress tolerance, Annu. Rev. Plant Biol., 43, 83, 10.1146/annurev.pp.43.060192.000503

Mittler, 2002, Oxidative stress, antioxidants and stress tolerance, Trends Plant Sci., 7, 405, 10.1016/S1360-1385(02)02312-9

Dat, 1998, Changes in salicylic acid and antioxidants during induced thermotolerance in mustard seedlings, Plant Physiol., 118, 1455, 10.1104/pp.118.4.1455

Gepstein, 2013, Strategies to ameliorate abiotic stress-induced plant senescence, Plant Mol. Biol., 82, 623, 10.1007/s11103-013-0038-z

Corpas, 2006, The expression of different superoxide dismutase forms is cell-type dependent in olive (Olea europaea L.) leaves, Plant Cell Physiol., 47, 984, 10.1093/pcp/pcj071

Blokhina, 2003, Antioxidants, oxidative damage and oxygen deprivation stress: A review, Ann. Bot., 91, 179, 10.1093/aob/mcf118

Pazmino, 2009, Cellular response of pea plants to cadmium toxicity: Cross talk between reactive oxygen species, nitric oxide, and calcium, Plant Physiol., 150, 229, 10.1104/pp.108.131524

Boguszewska, 2010, Drought-responsive antioxidant enzymes in potato (Solanum tuberosum L.), Potato Res., 53, 373, 10.1007/s11540-010-9178-6

Nicholls, 2000, Enzymology and Structure of Catalases, Adv Inorg Chem, 51, 51, 10.1016/S0898-8838(00)51001-0

Gasselhuber, 2012, Molecular evolution of hydrogen peroxide degrading enzymes, Arch. Biochem. Biophys., 525, 131, 10.1016/j.abb.2012.01.017

Loewen, 2000, Catalase—An “old” enzyme that continues to surprise us, ASM News, 66, 76

Mhamdi, 2010, Arabidopsis GLUTATHIONE REDUCTASE1 plays a crucial role in leaf responses to intracellular hydrogen peroxide and in ensuring appropriate gene expression through both salicylic acid and jasmonic acid signaling pathways, Plant Physiol., 153, 1144, 10.1104/pp.110.153767

Ogren, 1984, Photorespiration: Pathways, regulation, and modification, Annu. Rev. Plant Physiol., 35, 415, 10.1146/annurev.pp.35.060184.002215

Andre, 2013, Fusing catalase to an alkane-producing enzyme maintains enzymatic activity by converting the inhibitory byproduct H2O2 to the cosubstrate O2, Proc. Natl. Acad. Sci. USA, 110, 3191, 10.1073/pnas.1218769110

Singh, 2012, The plant host Brassica napus induces in the pathogen Verticillium longisporum the expression of functional catalase peroxidase which is required for the late phase of disease, Mol. Plant Microbe Interact., 25, 569, 10.1094/MPMI-08-11-0217

Scandalios, J.G. (2019). Causes of Photooxidative Stress and Amelioration of Defense Systems in Plants, CRC Press.

Sharma, I., and Ahmad, P. (2014). Oxidative Damage to Plants, Elsevier.

Gondim, 2012, Catalase plays a key role in salt stress acclimation induced by hydrogen peroxide pretreatment in maize, Plant Physiol. Biochem., 56, 62, 10.1016/j.plaphy.2012.04.012

Kerdnaimongkol, 1999, Inhibition of catalase by antisense RNA increases susceptibility to oxidative stress and chilling injury in transgenic tomato plants, J. Am. Soc. Hortic. Sci., 124, 330, 10.21273/JASHS.124.4.330

Nakano, 1981, Hydrogen peroxide is scavenged by ascorbate-specific peroxidase in spinach chloroplasts, Plant Cell Physiol., 22, 867

Caverzan, 2012, Plant responses to stresses: Role of ascorbate peroxidase in the antioxidant protection, Genet. Mol. Biol., 35, 1011, 10.1590/S1415-47572012000600016

Pandey, 2017, Abiotic stress tolerance in plants: Myriad roles of ascorbate peroxidase, Front. Plant Sci., 8, 581, 10.3389/fpls.2017.00581

Shigeoka, 2002, Regulation and function of ascorbate peroxidase isoenzymes, J. Exp. Bot., 53, 1305, 10.1093/jexbot/53.372.1305

Pnueli, 2003, Growth suppression, altered stomatal responses, and augmented induction of heat shock proteins in cytosolic ascorbate peroxidase (Apx1)-deficient Arabidopsis plants, Plant J., 34, 187, 10.1046/j.1365-313X.2003.01715.x

Neill, 2002, Hydrogen peroxide signalling, Curr. Opin. Plant Biol., 5, 388, 10.1016/S1369-5266(02)00282-0

Vandenabeele, 2004, Catalase deficiency drastically affects gene expression induced by high light in Arabidopsis thaliana, Plant J., 39, 45, 10.1111/j.1365-313X.2004.02105.x

Chen, 2003, Increasing vitamin C content of plants through enhanced ascorbate recycling, Proc. Natl. Acad. Sci. USA, 100, 3525, 10.1073/pnas.0635176100

Mittova, 2000, Activities of SOD and the ascorbate-glutathione cycle enzymes in subcellular compartments in leaves and roots of the cultivated tomato and its wild salt-tolerant relative Lycopersicon pennellii, Physiol. Plant., 110, 42, 10.1034/j.1399-3054.2000.110106.x

Eltayeb, 2007, Overexpression of monodehydroascorbate reductase in transgenic tobacco confers enhanced tolerance to ozone, salt and polyethylene glycol stresses, Planta, 225, 1255, 10.1007/s00425-006-0417-7

Hossain, 2010, Evidence for a role of exogenous glycinebetaine and proline in antioxidant defense and methylglyoxal detoxification systems in mung bean seedlings under salt stress, Physiol. Mol. Biol. Plants, 16, 19, 10.1007/s12298-010-0003-0

Chen, 2004, The ascorbic acid redox state controls guard cell signaling and stomatal movement, Plant Cell, 16, 1143, 10.1105/tpc.021584

Noshi, 2016, Redox regulation of ascorbate and glutathione by a chloroplastic dehydroascorbate reductase is required for high-light stress tolerance in Arabidopsis, Biosci. Biotechnol. Biochem., 80, 870, 10.1080/09168451.2015.1135042

Rahantaniaina, 2017, Cytosolic and chloroplastic DHARs cooperate in oxidative stress-driven activation of the salicylic acid pathway, Plant Physiol., 174, 956, 10.1104/pp.17.00317

Ding, 2020, The pivotal function of dehydroascorbate reductase in glutathione homeostasis in plants, J. Exp. Bot., 71, 3405, 10.1093/jxb/eraa107

Shimaoka, 2000, Purification and characterization of chloroplast dehydroascorbate reductase from spinach leaves, Plant Cell Physiol., 41, 1110, 10.1093/pcp/pcd035

Qin, 2011, Ascorbic acid contents in transgenic potato plants overexpressing two dehydroascorbate reductase genes, Mol. Biol. Rep., 38, 1557, 10.1007/s11033-010-0264-2

Sanmartin, 2003, Over-expression of ascorbate oxidase in the apoplast of transgenic tobacco results in altered ascorbate and glutathione redox states and increased sensitivity to ozone, Planta, 216, 918, 10.1007/s00425-002-0944-9

Dardalhon, 2012, Redox-sensitive YFP sensors monitor dynamic nuclear and cytosolic glutathione redox changes, Free Radic. Biol. Med., 52, 2254, 10.1016/j.freeradbiomed.2012.04.004

Rao, A., and Reddy, A.R. (2008). Sulfur Assimilation and Abiotic Stress in Plants, Springer.

Bass, 2004, A major fraction of endoplasmic reticulum-located glutathione is present as mixed disulfides with protein, J. Biol. Chem., 279, 5257, 10.1074/jbc.M304951200

Raturi, 2007, Characterization of redox state and reductase activity of protein disulfide isomerase under different redox environments using a sensitive fluorescent assay, Free Radic. Biol. Med., 43, 62, 10.1016/j.freeradbiomed.2007.03.025

Kubo, 1993, Primary structure and properties of glutathione reductase from Arabidopsis thaliana, Plant Cell Physiol., 34, 1259

Creissen, 1995, Cloning and characterisation of glutathione reductase cDNAs and identification of two genes encoding the tobacco enzyme, Planta, 197, 422, 10.1007/BF00202667

Kaminaka, 1998, Gene cloning and expression of cytosolic glutathione reductase in rice (Oryza sativa L.), Plant Cell Physiol., 39, 1269, 10.1093/oxfordjournals.pcp.a029330

2006, Glutathione reductase in leaves of cowpea: Cloning of two cDNAs, expression and enzymatic activity under progressive drought stress, desiccation and abscisic acid treatment, Ann. Bot., 98, 1279, 10.1093/aob/mcl217

Yousuf, P.Y., Hakeem, K.U.R., Chandna, R., and Ahmad, P. (2012). Abiotic Stress Responses in Plants, Springer.

Ghisla, 1989, Mechanisms of flavoprotein-catalyzed reactions, Eur. J. Biochem., 181, 1, 10.1111/j.1432-1033.1989.tb14688.x

Gill, 2013, Glutathione and glutathione reductase: A boon in disguise for plant abiotic stress defense operations, Plant Physiol. Biochem., 70, 204, 10.1016/j.plaphy.2013.05.032

Mika, 2003, Properties of guaiacol peroxidase activities isolated from corn root plasma membranes, Plant Physiol., 132, 1489, 10.1104/pp.103.020396

Hiraga, 2001, A large family of class III plant peroxidases, Plant Cell Physiol., 42, 462, 10.1093/pcp/pce061

Asada, 1999, The water-water cycle in chloroplasts: Scavenging of active oxygens and dissipation of excess photons, Annu. Rev. Plant Physiol. Plant Mol. Biol., 50, 601, 10.1146/annurev.arplant.50.1.601

Xiao, 2008, Oxidative stress induced by lead in chloroplast of spinach, Biol. Trace Elem. Res., 126, 257, 10.1007/s12011-008-8195-7

Castro, D., Contreras, L.M., Kurz, L., and Wilkesman, J. (2017). Zymography, Springer.

Parvanova, 2004, Transgenic tobacco plants accumulating osmolytes show reduced oxidative damage under freezing stress, Plant Physiol. Biochem., 42, 57, 10.1016/j.plaphy.2003.10.007

2004, Changes in the ascorbate metabolism of apoplastic and symplastic spaces are associated with cell differentiation, J. Exp. Bot., 55, 2559, 10.1093/jxb/erh253

Amaya, 2012, The challenge of increasing vitamin C content in plant foods, Biotechnol. J., 7, 1110, 10.1002/biot.201200041

Barnes, J., Zheng, Y., and Lyons, T. (2002). Air Pollution and Plant Biotechnology, Springer.

Gomez, 2004, Ascorbic acid biosynthesis: A precursor study on plants, Braz. J. Plant Physiol., 16, 147, 10.1590/S1677-04202004000300004

Horemans, 2000, Transport and action of ascorbate at the plant plasma membrane, Trends Plant Sci., 5, 263, 10.1016/S1360-1385(00)01649-6

Gallie, 2013, L-ascorbic acid: A multifunctional molecule supporting plant growth and development, Scientifica, 2013, 795964, 10.1155/2013/795964

HongBo, 2005, Investigation on dynamic changes of photosynthetic characteristics of 10 wheat (Triticum aestivum L.) genotypes during two vegetative-growth stages at water deficits, Colloids Surf. B, 43, 221, 10.1016/j.colsurfb.2005.05.005

Noctor, 1998, Ascorbate and glutathione: Keeping active oxygen under control, Annu. Rev. Plant Biol., 49, 249, 10.1146/annurev.arplant.49.1.249

Smirnof, 1993, Therole ofactive oxygenin theresponse of plants to water deficit and desiccation, New Phytol., 125, 27, 10.1111/j.1469-8137.1993.tb03863.x

Asada, 2006, Production and scavenging of reactive oxygen species in chloroplasts and their functions, Plant Physiol., 141, 391, 10.1104/pp.106.082040

Agarwal, 2007, Increased antioxidant activity in Cassia seedlings under UV-B radiation, Biol. Plant., 51, 157, 10.1007/s10535-007-0030-z

Bartoli, 2013, Glutathione and ascorbic acid protect Arabidopsis plants against detrimental effects of iron deficiency, J. Exp. Bot., 64, 3169, 10.1093/jxb/ert153

Szarka, 2012, The ascorbate-glutathione-α-tocopherol triad in abiotic stress response, Int. J. Mol. Sci., 13, 4458, 10.3390/ijms13044458

Lu, 2013, Glutathione synthesis, Biochim. Biophys. Acta Gen. Subj., 1830, 3143, 10.1016/j.bbagen.2012.09.008

Larson, 1988, The antioxidants of higher plants, Phytochemistry, 27, 969, 10.1016/0031-9422(88)80254-1

Mullineaux, 2005, Glutathione, photosynthesis and the redox regulation of stress-responsive gene expression, Photosynth. Res., 86, 459, 10.1007/s11120-005-8811-8

Ahmad, 2010, Roles of enzymatic and nonenzymatic antioxidants in plants during abiotic stress, Crit. Rev. Biotechnol., 30, 161, 10.3109/07388550903524243

Millar, 2003, Control of ascorbate synthesis by respiration and its implications for stress responses, Plant Physiol., 133, 443, 10.1104/pp.103.028399

May, 1998, Glutathione homeostasis in plants: Implications for environmental sensing and plant development, J. Exp. Bot., 49, 649

Roychoudhury, A., and Basu, S. (2012). Oxidative Stress in Plants: Causes, Consequences and Tolerance, IK International Publishers.

Appelqvist, 1996, The chemistry and antioxidant properties of tocopherols and tocotrienols, Lipids, 31, 671, 10.1007/BF02522884

Grabowski, 2005, Tocopherol content and activities of tyrosine aminotransferase and cystine lyase in Arabidopsis under stress conditions, J. Plant Physiol., 162, 767, 10.1016/j.jplph.2005.04.019

Kiffin, 2006, Oxidative stress and autophagy, Antioxid. Redox Signal., 8, 152, 10.1089/ars.2006.8.152

Kruk, 2005, Tocopherol as singlet oxygen scavenger in photosystem II, J. Plant Physiol., 162, 749, 10.1016/j.jplph.2005.04.020

Noctor, 2006, Metabolic signalling in defence and stress: The central roles of soluble redox couples, Plant Cell Environ., 29, 409, 10.1111/j.1365-3040.2005.01476.x

Igamberdiev, 2004, Nitrate, NO and haemoglobin in plant adaptation to hypoxia: An alternative to classic fermentation pathways, J. Exp. Bot., 55, 2473, 10.1093/jxb/erh272

Yu, 2004, MAP kinase cascades responding to environmental stress in plants, Acta Bot. Sin., 46, 127

Hare, 1998, Dissecting the roles of osmolyte accumulation during stress, Plant Cell Environ., 21, 535, 10.1046/j.1365-3040.1998.00309.x

Stahl, 2003, Antioxidant activity of carotenoids, Mol. Asp. Med., 24, 345, 10.1016/S0098-2997(03)00030-X

Muller, 2011, Antioxidant activity of β-carotene compounds in different in vitro assays, Molecules, 16, 1055, 10.3390/molecules16021055

Mortensen, 2001, The interaction of dietary carotenoids with radical species, Arch. Biochem. Biophys., 385, 13, 10.1006/abbi.2000.2172

Dewick, P.M. (2002). Medicinal Natural Products: A Biosynthetic Approach, John Wiley & Sons.

Gruszecki, 2005, Carotenoids as modulators of lipid membrane physical properties, Biochim. Biophys. Acta Mol. Basis Dis., 1740, 108, 10.1016/j.bbadis.2004.11.015

Heber, 2002, Overview of mechanisms of action of lycopene, Exp. Biol. Med., 227, 920, 10.1177/153537020222701013

Woodall, 1997, Oxidation of carotenoids by free radicals: Relationship between structure and reactivity, Biochim. Biophys. Acta Gen. Subj., 1336, 33, 10.1016/S0304-4165(97)00006-8

Ramel, 2012, Chemical quenching of singlet oxygen by carotenoids in plants, Plant Physiol., 158, 1267, 10.1104/pp.111.182394

Ozhogina, 1995, β-Carotene as an interceptor of free radicals, Free Radic. Biol. Med., 19, 575, 10.1016/0891-5849(95)00064-5

Das, 2014, Reactive oxygen species (ROS) and response of antioxidants as ROS-scavengers during environmental stress in plants, Front. Environ. Sci., 2, 53, 10.3389/fenvs.2014.00053

Polovka, 2003, Antioxidant properties of tea investigated by EPR spectroscopy, Biophys. Chem., 106, 39, 10.1016/S0301-4622(03)00159-5

Bravo, 1998, Polyphenols: Chemistry, dietary sources, metabolism, and nutritional significance, Nutr. Rev., 56, 317, 10.1111/j.1753-4887.1998.tb01670.x

Andersen, O.M., and Markham, K.R. (2005). Flavonoids: Chemistry, Biochemistry and Applications, CRC Press.

Chen, 1996, Antioxidant activity of natural flavonoids is governed by number and location of their aromatic hydroxyl groups, Chem. Phys. Lipids, 79, 157, 10.1016/0009-3084(96)02523-6

Morel, 1993, Antioxidant and iron-chelating activities of the flavonoids catechin, quercetin and diosmetin on iron-loaded rat hepatocyte cultures, Biochem. Pharmacol., 45, 13, 10.1016/0006-2952(93)90371-3

Yordi, 2012, Antioxidant and pro-oxidant effects of polyphenolic compounds and structure-activity relationship evidence, Nutr. Well-Being Health, 2, 23

Nijveldt, 2001, Flavonoids: A review of probable mechanisms of action and potential applications, Am. J. Clin. Nutr., 74, 418, 10.1093/ajcn/74.4.418

Foyer, 2005, Redox homeostasis and antioxidant signaling: A metabolic interface between stress perception and physiological responses, Plant Cell, 17, 1866, 10.1105/tpc.105.033589

Qamar, 2015, Role of proline and pyrroline-5-carboxylate metabolism in plant defense against invading pathogens, Front. Plant Sci., 6, 503, 10.3389/fpls.2015.00503

Verbruggen, 2008, Proline accumulation in plants: A review, Amino Acids, 35, 753, 10.1007/s00726-008-0061-6

Szabados, 2010, Proline: A multifunctional amino acid, Trends Plant Sci., 15, 89, 10.1016/j.tplants.2009.11.009

Decros, 2019, Get the balance right: ROS homeostasis and redox signalling in fruit, Front. Plant Sci., 10, 1091, 10.3389/fpls.2019.01091

Paciolla, C., Paradiso, A., and De Pinto, M. (2016). Redox State as a Central Regulator of Plant-Cell Stress Responses, Springer.

Finkel, 2011, Signal transduction by reactive oxygen species, J. Cell Biol., 194, 7, 10.1083/jcb.201102095

Rehman, 2019, Effect of salinity on cadmium tolerance, ionic homeostasis and oxidative stress responses in conocarpus exposed to cadmium stress: Implications for phytoremediation, Ecotoxicol. Environ. Saf., 171, 146, 10.1016/j.ecoenv.2018.12.077

Cheng, 2020, Thymol confers tolerance to salt stress by activating anti-oxidative defense and modulating Na+ homeostasis in rice root, Ecotoxicol. Environ. Saf., 188, 109894, 10.1016/j.ecoenv.2019.109894

Ahanger, 2020, Combined effects of brassinosteroid and kinetin mitigates salinity stress in tomato through the modulation of antioxidant and osmolyte metabolism, Plant Physiol. Biochem., 147, 31, 10.1016/j.plaphy.2019.12.007

Ahmad, 2019, Silicon (Si) supplementation alleviates NaCl toxicity in mung bean [Vigna radiata (L.) Wilczek] through the modifications of physio-biochemical attributes and key antioxidant enzymes, J. Plant Growth Regul., 38, 70, 10.1007/s00344-018-9810-2

Arora, 2020, Interaction between Piriformospora indica and Azotobacter chroococcum diminish the effect of salt stress in Artemisia annua L. by enhancing enzymatic and non-enzymatic antioxidants, Symbiosis, 80, 61, 10.1007/s13199-019-00656-w

Lalarukh, 2020, Response of antioxidants and lipid peroxidation to exogenous application of alpha-tocopherol in sunflower (Helianthus annuus L.) under salt stress, Pak. J. Bot., 52, 75, 10.30848/PJB2020-1(41)

Tariq, 2020, Glycinebetaine induced modulation in oxidative defense system and mineral nutrients sesame (Sesamum indicum L.) under saline regimes, Pak. J. Bot., 52, 775, 10.30848/PJB2020-3(34)

Mhadhbi, 2013, Alternative oxidase 1 (Aox1) gene expression in roots of Medicago truncatula is a genotype-specific component of salt stress tolerance, J. Plant Physiol., 170, 111, 10.1016/j.jplph.2012.08.017

Filippou, 2014, Proline and reactive oxygen/nitrogen species metabolism is involved in the tolerant response of the invasive plant species Ailanthus altissima to drought and salinity, Environ. Exp. Bot., 97, 1, 10.1016/j.envexpbot.2013.09.010

Cunha, 2016, Salinity and osmotic stress trigger different antioxidant responses related to cytosolic ascorbate peroxidase knockdown in rice roots, Environ. Exp. Bot., 131, 58, 10.1016/j.envexpbot.2016.07.002

Vighi, 2017, Functional characterization of the antioxidant enzymes in rice plants exposed to salinity stress, Biol. Plant., 61, 540, 10.1007/s10535-017-0727-6

Zeeshan, M., Lu, M., Sehar, S., Holford, P., and Wu, F. (2020). Comparison of biochemical, anatomical, morphological, and physiological responses to salinity stress in wheat and barley genotypes deferring in salinity tolerance. Agronomy, 10.

Alzahrani, 2019, Physiological, biochemical, and antioxidant properties of two genotypes of Vicia faba grown under salinity stress, Pak. J. Bot., 51, 786, 10.30848/PJB2019-3(3)

Alsahli, 2019, Salicylic acid alleviates salinity stress through the modulation of biochemical attributes and some key antioxidants in wheat seedlings, Pak. J. Bot., 51, 1551, 10.30848/PJB2019-5(12)

Ali, 2020, Exogenous jasmonic acid and humic acid increased salinity tolerance of sorghum, Agronomy, 112, 871, 10.1002/agj2.20072

Tanou, 2014, Polyamines reprogram oxidative and nitrosative status and the proteome of citrus plants exposed to salinity stress, Plant Cell Environ., 37, 864, 10.1111/pce.12204

Ahanger, 2019, Nitrogen availability prevents oxidative effects of salinity on wheat growth and photosynthesis by up-regulating the antioxidants and osmolytes metabolism, and secondary metabolite accumulation, BMC Plant Biol., 19, 1, 10.1186/s12870-019-2085-3

Chung, 2020, Silicon confers soybean resistance to salinity stress through regulation of reactive oxygen and reactive nitrogen species, Front. Plant Sci., 10, 1725, 10.3389/fpls.2019.01725

Christou, 2013, Hydrogen sulfide induces systemic tolerance to salinity and non-ionic osmotic stress in strawberry plants through modification of reactive species biosynthesis and transcriptional regulation of multiple defence pathways, J. Exp. Bot., 64, 1953, 10.1093/jxb/ert055

Santander, 2020, Efficiency of two arbuscular mycorrhizal fungal inocula to improve saline stress tolerance in lettuce plants by changes of antioxidant defense mechanisms, J. Sci. Food Agric., 100, 1577, 10.1002/jsfa.10166

Rady, 2019, Pretreatment with proline or an organic bio-stimulant induces salt tolerance in wheat plants by improving antioxidant redox state and enzymatic activities and reducing the oxidative stress, J. Plant Growth Regul., 38, 449, 10.1007/s00344-018-9860-5

Rady, 2019, Interplaying roles of silicon and proline effectively improve salt and cadmium stress tolerance in Phaseolus vulgaris plant, Plant Physiol. Biochem., 139, 558, 10.1016/j.plaphy.2019.04.025

Heydari, 2019, Role of Penconazole in salt stress amelioration in Sesamum indicum L., Soil Sci. Plant Nutr., 65, 243, 10.1080/00380768.2019.1595722

Aubert, 1999, Carbon metabolism in the subantarctic Kerguelen cabbage Pringlea antiscorbutica R. Br.: Environmental controls over carbohydrates and proline contents and relation to phenology, Plant Cell Environ., 22, 243, 10.1046/j.1365-3040.1999.00417.x

Yan, 2000, Effects of exogenous proline on the physiology of soyabean plantlets regenerated from embryos in vitro and on the ultrastructure of their mitochondria under NaCl stress, Soybean Sci., 19, 314

Djilianov, 2005, Improved abiotic stress tolerance in plants by accumulation of osmoprotectants—Gene transfer approach, Biotechnol. Biotechnol. Equip., 19, 63, 10.1080/13102818.2005.10817287

Jaarsma, R., de Vries, R.S., and de Boer, A.H. (2013). Effect of salt stress on growth, Na+ accumulation and proline metabolism in potato (Solanum tuberosum) cultivars. PLoS ONE, 8.

Rashwan, 2020, Mitigating salt stress effects by exogenous application of proline and yeast extract on morpho-physiological, biochemical and anatomical characters of calendula plants, Sci. J. Flowers Ornam. Plants, 7, 461, 10.21608/sjfop.2020.135166

Abdelaal, 2021, Salt tolerance activation in faba bean plants using proline and salicylic acid associated with physio-biochemical and yield characters improvement, Fresenius Environ. Bull., 30, 3175

Blasco, 2018, Influence of the proline metabolism and glycine betaine on tolerance to salt stress in tomato (Solanum lycopersicum L.) commercial genotypes, J. Plant Physiol., 231, 329, 10.1016/j.jplph.2018.10.013

Hannachi, 2018, Salt stress affects germination, seedling growth and physiological responses differentially in eggplant cultivars (Solanum melongena L.), Sci. Hortic., 228, 56, 10.1016/j.scienta.2017.10.002

Abdelaal, K., Attia, K.A., Niedbała, G., Wojciechowski, T., Hafez, Y., Alamery, S., Alateeq, T.K., and Arafa, S.A. (2021). Mitigation of Drought Damages by Exogenous Chitosan and Yeast Extract with Modulating the Photosynthetic Pigments, Antioxidant Defense System and Improving the Productivity of Garlic Plants. Horticulturae, 7.

Hare, 1997, Metabolic implications of stress-induced proline accumulation in plants, Plant Growth Regul., 21, 79, 10.1023/A:1005703923347

Zsigmond, 2008, Duplicated P5CS genes of Arabidopsis play distinct roles in stress regulation and developmental control of proline biosynthesis, Plant J., 53, 11, 10.1111/j.1365-313X.2007.03318.x

Chen, 2008, Glycinebetaine: An effective protectant against abiotic stress in plants, Trends Plant Sci., 13, 499, 10.1016/j.tplants.2008.06.007

Hoque, 2007, Exogenous proline and glycinebetaine increase NaCl-induced ascorbate–glutathione cycle enzyme activities, and proline improves salt tolerance more than glycinebetaine in tobacco Bright Yellow-2 suspension-cultured cells, J. Plant Physiol., 164, 1457, 10.1016/j.jplph.2006.10.004

Banu, 2009, Proline and glycinebetaine induce antioxidant defense gene expression and suppress cell death in cultured tobacco cells under salt stress, J. Plant Physiol., 166, 146, 10.1016/j.jplph.2008.03.002

Fotopoulos, 2006, Effect of ascorbate oxidase over-expression on ascorbate recycling gene expression in response to agents imposing oxidative stress, J. Exp. Bot., 57, 3933, 10.1093/jxb/erl147

Neill, 2002, Hydrogen peroxide and nitric oxide as signalling molecules in plants, J. Exp. Bot., 53, 1237, 10.1093/jexbot/53.372.1237

Molassiotis, 2011, Oxidative and nitrosative signaling in plants: Two branches in the same tree?, Plant Signal. Behav., 6, 210, 10.4161/psb.6.2.14878

Zhao, 2007, Nitric oxide synthase-dependent nitric oxide production is associated with salt tolerance in Arabidopsis, Plant Physiol., 144, 206, 10.1104/pp.107.096842

Qiao, 2009, Expression of a rice gene OsNOA1 re-establishes nitric oxide synthesis and stress-related gene expression for salt tolerance in Arabidopsis nitric oxide-associated 1 mutant Atnoa1, Environ. Exp. Bot., 65, 90, 10.1016/j.envexpbot.2008.06.002

Wahid, 2007, Pretreatment of seed with H2O2 improves salt tolerance of wheat seedlings by alleviation of oxidative damage and expression of stress proteins, J. Plant Physiol., 164, 283, 10.1016/j.jplph.2006.01.005

Lu, 2009, Abscisic acid improves drought tolerance of triploid bermudagrass and involves H2O2-and NO-induced antioxidant enzyme activities, Plant Physiol. Biochem., 47, 132, 10.1016/j.plaphy.2008.10.006

Zhang, 2009, Increased abscisic acid levels in transgenic tobacco over-expressing 9 cis-epoxycarotenoid dioxygenase influence H2O2 and NO production and antioxidant defences, Plant Cell Environ., 32, 509, 10.1111/j.1365-3040.2009.01945.x

Xie, 2008, Carbon monoxide enhances salt tolerance by nitric oxide-mediated maintenance of ion homeostasis and up-regulation of antioxidant defence in wheat seedling roots, Plant Cell Environ., 31, 1864, 10.1111/j.1365-3040.2008.01888.x

Zhao, G., Zhao, Y., Yu, X., Kiprotich, F., Han, H., Guan, R., Wang, R., and Shen, W. (2018). Nitric oxide is required for melatonin-enhanced tolerance against salinity stress in rapeseed (Brassica napus L.) seedlings. Int. J. Mol. Sci., 19.

Tari, 2011, Ethylene-regulated reactive oxygen species and nitric oxide under salt stress in tomato cell suspension culture, Acta Biol. Szeged., 55, 143

Molassiotis, 2010, NO says more than ‘YES’ to salt tolerance: Salt priming and systemic nitric oxide signaling in plants, Plant Signal. Behav., 5, 209, 10.4161/psb.5.3.10738

Xu, 2011, Involvement of auxin and nitric oxide in plant Cd-stress responses, Plant Soil, 346, 107, 10.1007/s11104-011-0800-4

Kong, 2016, Exogenous nitric oxide delays salt-induced leaf senescence in cotton (Gossypium hirsutum L.), Acta Physiol. Plant, 38, 1, 10.1007/s11738-016-2079-9

Wang, 2009, Ethylene and nitric oxide are involved in maintaining ion homeostasis in Arabidopsis callus under salt stress, Planta, 230, 293, 10.1007/s00425-009-0946-y

Simaei, 2011, Interactive effects of salicylic acid and nitric oxide on soybean plants under NaCl salinity, Russian J. Plant Physiol., 58, 783, 10.1134/S1021443711050220

Dong, 2015, Effects of rhizopheric nitric oxide (NO) on N uptake in Fagus sylvatica seedlings depend on soil CO2 concentration, soil N availability and N source, Tree Physiol., 35, 910, 10.1093/treephys/tpv051

Kesawat, 2012, Genome-wide identification, evolutionary and expression analyses of putative Fe–S biogenesis genes in rice (Oryza sativa), Genome, 55, 571, 10.1139/g2012-044

Kesawat, M.S., Das, B.K., Kumar, M., and Bhaganagare, G.R. (2015). Biological Nitrogen Fixation, Wiley Online Library.

Fatma, 2016, Nitric oxide alleviates salt stress inhibited photosynthetic performance by interacting with sulfur assimilation in mustard, Front. Plant Sci., 7, 521, 10.3389/fpls.2016.00521

Fan, 2012, Effect of nitric oxide on proline metabolism in cucumber seedlings under salinity stress, J. Am. Soc. Hortic. Sci., 137, 127, 10.21273/JASHS.137.3.127

Khan, 2012, Interactive role of nitric oxide and calcium chloride in enhancing tolerance to salt stress, Nitric Oxide, 27, 210, 10.1016/j.niox.2012.07.005

Campos, 2019, Nitric oxide and phytohormone interactions in the response of Lactuca sativa to salinity stress, Planta, 250, 1475, 10.1007/s00425-019-03236-w

Babaei, 2021, Comparative effects of nitric oxide and salicylic acid on salinity tolerance in saffron (Crocus sativus), Plant Biosyst., 155, 73, 10.1080/11263504.2020.1727975

Jiang, 2007, Conservation of the salt overly sensitive pathway in rice, Plant Physiol., 143, 1001, 10.1104/pp.106.092635

Ji, 2013, The salt overly sensitive (SOS) pathway: Established and emerging roles, Mol. Plant, 6, 275, 10.1093/mp/sst017

Kurusu, 2015, Plant signaling networks involving Ca2+ and Rboh/Nox-mediated ROS production under salinity stress, Front. Plant Sci., 6, 427, 10.3389/fpls.2015.00427

Coll, 2011, Programmed cell death in the plant immune system, Cell Death Differ., 18, 1247, 10.1038/cdd.2011.37

Kwak, 2003, NADPH oxidase AtrbohD and AtrbohF genes function in ROS-dependent ABA signaling in Arabidopsis, EMBO J., 22, 2623, 10.1093/emboj/cdg277

Szepesi, 2009, Salicylic acid improves acclimation to salt stress by stimulating abscisic aldehyde oxidase activity and abscisic acid accumulation, and increases Na+ content in leaves without toxicity symptoms in Solanum lycopersicum L., J. Plant Physiol., 166, 914, 10.1016/j.jplph.2008.11.012

Takeda, 2008, Local positive feedback regulation determines cell shape in root hair cells, Science, 319, 1241, 10.1126/science.1152505

Pottosin, 2014, Cross-talk between reactive oxygen species and polyamines in regulation of ion transport across the plasma membrane: Implications for plant adaptive responses, J. Exp. Bot., 65, 1271, 10.1093/jxb/ert423

Xu, 2013, De novo transcriptome sequencing and comparative analysis of differentially expressed genes in Gossypium aridum under salt stress, Gene, 525, 26, 10.1016/j.gene.2013.04.066

Choi, 2014, Salt stress-induced Ca2+ waves are associated with rapid, long-distance root-to-shoot signaling in plants, Proc. Natl. Acad. Sci. USA, 111, 6497, 10.1073/pnas.1319955111

Mostofa, 2015, Hydrogen sulfide modulates cadmium-induced physiological and biochemical responses to alleviate cadmium toxicity in rice, Sci. Rep., 5, 1, 10.1038/srep14078

Kodela, 2012, NOSH-Aspirin: A novel nitric oxide–hydrogen sulfide-releasing hybrid: A new class of anti-inflammatory pharmaceuticals, ACS Med. Chem. Lett., 3, 257, 10.1021/ml300002m

Zhang, 2015, Hydrogen sulfide alleviates cadmium-induced cell death through restraining ROS accumulation in roots of Brassica rapa L. ssp. pekinensis, Oxid. Med. Cell. Longev., 2015, 804603, 10.1155/2015/804603

Hancock, 2016, Hydrogen sulfide signaling: Interactions with nitric oxide and reactive oxygen species, Ann. N. Y. Acad. Sci., 1365, 5, 10.1111/nyas.12733

Paranhos, 2014, Interplay of calcium, cAMP and PKA in flavonoid accumulation by cell cultures of Hypericum androsaemum L., Planta Med., 80, P2O63, 10.1055/s-0034-1395053

Vafadar, 2020, Crosstalk between melatonin and Ca2+/CaM evokes systemic salt tolerance in Dracocephalum kotschyi, J. Plant Physiol., 252, 153237, 10.1016/j.jplph.2020.153237

Hajihashemi, 2020, Cross-talk between nitric oxide, hydrogen peroxide and calcium in salt-stressed Chenopodium quinoa Willd. At seed germination stage, Plant Physiol. Biochem., 154, 657, 10.1016/j.plaphy.2020.07.022

Souissi, 2017, Salicylic acid and hydrogen peroxide pretreatments alleviate salt stress in faba bean (Vicia faba) seeds during germination, Seed Sci. Technol., 45, 675

Pathak, 2014, Polyamines in response to abiotic stress tolerance through transgenic approaches, GM Crops Food, 5, 87, 10.4161/gmcr.28774

Shi, 2014, Improvement of plant abiotic stress tolerance through modulation of the polyamine pathway, J. Integr. Plant Biol., 56, 114, 10.1111/jipb.12128

Do, 2014, Changes in free polyamine levels, expression of polyamine biosynthesis genes, and performance of rice cultivars under salt stress: A comparison with responses to drought, Front. Plant Sci., 5, 182, 10.3389/fpls.2014.00182

Zheng, Q., Su, S., Wang, Z., Wang, Y., and Xu, X. (2021). Comprehensive Genome-Wide Identification and Transcript Profiling of GABA Pathway Gene Family in Apple (Malus domestica). Genes, 12.

Shabala, 2012, Salt-sensitive and salt-tolerant barley varieties differ in the extent of potentiation of the ROS-induced K+ efflux by polyamines, Plant Physiol. Biochem., 61, 18, 10.1016/j.plaphy.2012.09.002

Santolini, 2017, Nitric oxide synthase in plants: Where do we stand?, Nitric Oxide, 63, 30, 10.1016/j.niox.2016.09.005

Gupta, 2017, 24-Epibrassinolide and sodium nitroprusside alleviate the salinity stress in Brassica juncea L. cv. Varuna through cross talk among proline, nitrogen metabolism and abscisic acid, Plant Soil, 411, 483, 10.1007/s11104-016-3043-6

Siddiqui, 2012, Cumulative effect of nitrogen and sulphur on Brassica juncea L. genotypes under NaCl stress, Protoplasma, 249, 139, 10.1007/s00709-011-0273-6

Prakash, 2019, Crosstalk between nitric oxide (NO) and abscisic acid (ABA) signalling molecules in higher plants, Environ. Exp. Bot., 161, 41, 10.1016/j.envexpbot.2018.10.033

Kumar, M., Kherawat, B.S., Dey, P., Saha, D., Singh, A., Bhatia, S.K., Ghodake, G.S., Kadam, A.A., Kim, H.-U., and Chung, S.-M. (2021). Genome-wide identification and characterization of PIN-FORMED (PIN) gene family reveals role in developmental and various stress conditions in Triticum aestivum L.. Int. J. Mol. Sci., 22.

Iqbal, N., Masood, A., and Khan, N.A. (2012). Phytohormones and Abiotic Stress Tolerance in Plants, Springer.

Bialecka, 2009, Effect of ethephon and gibberellin A3 on Amaranthus caudatus seed germination and alpha-and beta-amylase activity under salinity stress, Acta Biol. Crac. Ser. Bot., 2, 119

Foo, 2006, A role for ethylene in the phytochrome-mediated control of vegetative development, Plant J., 46, 911, 10.1111/j.1365-313X.2006.02754.x

Lin, 2013, Ethylene promotes germination of Arabidopsis seed under salinity by decreasing reactive oxygen species: Evidence for the involvement of nitric oxide simulated by sodium nitroprusside, Plant Physiol. Biochem., 73, 211, 10.1016/j.plaphy.2013.10.003

Yalpani, 1994, Ultraviolet light and ozone stimulate accumulation of salicylic acid, pathogenesis-related proteins and virus resistance in tobacco, Planta, 193, 372, 10.1007/BF00201815

Durner, 1996, Salicylic acid is a modulator of tobacco and mammalian catalases, J. Biol. Chem., 271, 28492, 10.1074/jbc.271.45.28492

Szalai, 2000, Effects of cold acclimation and salicylic acid on changes in ACC and MACC contents in maize during chilling, Biol. Plant., 43, 637, 10.1023/A:1002824721597

Klessig, 1994, The salicylic acid signal in plants, Plant Mol. Biol., 26, 1439, 10.1007/BF00016484

Strnad, 2009, Salicylic acid-induced changes to growth and phenolic metabolism in Matricaria chamomilla plants, Plant Cell Rep., 28, 135, 10.1007/s00299-008-0627-5

Li, T., Hu, Y., Du, X., Tang, H., Shen, C., and Wu, J. (2014). Salicylic acid alleviates the adverse effects of salt stress in Torreya grandis cv. Merrillii seedlings by activating photosynthesis and enhancing antioxidant systems. PLoS ONE, 9.

Arfan, 2007, Does exogenous application of salicylic acid through the rooting medium modulate growth and photosynthetic capacity in two differently adapted spring wheat cultivars under salt stress?, J. Plant Physiol., 164, 685, 10.1016/j.jplph.2006.05.010

Cao, 2009, Lack of salicylic acid in Arabidopsis protects plants against moderate salt stress, Z. Naturforsch. C J. Biosci., 64, 231, 10.1515/znc-2009-3-414

Friml, 2003, Auxin transport—Shaping the plant, Curr. Opin. Plant Biol., 6, 7, 10.1016/S1369526602000031

Ganguly, 2014, Functional analysis of the hydrophilic loop in intracellular trafficking of Arabidopsis PIN-FORMED proteins, Plant Cell, 26, 1570, 10.1105/tpc.113.118422

Wang, 2021, OsIAA18, an aux/IAA transcription factor gene, is involved in salt and drought tolerance in rice, Front. Plant Sci., 12, 2571, 10.3389/fpls.2021.738660

Sun, 2008, Salt modulates gravity signaling pathway to regulate growth direction of primary roots in Arabidopsis, Plant Physiol., 146, 178, 10.1104/pp.107.109413

Shen, 2010, Expression profile of PIN, AUX/LAX and PGP auxin transporter gene families in Sorghum bicolor under phytohormone and abiotic stress, FEBS J., 277, 2954, 10.1111/j.1742-4658.2010.07706.x

Julkowska, 2015, Tuning plant signaling and growth to survive salt, Trends Plant Sci., 20, 586, 10.1016/j.tplants.2015.06.008

Song, 2009, Comprehensive expression profiling analysis of OsIAA gene family in developmental processes and in response to phytohormone and stress treatments, Planta, 229, 577, 10.1007/s00425-008-0853-7

Li, 2020, The rice Aux/IAA transcription factor gene OsIAA18 enhances salt and osmotic tolerance in Arabidopsis, Biol. Plant, 64, 454, 10.32615/bp.2019.069

Jung, 2015, OsIAA6, a member of the rice Aux/IAA gene family, is involved in drought tolerance and tiller outgrowth, Plant Sci., 236, 304, 10.1016/j.plantsci.2015.04.018

Zhang, 2021, OsIAA20, an Aux/IAA protein, mediates abiotic stress tolerance in rice through an ABA pathway, Plant Sci., 308, 110903, 10.1016/j.plantsci.2021.110903

Li, W., Dang, C., Ye, Y., Wang, Z., Hu, L., Zhang, F., Zhang, Y., Qian, X., Shi, J., and Guo, Y. (2020). Overexpression of grapevine VvIAA18 gene enhanced salt tolerance in tobacco. Int. J. Mol. Sci., 21.

Wang, 2017, Overexpression of a chrysanthemum transcription factor gene DgNAC1 improves the salinity tolerance in chrysanthemum, Plant Cell Rep., 36, 571, 10.1007/s00299-017-2103-6

Shafi, 2017, Transgenic potato plants overexpressing SOD and APX exhibit enhanced lignification and starch biosynthesis with improved salt stress tolerance, Plant Mol. Biol. Rep., 35, 504, 10.1007/s11105-017-1041-3

Tang, 2019, Overexpression of a MYB family gene, OsMYB6, increases drought and salinity stress tolerance in transgenic rice, Front. Plant Sci., 10, 168, 10.3389/fpls.2019.00168

Liu, 2020, Overexpression of StCYS1 gene enhances tolerance to salt stress in the transgenic potato (Solanum tuberosum L.) plant, J. Integr. Agric., 19, 2239, 10.1016/S2095-3119(20)63262-2

Wang, 2020, Overexpression of the transcription factor gene OsSTAP1 increases salt tolerance in rice, Rice, 13, 1, 10.1186/s12284-020-00405-4

Xu, 2014, Overexpression of DnWRKY11 enhanced salt and drought stress tolerance of transgenic tobacco, Biologia, 69, 994, 10.2478/s11756-014-0398-0

Zhang, 2020, The salt-induced transcription factor GmMYB84 confers salinity tolerance in soybean, Plant Sci., 291, 110326, 10.1016/j.plantsci.2019.110326

Jadamba, C., Kang, K., Paek, N.-C., Lee, S.I., and Yoo, S.-C. (2020). Overexpression of rice expansin7 (Osexpa7) confers enhanced tolerance to salt stress in rice. Int. J. Mol. Sci., 21.

Wen, 2021, MsWRKY11, activated by MsWRKY22, functions in drought tolerance and modulates lignin biosynthesis in alfalfa (Medicago sativa L.), Environ. Exp. Bot., 184, 104373, 10.1016/j.envexpbot.2021.104373

Trinh, 2019, Overexpression of the dominant negative nbexo70d1 mutantion confers tolerance to salt stress in transgenic tobacco, Biol. Plant., 63, 484

Zhu, 2019, VvWRKY30, a grape WRKY transcription factor, plays a positive regulatory role under salinity stress, Plant Sci., 280, 132, 10.1016/j.plantsci.2018.03.018

Wang, Q., Ni, J., Shah, F., Liu, W., Wang, D., Yao, Y., Hu, H., Huang, S., Hou, J., and Fu, S. (2019). Overexpression of the stress-inducible SsMAX2 promotes drought and salt resistance via the regulation of redox homeostasis in Arabidopsis. Int. J. Mol. Sci., 20.

Xiang, X.-Y., Chen, J., Xu, W.-X., Qiu, J.-R., Song, L., Wang, J.-T., Tang, R., Chen, D., Jiang, C.-Z., and Huang, Z. (2021). Dehydration-induced WRKY transcriptional factor MfWRKY70 of Myrothamnus flabellifolia enhanced drought and salinity tolerance in Arabidopsis. Biomolecules, 11.

Shen, 2013, Overexpression of PeHSF mediates leaf ROS homeostasis in transgenic tobacco lines grown under salt stress conditions, Plant Cell Tissue Organ Cult., 115, 299, 10.1007/s11240-013-0362-7

Sun, T.-T., Wang, C., Liu, R., Zhang, Y., Wang, Y.-C., and Wang, L.-Q. (2021). ThHSFA1 confers salt stress tolerance through modulation of reactive oxygen species scavenging by directly regulating ThWRKY4. Int. J. Mol. Sci., 22.

Yang, D.-Y., Zhuang, K.-Y., and Ma, N.-N. (2022). Protoplasma, Springer.