{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T18:17:27Z","timestamp":1791569847183,"version":"4.3.4"},"reference-count":125,"publisher":"MDPI AG","issue":"7","license":[{"start":{"date-parts":[[2020,7,10]],"date-time":"2020-07-10T00:00:00Z","timestamp":1594339200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"CONACYT-M\u00e9xico","award":["PN-4631 a project supported by FOINS"],"award-info":[{"award-number":["PN-4631 a project supported by FOINS"]}]},{"DOI":"10.13039\/501100007071","name":"Harran \u00dcniversitesi","doi-asserted-by":"publisher","award":["HUBAP-17225"],"award-info":[{"award-number":["HUBAP-17225"]}],"id":[{"id":"10.13039\/501100007071","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Antioxidants"],"abstract":"<jats:p>The aim of this study is to assess the role of l-cysteine desulfhydrase (l-DES) and endogenous hydrogen sulfide (H2S) in glutathione (GSH)-induced tolerance to salinity stress (SS) in sweet pepper (Capsicum annuum L.). Two weeks after germination, before initiating SS, half of the pepper seedlings were retained for 12 h in a liquid solution containing H2S scavenger, hypotaurine (HT), or the l-DES inhibitor dl-propargylglycine (PAG). The seedlings were then exposed for three weeks to control or SS (100 mmol L\u22121 NaCl) and supplemented with or without GSH or GSH+NaHS (sodium hydrosulfide, H2S donor). Salinity suppressed dry biomass, leaf water potential, chlorophyll contents, maximum quantum efficiency, ascorbate, and the activities of dehydroascorbate reductase, monodehydroascorbate reductase, and glyoxalase II in plants. Contrarily, it enhanced the accumulation of hydrogen peroxide, malondialdehyde, methylglyoxal, electrolyte leakage, proline, GSH, the activities of glutathione reductase, peroxidase, catalase, superoxide dismutase, ascorbate peroxidase, glyoxalase I, and l-DES, as well as endogenous H2S content. Salinity enhanced leaf Na+ but reduced K+; however, the reverse was true with GSH application. Overall, the treatments, GSH and GSH+NaHS, effectively reversed the oxidative stress and upregulated salt tolerance in pepper plants by controlling the activities of the AsA-GSH and glyoxalase-system-related enzymes as well as the levels of osmolytes.<\/jats:p>","DOI":"10.3390\/antiox9070603","type":"journal-article","created":{"date-parts":[[2020,7,10]],"date-time":"2020-07-10T09:25:28Z","timestamp":1594373128000},"page":"603","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":34,"title":["Involvement of l-Cysteine Desulfhydrase and Hydrogen Sulfide in Glutathione-Induced Tolerance to Salinity by Accelerating Ascorbate-Glutathione Cycle and Glyoxalase System in Capsicum"],"prefix":"10.3390","volume":"9","author":[{"given":"Cengiz","family":"Kaya","sequence":"first","affiliation":[{"name":"Soil Science and Plant Nutrition Department, Agriculture Faculty, Harran University, Sanliurfa 6300, Turkey"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9489-4054","authenticated-orcid":false,"given":"Bernardo","family":"Murillo-Amador","sequence":"additional","affiliation":[{"name":"Centro de Investigaciones Biol\u00f3gicas del Noroeste, S.C. Avenida Instituto Polit\u00e9cnico Nacional No. 195, Colonia Playa Palo de Santa Rita Sur, La Paz 23096, Baja California Sur, Mexico"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Muhammad","family":"Ashraf","sequence":"additional","affiliation":[{"name":"Department of Botany, University of Agriculture, Faisalabad 38040, Pakistan"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2020,7,10]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"2180","DOI":"10.3389\/fpls.2017.02180","article-title":"Interactive effects of CO2 concentration and water regime on stable isotope signatures, nitrogen assimilation and growth in sweet pepper","volume":"8","author":"Serret","year":"2018","journal-title":"Front. Plant Sci."},{"key":"ref_2","unstructured":"Kijne, W. (2020, April 20). Working Towards Unlocking the Water Potential of Agriculture. Available online: http:\/\/agris.fao.org\/agris-search\/search.do?recordID=SO2005100464."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"227","DOI":"10.17221\/3433-PSE","article-title":"Hot pepper response to interactive effects of salinity and boron","volume":"52","author":"Lee","year":"2006","journal-title":"Plant Soil Environ."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Hussain, S., Shaukat, M., Ashraf, M., Zhu, C., Jin, Q., and Zhang, J. (2019). Salinity Stress in Arid and Semi-Arid Climates: Effects and Management in Field Crops, Intech Open.","DOI":"10.5772\/intechopen.87982"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"36","DOI":"10.3906\/bot-1907-27","article-title":"The endogenous l-cysteine desulfhydrase and hydrogen sulfide participate in supplemented phosphorus-induced tolerance to salinity stress in maize (Zea mays) plants","volume":"44","author":"Kaya","year":"2020","journal-title":"Turk. J. Bot."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"381","DOI":"10.1080\/03650340.2014.936400","article-title":"Individual and additive effects of Na+ and Cl\u2212 ions on rice under salinity stress","volume":"61","author":"Kumar","year":"2015","journal-title":"Arch. Agron. Soil Sci."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"825","DOI":"10.1111\/plb.12999","article-title":"Potassium-induced decrease in cytosolic Na+ alleviates deleterious effects of salt stress on wheat (Triticum aestivum L.)","volume":"21","author":"Gul","year":"2019","journal-title":"Plant Biol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"796","DOI":"10.1111\/jipb.12689","article-title":"Unraveling salt stress signalling in plants","volume":"60","author":"Yang","year":"2018","journal-title":"J. Integr. Plant Biol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"521","DOI":"10.1016\/j.jenvman.2016.09.009","article-title":"Silicon occurrence, uptake, transport and mechanisms of heavy metals, minerals and salinity enhanced tolerance in plants with future prospects: A review","volume":"183","author":"Imtiaz","year":"2016","journal-title":"J. Environ. Manag."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Parvin, K., Hasanuzzaman, M., Bhuyan, M.H.M., Mohsin, S.M., and Fujita, M. (2019). Quercetin mediated salt tolerance in tomato through the enhancement of plant antioxidant defense and glyoxalase systems. Plants, 8.","DOI":"10.3390\/plants8080247"},{"key":"ref_11","unstructured":"Ahmad, P., Ahanger, M.A., and Alyemeni, M.N. (2018). Environmental Stress and secondary metabolites in plants: An overview. Plant Metabolites and Regulation Under Environmental Stress, Academic Press."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1016\/j.sajb.2016.11.003","article-title":"Drought and salinity stress alter ROS accumulation, water retention, and osmolyte content in sorghum plants","volume":"108","author":"Nxele","year":"2017","journal-title":"S. Afr. J. Bot."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Zargar, S., and Zargar, M. (2018). Reactive Oxygen Species (ROS): A Way to Stress Survival. Plants in Abiotic Stress-Mediated Sensing and Signaling in Plants: An Omics Perspective, Springer.","DOI":"10.1007\/978-981-10-7479-0"},{"key":"ref_14","unstructured":"J\u00fcrgens, G., and Pimpl, P. (2018). ROS-related redox regulation and signaling in plants. Seminars in Cell Developmental Biology, Academic Press."},{"key":"ref_15","unstructured":"Khan, M.I.R., Ferrante, A., Reddy, P.S., and Khan, N.A. (2019). Role and Regulation of Osmolytes and ABA Interaction in Salt and Drought Stress Tolerance. Plant Signaling Molecules, Woodhead Publishing."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Parvin, K., Hasanuzzaman, M., Bhuyan, M.H.M., Nahar, K., Mohsin, S.M., and Fujita, M. (2019). Comparative physiological and biochemical changes in tomato (Solanum lycopersicum L.) under salt stress and recovery: Role of Antioxidant Defense and Glyoxalase Systems. Antioxidants, 8.","DOI":"10.3390\/antiox8090350"},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Duhan, S., Kumari, A., and Lal, M. (2019). Oxidative Stress and Antioxidant Defense Under Combined Waterlogging and Salinity Stresses in Reactive Oxygen, Nitrogen and Sulfur Species. Plants: Production, Metabolism, Signaling and Defense Mechanisms, Wiley.","DOI":"10.1002\/9781119468677.ch5"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1016\/j.freeradbiomed.2018.03.009","article-title":"Methylglyoxal\u2014A signaling molecule in plant abiotic stress responses","volume":"122","author":"Mostofa","year":"2018","journal-title":"Free Radic. Biol. Med."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1186","DOI":"10.1111\/pce.12968","article-title":"Manipulation of glyoxalase pathway confers tolerance to multiple stresses in rice","volume":"41","author":"Gupta","year":"2018","journal-title":"Plant Cell Environ."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1007\/s10343-018-0430-3","article-title":"Exogenous Silicon Protects Brassica napus Plants from Salinity-Induced Oxidative Stress Through the Modulation of AsA-GSH Pathway, Thiol-Dependent Antioxidant Enzymes and Glyoxalase Systems","volume":"70","author":"Hasanuzzaman","year":"2018","journal-title":"Gesunde Pflanz."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Alam, P., Albalawi, T.H., Altalayan, F.H., Bakht, M.A., Ahanger, M.A., Raja, V., and Ahmad, P. (2019). 24-Epibrassinolide (EBR) confers tolerance against NaCl stress in soybean plants by up-regulating antioxidant system, ascorbate-glutathione cycle, and glyoxalase system. Biomolecules, 9.","DOI":"10.3390\/biom9110640"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"454","DOI":"10.1111\/j.1365-3040.2011.02400.x","article-title":"Glutathione in plants: An integrated overview","volume":"35","author":"Noctor","year":"2012","journal-title":"Plant Cell Environ."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1154","DOI":"10.1016\/j.freeradbiomed.2015.09.023","article-title":"Glutathione\u2013linking cell proliferation to oxidative stress","volume":"89","author":"Kiddle","year":"2015","journal-title":"Free Radic. Biol. Med."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"877","DOI":"10.1007\/s00344-017-9691-9","article-title":"Exogenous glutathione modulates salinity tolerance of soybean [Glycine max (L.) Merrill] at reproductive stage","volume":"36","author":"Akram","year":"2017","journal-title":"J. Plant Growth Regul."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1016\/j.scienta.2019.02.026","article-title":"Application of exogenous glutathione confers salinity stress tolerance in tomato seedlings by modulating ions homeostasis and polyamine metabolism","volume":"250","author":"Zhou","year":"2019","journal-title":"Sci. Hortic."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Ahanger, M.A., Alyemeni, M.N., Wijaya, L., Alamri, S.A., Alam, P., Ashraf, M., and Ahmad, P. (2018). Potential of exogenously sourced kinetin in protecting Solanum lycopersicum from NaCl-induced oxidative stress through up-regulation of the antioxidant system, ascorbate-glutathione cycle and glyoxalase system. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0202175"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1007\/s12298-017-0422-2","article-title":"Glutathione in plants: Biosynthesis and physiological role in environmental stress tolerance","volume":"23","author":"Hasanuzzaman","year":"2017","journal-title":"Physiol. Mol. Biol. Plants"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1016\/j.envexpbot.2014.12.001","article-title":"Exogenous glutathione confers high temperature stress tolerance in mung bean (Vigna radiata L.) by modulating antioxidant defense and methylglyoxal detoxification system","volume":"112","author":"Nahar","year":"2015","journal-title":"Environ. Exp. Bot."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1080\/17429145.2018.1458913","article-title":"Exogenous glutathione attenuates lead-induced oxidative stress in wheat by improving antioxidant defense and physiological mechanisms","volume":"13","author":"Hasanuzzaman","year":"2018","journal-title":"J. Plant Interact."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1007\/s10535-015-0542-x","article-title":"Roles of exogenous glutathione in antioxidant defense system and methylglyoxal detoxification during salt stress in mung bean","volume":"59","author":"Nahar","year":"2015","journal-title":"Biol. Plant."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1016\/j.scienta.2017.02.021","article-title":"Exogenous glutathione alleviates salt-induced oxidative stress in tomato seedlings by regulating glutathione metabolism, redox status, and the antioxidant system","volume":"220","author":"Zhou","year":"2017","journal-title":"Sci. Hortic."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"4621","DOI":"10.4161\/psb.24007","article-title":"l-Cysteine Desulfhydrase 1 modulates the generation of the signaling molecule sulfide in plant cytosol","volume":"8","author":"Romero","year":"2013","journal-title":"Plant Signal. Behav."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1777","DOI":"10.1111\/pce.12982","article-title":"l-cysteine desulfhydrase-related H2S production is involved in OsSE5-promoted ammonium tolerance in roots of Oryza sativa","volume":"40","author":"Guo","year":"2017","journal-title":"Plant Cell Environ."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1016\/bs.mie.2014.11.035","article-title":"Analysis of some enzymes activities of hydrogen sulfide metabolism in plants","volume":"Volume 555","author":"Cardenas","year":"2015","journal-title":"Methods in Enzymology"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1016\/j.plantsci.2014.06.006","article-title":"Endogenous hydrogen sulfide enhances salt tolerance by coupling the reestablishment of redox homeostasis and preventing salt-induced K+ loss in seedlings of Medicago sativa","volume":"225","author":"Lai","year":"2014","journal-title":"Plant Sci."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1016\/j.plantsci.2016.12.011","article-title":"Salinity-induced accumulation of endogenous H2S and NO is associated with modulation of the antioxidant and redox defense systems in Nicotiana tabacum L. cv. Havana","volume":"256","author":"Fontes","year":"2017","journal-title":"Plant Sci."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"983","DOI":"10.1016\/j.tplants.2019.08.003","article-title":"Hydrogen sulfide: A new warrior against abiotic stress","volume":"24","author":"Corpas","year":"2019","journal-title":"Trends Plant Sci."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Corpas, F.J., and Palma, J.M. (2020). H2S signaling in plants and applications in agriculture. J. Adv. Res.","DOI":"10.1016\/j.jare.2020.03.011"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1016\/j.plaphy.2016.10.006","article-title":"Hydrogen sulfide and proline cooperate to alleviate cadmium stress in foxtail millet seedlings","volume":"109","author":"Tian","year":"2016","journal-title":"Plant Physiol. Biochem."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1111\/ppl.13001","article-title":"Hydrogen sulfide alleviates chromium stress on cauliflower by restricting its uptake and enhancing antioxidative system","volume":"168","author":"Ahmad","year":"2019","journal-title":"Physiol. Plant."},{"key":"ref_41","first-page":"150","article-title":"Hydrogen sulfide: A new endogenous player in an old mechanism of plant tolerance to high salinity","volume":"32","author":"Modolo","year":"2018","journal-title":"Acta Bot. Bras."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"678","DOI":"10.3389\/fpls.2019.00678","article-title":"H2S alleviates salinity stress in cucumber by maintaining the Na+\/K+ balance and regulating H2S metabolism and oxidative stress response","volume":"10","author":"Jiang","year":"2019","journal-title":"Front. Plant Sci."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1016\/j.scienta.2015.01.009","article-title":"Exogenous application of nitric oxide promotes growth and oxidative defense system in highly boron stressed tomato plants bearing fruit","volume":"185","author":"Kaya","year":"2015","journal-title":"Sci. Hortic."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1016\/j.jplph.2014.12.018","article-title":"Hydrogen sulfide acts as a downstream signal molecule in salicylic acid-induced heat tolerance in maize (Zea mays L.) seedlings","volume":"177","author":"Li","year":"2015","journal-title":"J. Plant Physiol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"3199","DOI":"10.1007\/s11738-014-1686-6","article-title":"The alleviating effects of selenium and salicylic acid in salinity exposed soybean","volume":"36","author":"Ardebili","year":"2014","journal-title":"Acta Physiol. Plant."},{"key":"ref_46","unstructured":"Vernon, L.P., and Seely, G.R. (1966). Extraction separation estimation and isolation of the chlorophylls. The Chlorophylls, Academic Press."},{"key":"ref_47","first-page":"69","article-title":"Measurements of leaf relative water content in Araucaria angustifolia","volume":"11","author":"Yamasaki","year":"1999","journal-title":"Rev. Bras. Fisiol. Veg."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1016\/j.envexpbot.2006.12.008","article-title":"Improved salt tolerance of melon (Cucumis melo L) by the addition of proline and potassium nitrate. Environ","volume":"60","author":"Kaya","year":"2007","journal-title":"Exp. Bot."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1007\/BF00018060","article-title":"Rapid determination of free proline for water stress studies","volume":"39","author":"Bates","year":"1973","journal-title":"Plant Soil"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1007\/BF02374789","article-title":"Rapid assay for determination of water-soluble quaternary ammonium compounds","volume":"70","author":"Grieve","year":"1983","journal-title":"Plant Soil"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"2207","DOI":"10.1007\/s11738-012-1021-z","article-title":"Hydrogen sulfide is a mediator in H2O2-induced seed germination in Jatropha curcas","volume":"34","author":"Li","year":"2012","journal-title":"Acta Physiol. Plant."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1016\/S0168-9452(99)00197-1","article-title":"Oxidative stress and some antioxidant system in acid rain treated bean plants: Protective role of exogenous polyamines","volume":"151","author":"Velikova","year":"2000","journal-title":"Plant Sci."},{"key":"ref_53","first-page":"180","article-title":"Photoperoxidation in isolated chloroplasts. I. Kinetics and stoichiometry of fatty acid peroxidation","volume":"125","author":"Heath","year":"1968","journal-title":"Arch. Biochem. Biophys."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0168-9452(98)00025-9","article-title":"Antioxidant responses of rice seedlings to salinity stress","volume":"135","author":"Tobita","year":"1998","journal-title":"Plant Sci."},{"key":"ref_55","first-page":"45","article-title":"Paclobutrazol protects wheat seedlings from heat andparaquat injury Is detoxification of active oxygen involved?","volume":"35","author":"Kraus","year":"1994","journal-title":"Plant Cell Physiol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"764","DOI":"10.1016\/S0076-6879(55)02300-8","article-title":"Assay of catalase and peroxidases","volume":"2","author":"Chance","year":"1955","journal-title":"Method. Enzymol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/S0168-9452(97)00215-X","article-title":"Role of oxidative damage in tulip bulb scale micropropagation","volume":"130","author":"Alberda","year":"1997","journal-title":"Plant Sci."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1016\/0003-2697(76)90527-3","article-title":"A rapid and sensitive method for the quantitation of micro gram quantities of protein utilizing the principle of protein-dye binding","volume":"72","author":"Bradford","year":"1976","journal-title":"Anal. Biochem."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"3041","DOI":"10.1093\/jxb\/eri301","article-title":"Increased sensitivity to salt stress in ascorbate deficient Arabidopsis mutant","volume":"56","author":"Huang","year":"2005","journal-title":"J. Exp. Bot."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"955","DOI":"10.1071\/FP03091","article-title":"Hydrogen peroxide-induced chilling tolerance in mung beans mediated through ABA-independent glutathione accumulation","volume":"30","author":"Yu","year":"2003","journal-title":"Funct. Plant Biol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1016\/j.plantsci.2009.11.002","article-title":"Jasmonic acid regulates ascorbate and glutathione metabolism in Agropyron cristatum leaves under water stress","volume":"178","author":"Shan","year":"2010","journal-title":"Plant Sci."},{"key":"ref_62","first-page":"867","article-title":"Hydrogen peroxide is scavenged by ascorbate specific peroxidase in spinach chloroplasts","volume":"22","author":"Nakano","year":"1981","journal-title":"Plant Cell. Physiol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"1631","DOI":"10.1104\/pp.112.4.1631","article-title":"Acclimation of foliar antioxidant systems to growth irradiance in three broad-leaved evergreen species","volume":"112","author":"Grace","year":"1996","journal-title":"Plant Physiol."},{"key":"ref_64","first-page":"541","article-title":"Thylakoid-bound ascorbate peroxidase in spinach chloroplasts and photoreduction of its primary oxidation product monodehydroascorbate radicals in thylakoids","volume":"33","author":"Miyake","year":"1992","journal-title":"Plant Cell. Physiol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"3811","DOI":"10.1073\/pnas.83.11.3811","article-title":"Enzymatic reactions of ascorbate and glutathione that prevent peroxide damage in soybean root nodules","volume":"83","author":"Dalton","year":"1986","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1007\/s12011-014-0120-7","article-title":"Modulation of antioxidant machinery and the methylglyoxal detoxification system in selenium-supplemented Brassica napus seedlings confers tolerance to high temperature stress","volume":"161","author":"Hasanuzzaman","year":"2014","journal-title":"Biol. Trace. Elem. Res."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"2577","DOI":"10.1007\/s00216-012-6086-4","article-title":"A quick, convenient and economical method for the reliable determination of methylglyoxal in millimolar concentrations: The N-acetyll-cysteine assay","volume":"403","author":"Wild","year":"2012","journal-title":"Anal. Bioanal. Chem."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"60","DOI":"10.1097\/00010694-196201000-00015","article-title":"Methods analysis for Soils, plants and waters","volume":"93","author":"Chapman","year":"1962","journal-title":"Soil Sci."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"771","DOI":"10.1007\/s10725-014-9980-2","article-title":"In vitro rice shoot apices as simple model to study the effect of NaCl and the potential of exogenous proline and glutathione in mitigating salinity stress","volume":"75","author":"Teh","year":"2015","journal-title":"Plant Growth Regul."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1276","DOI":"10.1080\/01904167.2010.484089","article-title":"Potassium sulfate improves water deficit tolerance in melon plants grown under glasshouse conditions","volume":"33","author":"Tuna","year":"2010","journal-title":"J. Plant Nutr."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1080\/01904160500526717","article-title":"Calcium sulfate improves salinity tolerance in rootstocks of plum","volume":"29","author":"Bolat","year":"2006","journal-title":"J. Plant Nutr."},{"key":"ref_72","first-page":"319","article-title":"Responses of the tomato (Lycopersicon esculentum Mill.) plant to exposure to different salt forms and rates","volume":"32","author":"Tuna","year":"2008","journal-title":"Turk. J. Agric. For."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1080\/17429145.2018.1441452","article-title":"Upregulation of antioxidant and glyoxalase systems mitigates NaCl stress in Brassica juncea by supplementation of zinc and calcium","volume":"13","author":"Ahmad","year":"2018","journal-title":"J. Plant Interact."},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Adhikari, B., Dhungana, S.K., Kim, I.D., and Shin, D.H. (2019). Effect of foliar application of potassium fertilizers on soybean plants under salinity stress. J. Saudi Soc. Agric. Sci.","DOI":"10.1016\/j.jssas.2019.02.001"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1016\/j.scienta.2018.01.060","article-title":"Salicylic acid and calcium pretreatments alleviate the toxic effect of salinity in the Oueslati olive variety","volume":"233","author":"Methenni","year":"2018","journal-title":"Sci. Hortic."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"19","DOI":"10.5897\/IJPPB2018.0280","article-title":"Effects of salinity stress on growth in relation to gas exchanges parameters and water status in amaranth (Amaranthus cruentus)","volume":"10","author":"Gandonou","year":"2018","journal-title":"Int. J. Plant Physiol. Biochem."},{"key":"ref_77","first-page":"49","article-title":"Effect of silica nanoparticles on basil (Ocimum basilicum) under salinity stress","volume":"4","author":"Kalteh","year":"2018","journal-title":"J. Chem. Health Risks"},{"key":"ref_78","first-page":"1","article-title":"Exogenous glutathione improves salinity stress tolerance in rice (Oryza sativa L.)","volume":"7","author":"Hussain","year":"2016","journal-title":"Plant Gene Trait."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"plv069","DOI":"10.1093\/aobpla\/plv069","article-title":"Glutathione-induced drought stress tolerance in mung bean: Coordinated roles of the antioxidant defence and methylglyoxal detoxification systems","volume":"7","author":"Nahar","year":"2015","journal-title":"AoB Plants"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"9417","DOI":"10.1007\/s11356-017-8611-7","article-title":"Genotype-dependent alleviation effects of exogenous GSH on salinity stress in cotton is related to improvement in chlorophyll content, photosynthetic performance, and leaf\/root ultrastructure","volume":"24","author":"Ibrahim","year":"2017","journal-title":"Environ. Sci. Pollut. Res."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"2257","DOI":"10.1016\/S2095-3119(18)62068-4","article-title":"Exogenous GSH protects tomatoes against salt stress by modulating photosystem II efficiency, absorbed light allocation and H2O2-scavenging system in chloroplasts","volume":"17","author":"Yan","year":"2018","journal-title":"J. Integr. Agric."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"656","DOI":"10.1104\/pp.109.147975","article-title":"An Oacetylserine (thiol) lyase homolog with l-cysteine desulfhydrase activity regulates cysteine homeostasis in Arabidopsis","volume":"152","author":"Calo","year":"2010","journal-title":"Plant Physiol."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1556\/CRC.2014.0037","article-title":"Hydrogen sulfide stimulates wheat grain germination and counteracts the effect of oxidative damage caused by salinity stress","volume":"43","author":"Ye","year":"2015","journal-title":"Cereal Res. Commun."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"1055","DOI":"10.3389\/fpls.2015.01055","article-title":"Hydrogen sulfide regulates salt tolerance in rice by maintaining Na+\/K+ balance, mineral homeostasis and oxidative metabolism under excessive salt stress","volume":"6","author":"Mostofa","year":"2015","journal-title":"Front. Plant Sci."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"14078","DOI":"10.1038\/srep14078","article-title":"Hydrogen sulfide modulates cadmium-induced physiological and biochemical responses to alleviate cadmium toxicity in rice","volume":"5","author":"Mostofa","year":"2015","journal-title":"Sci. Rep."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"1173","DOI":"10.3389\/fpls.2016.01173","article-title":"Hydrogen sulfide-mediated polyamines and sugar changes are involved in hydrogen sulfide-induced drought tolerance in Spinacia oleracea seedlings","volume":"7","author":"Chen","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1016\/j.jplph.2017.09.010","article-title":"Alleviation of Cr (VI)-induced oxidative stress in maize (Zea mays L.) seedlings by NO and H2S donors through differential organ-dependent regulation of ROS and NADPH-recycling metabolisms","volume":"219","author":"Kharbech","year":"2017","journal-title":"J. Plant Physiol."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"306","DOI":"10.1016\/j.sajb.2016.03.011","article-title":"Effect of salt stress on growth, chlorophyll content, lipid peroxidation and antioxidant defence systems in Phaseolus vulgaris L.","volume":"105","author":"Abderrahim","year":"2016","journal-title":"S. Afr. J. Bot."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"635","DOI":"10.3389\/fpls.2018.00635","article-title":"5-Aminolevulinic Acid (ALA) alleviated salinity stress in cucumber seedlings by enhancing chlorophyll synthesis pathway","volume":"9","author":"Wu","year":"2018","journal-title":"Front. Plant Sci."},{"key":"ref_90","first-page":"167","article-title":"Bioinformatics evaluation of plant chlorophyllase, the key enzyme in chlorophyll degradation","volume":"4","author":"Sharafi","year":"2017","journal-title":"Appl. Food Biotechnol."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1016\/j.bbrc.2016.02.067","article-title":"Co-transforming bar and CsLEA enhanced tolerance to drought and salt stress in transgenic alfalfa (Medicago sativa L.)","volume":"472","author":"Zhang","year":"2016","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"368","DOI":"10.1134\/S1021443717030086","article-title":"Selenium ameliorates salinity stress in Petroselinum crispum by modulation of photosynthesis and by reducing shoot Na accumulation","volume":"64","author":"Habibi","year":"2017","journal-title":"Russ. J. Plant Physiol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"753","DOI":"10.32615\/ps.2019.053","article-title":"How glycine betaine induces tolerance of cucumber plants to salinity stress?","volume":"57","author":"Estaji","year":"2019","journal-title":"Photosynthetica"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"1815","DOI":"10.1093\/pcp\/pcs124","article-title":"Enhancement of non-photochemical quenching in the bryophyte Physcomitrella patens during acclimation to salt and osmotic stress","volume":"53","author":"Azzabi","year":"2012","journal-title":"Plant Cell Physiol."},{"key":"ref_95","first-page":"2637","article-title":"Effects of exogenous GSH on photosynthetic characteristics and expression of key enzyme genes of CO2 assimilation in leaves of tomato seedlings under NaCl stress","volume":"25","author":"Liu","year":"2014","journal-title":"China J. App. Ecol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"254","DOI":"10.1071\/FP15330","article-title":"Salt effects on proline and glycine betaine levels and photosynthetic performance in Melilotus siculus, Tecticornia pergranulata and Thinopyrum ponticum measured in simulated saline conditions","volume":"43","author":"Bhuiyan","year":"2016","journal-title":"Funct. Plant Biol."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"723","DOI":"10.1007\/s10265-017-0929-1","article-title":"Physiological responses of three soybean species (Glycine soja, G. gracilis, and G. max cv. Melrose) to salinity stress","volume":"130","author":"Liu","year":"2017","journal-title":"J. Plant Res."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1016\/j.niox.2016.07.001","article-title":"Melatonin and nitric oxide modulate glutathione content and glutathione reductase activity in sunflower seedling cotyledons accompanying salt stress","volume":"59","author":"Kaur","year":"2016","journal-title":"Nitric Oxide"},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"614","DOI":"10.1016\/j.scienta.2018.06.069","article-title":"Up-regulation of antioxidative defense systems by glycine betaine foliar application in onion plants confer tolerance to salinity stress","volume":"240","author":"Rady","year":"2018","journal-title":"Sci. Hortic."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"1953","DOI":"10.1093\/jxb\/ert055","article-title":"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","volume":"64","author":"Christou","year":"2013","journal-title":"J. Exp. Bot."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"868","DOI":"10.1038\/s41598-017-01046-2","article-title":"Hydrogen sulfide toxicity inhibits primary root growth through the ROS-NO pathway","volume":"7","author":"Zhang","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1007\/s10535-013-0366-5","article-title":"Effects of exogenous hydrogen sulfide on the redox states of ascorbate and glutathione in maize leaves under salt stress","volume":"58","author":"Shan","year":"2014","journal-title":"Biol. Plant."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"256","DOI":"10.1111\/ppl.12976","article-title":"Integrative roles of nitric oxide and hydrogen sulfide in melatonin-induced tolerance of pepper (Capsicum annuum L.) plants to iron deficiency and salt stress alone or in combination","volume":"168","author":"Kaya","year":"2019","journal-title":"Physiol. Plant"},{"key":"ref_104","doi-asserted-by":"crossref","unstructured":"Kumar, V., Wani, S.H., Suprasanna, P., and Tran, L.S.P. (2018). Salinity Stress Tolerance in Crops in Salinity Responses and Tolerance in Plants. Targeting Sensory, Transport and Signaling Mechanisms, Springer.","DOI":"10.1007\/978-3-319-90318-7"},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"347","DOI":"10.3389\/fpls.2016.00347","article-title":"Nitric Oxide Mitigates Salt stress by regulating levels of osmolytes and antioxidant enzymes in chickpea","volume":"7","author":"Ahmad","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1007\/s11738-019-2918-6","article-title":"Exogenous hydrogen sulfide alleviates salt stress by improving antioxidant defenses and the salt overly sensitive pathway in wheat seedlings","volume":"41","author":"Ding","year":"2019","journal-title":"Acta Physiol. Plant."},{"key":"ref_107","first-page":"323","article-title":"Role of glutathione and polyadenylic acid on the oxidative defense systems of two different cultivars of canola seedlings grown under saline conditions","volume":"1","author":"Khattab","year":"2007","journal-title":"Aust. J. Basic Appl. Sci."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1007\/s11104-011-0936-2","article-title":"Hydrogen sulfide enhances alfalfa (Medicago sativa) tolerance against salinity during seed germination by nitric oxide pathway","volume":"351","author":"Wang","year":"2012","journal-title":"Plant Soil"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"613","DOI":"10.3389\/fpls.2017.00613","article-title":"Ascorbic acid-a potential oxidant scavenger and its role in plant development and abiotic stress tolerance","volume":"8","author":"Akram","year":"2017","journal-title":"Front. Plant Sci."},{"key":"ref_110","doi-asserted-by":"crossref","unstructured":"Hossain, M.A., Munn\u00e9-Bosch, S., Burrit, D.J., Diaz-Vivancos, P., Fujita, M., and Lorence, A. (2017). The Role of Plant High-Throughput Phenotyping in the Characterization of the Response of High Ascorbate Plants to Abiotic Stresses. Ascorbic Acid in Plant Growth, Development and Stress Tolerance, Springer.","DOI":"10.1007\/978-3-319-74057-7"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"e1297000","DOI":"10.1080\/15592324.2017.1297000","article-title":"Ethephon increases photosynthetic-nitrogen use efficiency, proline and antioxidant metabolism to alleviate decrease in photosynthesis under salinity stress in mustard","volume":"12","author":"Iqbal","year":"2017","journal-title":"Plant Signal. Behav."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1007\/s12298-016-0371-1","article-title":"Manganese-induced salt stress tolerance in rice seedlings: Regulation of ion homeostasis, antioxidant defense and glyoxalase systems","volume":"22","author":"Rahman","year":"2016","journal-title":"Physiol. Mol. Biol. Plant"},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1556\/0806.45.2017.021","article-title":"Hydrogen sulfide is involved in the regulation of ascorbate-glutathione cycle by exogenous ABA in wheat seedling leaves under osmotic stress","volume":"45","author":"Shan","year":"2017","journal-title":"Cereal Res. Commun."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"2169","DOI":"10.1111\/pbi.13130","article-title":"Pe STZ 1, a C2H2-type zinc finger transcription factor from Populus euphratica, enhances freezing tolerance through modulation of ROS scavenging by directly regulating Pe APX 2","volume":"17","author":"He","year":"2019","journal-title":"Plant Biotechnol. J."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"334","DOI":"10.1104\/pp.15.00009","article-title":"S-sulfhydration: A cysteine posttranslational modification in plant systems","volume":"168","author":"Aroca","year":"2015","journal-title":"Plant Physiol."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1007\/s11032-017-0720-y","article-title":"Cytosolic monodehydroascorbate reductase gene affects stress adaptation and grain yield under paddy field conditions in Oryza sativa L. japonica","volume":"37","author":"Kim","year":"2017","journal-title":"Mol. Breed."},{"key":"ref_117","unstructured":"Marschner, H. (2011). Marschner\u2019s Mineral Nutrition of Higher Plants, Academic Press."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"650","DOI":"10.1016\/j.envexpbot.2018.08.016","article-title":"Long-term exogenous application of melatonin improves nutrient uptake fluxes in apple plants under moderate drought stress","volume":"155","author":"Liang","year":"2018","journal-title":"Environ. Exp. Bot."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"523","DOI":"10.1080\/03650340.2015.1065972","article-title":"Effect of NaCl-induced saline sodicity on the interpretation of soil potassium dynamics","volume":"62","author":"Gul","year":"2016","journal-title":"Arch. Agron. Soil Sci."},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"1787","DOI":"10.3389\/fpls.2016.01787","article-title":"New insights on plant salt tolerance mechanisms and their potential use for breeding","volume":"7","author":"Hanin","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"509","DOI":"10.3389\/fphys.2017.00509","article-title":"The role of Na+ and K+ transporters in salt stress adaptation in glycophytes","volume":"8","author":"Assaha","year":"2017","journal-title":"Front. Physiol."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"352","DOI":"10.21475\/poj.09.05.16.pne232","article-title":"Salinity and drought-induced methylglyoxal detoxification in Brassica spp. and purification of a high active glyoxalase I from tolerant genotype","volume":"9","author":"Hossain","year":"2016","journal-title":"Plant Omics"},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"1341","DOI":"10.3389\/fpls.2016.01341","article-title":"Methylglyoxal: An emerging signaling molecule in plant abiotic stress responses and tolerance","volume":"7","author":"Hoque","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref_124","doi-asserted-by":"crossref","unstructured":"Sankaranarayanan, S., Jamshed, M., Kumar, A., Skori, L., Scandola, S., Wang, T., and Samuel, M.A. (2017). Glyoxalase goes green: The expanding roles of glyoxalase in plants. Int. J. Mol. Sci., 18.","DOI":"10.3390\/ijms18040898"},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"13515","DOI":"10.1038\/s41598-018-31917-1","article-title":"Exogenous application of calcium to 24-epibrassinosteroid pre-treated tomato seedlings mitigates NaCl toxicity by modifying ascorbate-glutathione cycle and secondary metabolites","volume":"8","author":"Ahmad","year":"2018","journal-title":"Sci. Rep."}],"container-title":["Antioxidants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-3921\/9\/7\/603\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T09:49:43Z","timestamp":1760176183000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-3921\/9\/7\/603"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,7,10]]},"references-count":125,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2020,7]]}},"alternative-id":["antiox9070603"],"URL":"https:\/\/doi.org\/10.3390\/antiox9070603","relation":{},"ISSN":["2076-3921"],"issn-type":[{"value":"2076-3921","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,7,10]]}}}