{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,3]],"date-time":"2026-11-03T20:39:28Z","timestamp":1793738368633,"version":"4.6.1"},"reference-count":110,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2021,8,26]],"date-time":"2021-08-26T00:00:00Z","timestamp":1629936000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Photosynthesis is a pivotal process that determines the synthesis of carbohydrates required for sustaining growth under normal or stress situation. Stress exposure reduces the photosynthetic potential owing to the excess synthesis of reactive oxygen species that disturb the proper functioning of photosynthetic apparatus. This decreased photosynthesis is associated with disturbances in carbohydrate metabolism resulting in reduced growth under stress. We evaluated the importance of melatonin in reducing heat stress-induced severity in wheat (Triticum aestivum L.) plants. The plants were subjected to 25 \u00b0C (optimum temperature) or 40 \u00b0C (heat stress) for 15 days at 6 h time duration and then developed the plants for 30 days. Heat stress led to oxidative stress with increased production of thiobarbituric acid reactive substances (TBARS) and hydrogen peroxide (H2O2) content and reduced accrual of total soluble sugars, starch and carbohydrate metabolism enzymes which were reflected in reduced photosynthesis. Application of melatonin not only reduced oxidative stress through lowering TBARS and H2O2 content, augmenting the activity of antioxidative enzymes but also increased the photosynthesis in plant and carbohydrate metabolism that was needed to provide energy and carbon skeleton to the developing plant under stress. However, the increase in these parameters with melatonin was mediated via hydrogen sulfide (H2S), as the inhibition of H2S by hypotaurine (HT; H2S scavenger) reversed the ameliorative effect of melatonin. This suggests a crosstalk of melatonin and H2S in protecting heat stress-induced photosynthetic inhibition via regulation of carbohydrate metabolism.<\/jats:p>","DOI":"10.3390\/plants10091778","type":"journal-article","created":{"date-parts":[[2021,8,31]],"date-time":"2021-08-31T04:07:28Z","timestamp":1630382848000},"page":"1778","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":109,"title":["The Crosstalk of Melatonin and Hydrogen Sulfide Determines Photosynthetic Performance by Regulation of Carbohydrate Metabolism in Wheat under Heat Stress"],"prefix":"10.3390","volume":"10","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6691-496X","authenticated-orcid":false,"given":"Noushina","family":"Iqbal","sequence":"first","affiliation":[{"name":"Department of Botany, Jamia Hamdard, New Delhi 110062, India"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4569-2082","authenticated-orcid":false,"given":"Mehar","family":"Fatma","sequence":"additional","affiliation":[{"name":"Plant Physiology and Biochemistry Laboratory, Department of Botany, Aligarh Muslim University, Aligarh 202002, India"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4476-5676","authenticated-orcid":false,"given":"Harsha","family":"Gautam","sequence":"additional","affiliation":[{"name":"Plant Physiology and Biochemistry Laboratory, Department of Botany, Aligarh Muslim University, Aligarh 202002, India"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shahid","family":"Umar","sequence":"additional","affiliation":[{"name":"Department of Botany, Jamia Hamdard, New Delhi 110062, India"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0305-308X","authenticated-orcid":false,"given":"Adriano","family":"Sofo","sequence":"additional","affiliation":[{"name":"Department of European and Mediterranean Cultures: Architecture, Environment, Cultural Heritage (DiCEM), University of Basilicata, 75100 Matera, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ilaria","family":"D\u2019ippolito","sequence":"additional","affiliation":[{"name":"Department of European and Mediterranean Cultures: Architecture, Environment, Cultural Heritage (DiCEM), University of Basilicata, 75100 Matera, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7957-5832","authenticated-orcid":false,"given":"Nafees A.","family":"Khan","sequence":"additional","affiliation":[{"name":"Plant Physiology and Biochemistry Laboratory, Department of Botany, Aligarh Muslim University, Aligarh 202002, India"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,8,26]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1080\/11263504.2020.1727987","article-title":"Heat stress in cultivated plants: Nature, impact, mechanisms, and mitigation strategies\u2014A review","volume":"155","author":"Hassan","year":"2021","journal-title":"Plant Biosys."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1016\/j.tplants.2016.08.015","article-title":"Transcriptional regulatory network of plant heat stress response","volume":"22","author":"Ohama","year":"2017","journal-title":"Trends Plant Sci."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1147","DOI":"10.3389\/fpls.2017.01147","article-title":"Crop production under drought and heat stress: Plant responses and management options","volume":"8","author":"Fahad","year":"2017","journal-title":"Front. Plant Sci."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Iqbal, N., Umar, S., Khan, N.A., and Corpas, F.J. (2021). Nitric oxide and hydrogen sulfide coordinately reduce glucose sensitivity and decrease oxidative stress via ascorbate-glutathione cycle in heat-stressed wheat (Triticum aestivum L.) Plants. Antioxidants, 10.","DOI":"10.3390\/antiox10010108"},{"key":"ref_5","unstructured":"Wani, S.H., and Kumar, V. (2020). Alteration in carbohydrate metabolism modulates thermotolerance of plant under heat stress. Heat Stress Tolerance in Plants: Physiological, Molecular and Genetic Perspectives, John Wiley & Sons Ltd."},{"key":"ref_6","first-page":"557","article-title":"Effects of exogenous melatonin on nitrogen metabolism in cucumber seedlings under high temperature stress","volume":"48","author":"Zhao","year":"2012","journal-title":"Plant Physiol. Commun."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Liang, D., Gao, F., Ni, Z., Lin, L., Deng, Q., Tang, Y., and Xia, H. (2018). Melatonin improves heat tolerance in kiwifruit seedlings through promoting antioxidant enzymatic activity and glutathione S-transferase transcription. Molecules, 23.","DOI":"10.3390\/molecules23030584"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1007\/s00709-018-1311-4","article-title":"Melatonin enhances thermotolerance of maize seedlings (Zea mays L.) by modulating antioxidant defense, methylglyoxal detoxification, and osmoregulation systems","volume":"256","author":"Li","year":"2018","journal-title":"Protoplasma"},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Buttar, Z.A., Wu, S.N., Arnao, M.B., Wang, C., Ullah, I., and Wang, C. (2020). Melatonin suppressed the heat stress-induced damage in wheat seedlings by modulating the antioxidant machinery. Plants, 9.","DOI":"10.3390\/plants9070809"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1111\/jpi.12167","article-title":"Melatonin promotes seed germination under high salinity by regulating antioxidant systems, ABA and GA 4 interaction in cucumber (Cucumis sativus L.)","volume":"57","author":"Zhang","year":"2014","journal-title":"J. Pineal Res."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"695","DOI":"10.1093\/jxb\/eru392","article-title":"Melatonin enhances plant growth and abiotic stress tolerance in soybean plants","volume":"66","author":"Wei","year":"2015","journal-title":"J. Exp. Bot."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s11738-008-0213-z","article-title":"Melatonin in plants","volume":"31","author":"Posmyk","year":"2009","journal-title":"Acta Physiol. Plant."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"681","DOI":"10.1093\/jxb\/eru373","article-title":"Comparative physiological, metabolomic, and transcriptomic analyses reveal mechanisms of improved abiotic stress resistance in bermudagrass [Cynodon dactylon (L). Pers.] by exogenous melatonin","volume":"66","author":"Shi","year":"2014","journal-title":"J. Exp. Bot."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Nawaz, K., Chaudhary, R., Sarwar, A., Ahmad, B., Gul, A., Hano, C., and Anjum, S. (2021). Melatonin as master regulator in plant growth, development and stress alleviator for sustainable agricultural production: Current status and future perspectives. Sustainability, 13.","DOI":"10.3390\/su13010294"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1159\/000014635","article-title":"Significance of melatonin in antioxidative defense system: Reactions and products","volume":"9","author":"Tang","year":"2000","journal-title":"Neurosignals"},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Varghese, N., Alyammahi, O., Nasreddine, S., Alhassani, A., and Gururani, M.A. (2019). Melatonin positively influences the photosynthetic machinery and antioxidant system of Avena sativa during salinity stress. Plants, 8.","DOI":"10.3390\/plants8120610"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1093\/aob\/mcx114","article-title":"Melatonin in its relationship to plant hormones","volume":"121","author":"Arnao","year":"2018","journal-title":"Ann. Bot."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Martinez, V., Lopez-Delacalle, M., Rodenas, R., Mestre, T.C., Garcia-Sanchez, F., Rubio, F., Nortes, P.A., Mittler, R., and Rivero, R.M. (2018). Tolerance to stress combinations in tomato plants: New insights in the protective role of melatonin. Molecules, 23.","DOI":"10.3390\/molecules23030535"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"244","DOI":"10.3389\/fpls.2017.00244","article-title":"Exogenous melatonin mitigates photoinhibition by accelerating non-photochemical quenching in tomato seedlings exposed to moderate light during chilling","volume":"8","author":"Ding","year":"2017","journal-title":"Front. Plant Sci."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1111\/jipb.12993","article-title":"Melatonin: A master regulator of plant development and stress responses","volume":"63","author":"Sun","year":"2021","journal-title":"J. Integr. Plant Biol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"627","DOI":"10.1016\/j.chemosphere.2019.03.026","article-title":"Melatonin-mediated nitric oxide improves tolerance to cadmium toxicity by reducing oxidative stress in wheat plants","volume":"225","author":"Kaya","year":"2019","journal-title":"Chemosphere"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1111\/ppl.13143","article-title":"Crosstalk between melatonin and nitric oxide in plant development and stress responses","volume":"170","author":"He","year":"2020","journal-title":"Physiol. Plant."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1111\/nyas.12733","article-title":"Hydrogen sulphide signaling: Interactions with nitric oxide and reactive oxygen species","volume":"1365","author":"Hancock","year":"2016","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.envexpbot.2018.08.034","article-title":"Hydrogen sulfide and environmental stresses","volume":"161","author":"Hancock","year":"2019","journal-title":"Environ. Exp. Bot."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Mukherjee, S., and Bhatla, S.C. (2020). Exogenous melatonin modulates endogenous H2S homeostasis and l-cysteine desulfhydrase activity in salt-stressed tomato (Solanum lycopersicum L. var. cherry) seedling cotyledons. J. Plant Growth Regul.","DOI":"10.1007\/s00344-020-10261-7"},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Li, Z.G., Xiang, R.H., and Wang, J.Q. (2021). Hydrogen sulfide\u2014Phytohormone interaction in plants under physiological and stress conditions. J. Plant Growth Regul.","DOI":"10.1007\/s00344-021-10350-1"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"511","DOI":"10.1111\/ppl.13064","article-title":"Stress responsive gene regulation in relation to hydrogen sulfide in plants under abiotic stress","volume":"168","author":"Pandey","year":"2020","journal-title":"Physiol. Plant."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"2745","DOI":"10.1016\/S2095-3119(16)61358-8","article-title":"Foliar application of sodium hydrosulfide (NaHS), a hydrogen sulfide (H2S) donor, can protect seedlings against heat stress in wheat (Triticum aestivum L.)","volume":"15","author":"Yang","year":"2016","journal-title":"J. Integr. Agric."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"4481","DOI":"10.1093\/jxb\/err145","article-title":"Hydrogen sulphide enhances photosynthesis through promoting chloroplast biogenesis, photosynthetic enzyme expression, and thiol redox modification in Spinacia oleracea seedlings","volume":"62","author":"Chen","year":"2011","journal-title":"J. Exp. Bot."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"943","DOI":"10.1111\/nph.14491","article-title":"Starch as a determinant of plant fitness under abiotic stress","volume":"214","author":"Thalmann","year":"2017","journal-title":"New Phytol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"4433","DOI":"10.1093\/jxb\/erx291","article-title":"Starch as a source, starch as a sink: The bifunctional role of starch in carbon allocation","volume":"68","author":"MacNeill","year":"2017","journal-title":"J. Exp. Bot."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"837","DOI":"10.1104\/pp.104.056374","article-title":"Starch-related alpha-glucan\/water dikinase is involved in the cold-induced development of freezing tolerance in Arabidopsis","volume":"138","author":"Yano","year":"2005","journal-title":"Plant Physiol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1860","DOI":"10.1105\/tpc.16.00143","article-title":"Regulation of leaf starch degradation by abscisic acid is important for osmotic stress tolerance in plants","volume":"28","author":"Thalmann","year":"2016","journal-title":"Plant Cell"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1046\/j.1365-313X.1995.07010097.x","article-title":"Evidence of the crucial role of sucrose synthase for sink strength using transgenic potato plants (Solanum tuberosum L.)","volume":"7","author":"Zrenner","year":"1995","journal-title":"Plant J."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1023\/A:1006327606696","article-title":"Antisense repression of sucrose synthase in carrot (Daucus carota L.) affects growth rather than sucrose partitioning","volume":"41","author":"Tang","year":"1999","journal-title":"Plant Mol. Biol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1593","DOI":"10.1093\/jxb\/err460","article-title":"Drought, salt, and temperature stress-induced metabolic rearrangements and regulatory networks","volume":"63","author":"Krasensky","year":"2012","journal-title":"J. Exp. Bot."},{"key":"ref_37","first-page":"223","article-title":"Alleviation of salinity stress on Viciafaba L. plants via seed priming with melatonin","volume":"20","author":"Dawood","year":"2015","journal-title":"Acta Biol. Colomb."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1016\/j.plaphy.2019.07.012","article-title":"Physiological and iTRAQ-based proteomic analyses reveal that melatonin alleviates oxidative damage in maize leaves exposed to drought stress","volume":"142","author":"Su","year":"2019","journal-title":"Plant Physiol. Biochem."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"150","DOI":"10.3389\/fpls.2019.00150","article-title":"Melatonin-mediated sugar accumulation and growth inhibition in apple plants involves down-regulation of fructokinase 2 expression and activity","volume":"10","author":"Yang","year":"2019","journal-title":"Front. Plant Sci."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1007\/s12571-013-0263-y","article-title":"Crops that feed the world 10. Past successes and future challenges to the role played by wheat in global food security","volume":"5","author":"Shiferaw","year":"2013","journal-title":"Food Secur."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1007\/s13593-017-0443-9","article-title":"Heat stress effects and management in wheat: A review","volume":"37","author":"Akter","year":"2017","journal-title":"Agron. Sustain. Dev."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"997","DOI":"10.1111\/j.1365-2486.2010.02262.x","article-title":"The impact of temperature variability on wheat yields","volume":"17","author":"Asseng","year":"2011","journal-title":"Global Change Biol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1007\/s00122-012-1831-9","article-title":"Genetic dissection of grain yield and physical grain quality in bread wheat (Triticum aestivum L.) under water-limited environments","volume":"125","author":"Bennett","year":"2012","journal-title":"Theor. Appl. Genet."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"518","DOI":"10.1002\/joc.3704","article-title":"Year patterns of climate impact on wheat yields","volume":"34","author":"Yu","year":"2014","journal-title":"Int. J. Climatol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1186\/s13007-020-00565-w","article-title":"Drought and heat stress tolerance screening in wheat using computed tomography","volume":"16","author":"Schmidt","year":"2020","journal-title":"Plant Methods"},{"key":"ref_46","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":"2020","journal-title":"Physiol. Plant."},{"key":"ref_47","first-page":"8","article-title":"Salicylic acid-induced hydrogen sulphide improves lead stress tolerance in pepper plants by upraising the ascorbate-glutathione cycle","volume":"173","author":"Kaya","year":"2021","journal-title":"Physiol. Plant."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1016\/j.jplph.2013.09.018","article-title":"Hydrogen sulfide delays GA-triggered programmed cell death in wheat aleurone layers by the modulation of glutathione homeostasis and heme oxygenase-1 expression","volume":"171","author":"Xie","year":"2014","journal-title":"J. Plant Physiol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1265","DOI":"10.1104\/pp.97.3.1265","article-title":"Abrupt increase in the level of hydrogen peroxide in leaves of winter wheat is caused by cold treatment","volume":"97","author":"Okuda","year":"1991","journal-title":"Plant Physiol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1093\/jxb\/32.1.93","article-title":"Leaf senescence correlated within creased level of membrane permeability, lipid peroxidation and decreased level of SOD and CAT","volume":"32","author":"Dhindsa","year":"1981","journal-title":"J. Exp. Bot."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1104\/pp.59.2.309","article-title":"Superoxide dismutases: I. Occurrence in higher plants","volume":"59","author":"Giannopolitis","year":"1977","journal-title":"Plant Physiol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"559","DOI":"10.1016\/0003-2697(87)90489-1","article-title":"Assaying for superoxide dismutase activity: Some large consequences of minor changes in conditions","volume":"161","author":"Beyer","year":"1987","journal-title":"Anal. Biochem."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1016\/S0076-6879(84)05016-3","article-title":"Catalase in vitro","volume":"105","author":"Aebi","year":"1984","journal-title":"Meth. Enzymol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1007\/BF00386001","article-title":"The presence of glutathione and glutathione reductase in chloroplasts: A proposed role in ascorbic acid metabolism","volume":"133","author":"Foyer","year":"1976","journal-title":"Planta"},{"key":"ref_55","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_56","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1016\/j.envexpbot.2014.05.008","article-title":"Excess sulfur supplementation improves photosynthesis and growth in mustard under salt stress through increased production of glutathione","volume":"107","author":"Fatma","year":"2014","journal-title":"Environ. Exp. Bot."},{"key":"ref_57","first-page":"1455","article-title":"The activation state of ribulose 1, 5-bisphosphate carboxylase in maize leaves in dark and light","volume":"26","author":"Usuda","year":"1985","journal-title":"Plant Cell Physiol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"1982","DOI":"10.1007\/s12010-013-0622-3","article-title":"Functional analysis of the potential enzymes involved in sugar modulation in high and low sugarcane cultivars","volume":"172","author":"Kalwade","year":"2013","journal-title":"Appl. Biochem. Biotechnol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1104\/pp.101.1.179","article-title":"Insensitivity of barley endosperm ADP glucose pyrophosphorylase to 3-phosphoglycerate and orthophosphate regulation","volume":"101","author":"Kleczkowski","year":"1993","journal-title":"Plant Physiol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1016\/j.plantsci.2014.03.010","article-title":"Waterlogging during flowering and boll forming stages affects sucrose metabolism in the leaves subtending the cotton boll and its relationship with boll weight","volume":"223","author":"Kuai","year":"2014","journal-title":"Plant Sci."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1007\/s11738-014-1760-0","article-title":"Drought stress condition increases root to shoot ratio via alteration of carbohydrate partitioning and enzymatic activity in rice seedlings","volume":"37","author":"Xu","year":"2015","journal-title":"Acta Physiol. Plant."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s11104-020-04471-x","article-title":"The role of hydrogen sulfide in plant alleviates heavy metal stress","volume":"449","author":"Luo","year":"2020","journal-title":"Plant Soil"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1016\/j.plaphy.2020.08.014","article-title":"Appraisal of H2S metabolism in Arabidopsis thaliana: In silico analysis at the subcellular level","volume":"155","author":"Palma","year":"2020","journal-title":"Plant Physiol. Biochem."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"110363","DOI":"10.1016\/j.plantsci.2019.110363","article-title":"Physiological response and transcription profiling analysis reveal the role of glutathione in H2S-induced chilling stress tolerance of cucumber seedlings","volume":"291","author":"Liu","year":"2020","journal-title":"Plant Sci."},{"key":"ref_65","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_66","doi-asserted-by":"crossref","first-page":"1257","DOI":"10.1007\/s00709-018-1213-5","article-title":"The roles of H2S and H2O2 in regulating AsA-GSH cycle in the leaves of wheat seedlings under drought stress","volume":"255","author":"Shan","year":"2018","journal-title":"Protoplasma"},{"key":"ref_67","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_68","first-page":"1","article-title":"Hydrogen sulfide (H2S) signaling in plant development and stress responses","volume":"1","author":"Liu","year":"2021","journal-title":"Abiotech"},{"key":"ref_69","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_70","doi-asserted-by":"crossref","first-page":"733","DOI":"10.1134\/S1021443713060058","article-title":"Hydrogen sulfide: A multifunctional gaseous molecule in plants","volume":"60","author":"Li","year":"2013","journal-title":"Russ. J. Plant Physiol."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1016\/j.niox.2014.02.005","article-title":"Hydrogen sulfide in plants: From dissipation of excess sulfur to signaling mole cule","volume":"41","author":"Calderwood","year":"2014","journal-title":"Nitric Oxide"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1007\/s11738-015-1971-z","article-title":"Endogenous hydrogen sulfide regulated by calcium is involved in thermotolerance in tobacco Nicotiana tabacum L. suspension cell cultures","volume":"37","author":"Li","year":"2015","journal-title":"Acta Physiol. Plant."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1016\/j.plantsci.2011.10.006","article-title":"Hydrogen sulfide donor sodium hydrosulfide-induced heat tolerance in tobacco (Nicotiana tabacum L.) suspension cultured cells and involvement of Ca2+ and calmodulin","volume":"185","author":"Li","year":"2012","journal-title":"Plant Sci."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1415","DOI":"10.1007\/s00709-020-01516-x","article-title":"Interplay between hydrogen sulfide and methylglyoxal initiates thermotolerance in maize seedlings by modulating reactive oxidative species and osmolyte metabolism","volume":"257","author":"Ye","year":"2020","journal-title":"Protoplasma"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"595","DOI":"10.1007\/s10535-016-0612-8","article-title":"Hydrogen sulfide as a signal molecule in hematin-induced heat tolerance of tobacco cell suspension","volume":"60","author":"Li","year":"2016","journal-title":"Biol. Plant."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1007\/s11240-015-0939-4","article-title":"Hydrogen sulfide partly mediates abscisic acid-induced heat tolerance in tobacco (Nicotiana tabacum L.) suspension cultured cells","volume":"125","author":"Li","year":"2016","journal-title":"Plant Cell Tiss. Org."},{"key":"ref_77","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_78","doi-asserted-by":"crossref","first-page":"732","DOI":"10.1016\/j.molp.2020.01.004","article-title":"Hydrogen sulfide positively regulates abscisic acid signaling through persulfidation of SnRK2.6 in guard cells","volume":"13","author":"Chen","year":"2020","journal-title":"Mol. Plant"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"1517","DOI":"10.3389\/fpls.2018.01517","article-title":"Ethylene-induced hydrogen sulfide negatively regulates ethylene biosynthesis by persulfidation of ACO in tomato under osmotic stress","volume":"9","author":"Jia","year":"2018","journal-title":"Front. Plant Sci."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1071\/FP20205","article-title":"Hydrogen sulfide induced by hydrogen peroxide mediates brassinosteroid-induced stomatal closure of Arabidopsis thaliana","volume":"48","author":"Ma","year":"2020","journal-title":"Funct. Plant Biol."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"381","DOI":"10.3389\/fpls.2021.650955","article-title":"Melatonin pretreatment confers heat tolerance and repression of heat-induced senescence in tomato through the modulation of ABA-and GA-mediated pathways","volume":"12","author":"Jahan","year":"2021","journal-title":"Front. Plant Sci."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1016\/j.envexpbot.2018.06.006","article-title":"Endogenous melatonin deficiency aggravates high temperature-induced oxidative stress in Solanum lycopersicum L","volume":"161","author":"Ahammed","year":"2019","journal-title":"Environ. Exp. Bot."},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Aftab, T., and Hakeem, K.R. (2021). Melatonin: Role in abiotic stress resistance and tolerance. Plant Growth Regulators, Signalling under Stress Conditions, Springer.","DOI":"10.1007\/978-3-030-61153-8"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1002\/jpln.201400476","article-title":"Melatonin alleviates cold-induced oxidative damage in maize seedlings by up-regulating mineral elements and enhancing antioxidant activity","volume":"178","author":"Turk","year":"2015","journal-title":"J. Plant Nutr. Soil Sci."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1111\/j.1439-037X.1997.tb00486.x","article-title":"Tolerance to drought and temperature stress in relation to increased antioxidant enzyme activity in wheat","volume":"178","author":"Sairam","year":"1997","journal-title":"J. Agron. Crop Sci."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"1837","DOI":"10.1007\/s11738-013-1221-1","article-title":"Effect of short-term heat stress on growth, physiology and antioxidative defence system in wheat seedlings","volume":"35","author":"Gupta","year":"2013","journal-title":"Acta Physiol. Plant."},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Sharif, R., Xie, C., Zhang, H., Arnao, M.B., Ali, M., Ali, Q., Muhammad, I., Shalmani, A., Nawaz, M.A., and Chen, P. (2018). Melatonin and its effects on plant systems. Molecules, 23.","DOI":"10.3390\/molecules23092352"},{"key":"ref_88","doi-asserted-by":"crossref","unstructured":"Arnao, M.B., and Hern\u00e1ndez-Ruiz, J. (2020). Isphytomelatonin a new plant hormone?. Agronomy, 10.","DOI":"10.3390\/agronomy10010095"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1111\/plb.13202","article-title":"Melatonin as a regulatory hub of plant hormone levels and action in stress situations","volume":"23","author":"Arnao","year":"2020","journal-title":"Plant Biol."},{"key":"ref_90","doi-asserted-by":"crossref","unstructured":"Siddiqui, M.H., Alamri, S., Al-Khaishany, M.Y., Khan, M.N., Al-Amri, A., Ali, H.M., and Alsahli, A.A. (2019). Exogenous melatonin counteracts NaCl-induced damage by regulating the antioxidant system, proline and carbohydrates metabolism in tomato seedlings. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20020353"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/j.niox.2018.11.003","article-title":"Recent advancements in the mechanism of nitric oxide signaling associated with hydrogen sulfide and melatonin crosstalk during ethylene-induced fruit ripening in plants","volume":"82","author":"Mukherjee","year":"2019","journal-title":"Nitric Oxide"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"1460","DOI":"10.1104\/pp.103.038323","article-title":"Relationship between the heat tolerance of photosynthesis and the thermal stability of Rubisco activase in plants from contrasting thermal environments","volume":"134","author":"Salvucci","year":"2004","journal-title":"Plant Physiol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"942","DOI":"10.1093\/mp\/ssq044","article-title":"Sugar input, metabolism, and signaling mediated by invertase: Roles in development, yield potential, and response to drought and heat","volume":"3","author":"Ruan","year":"2010","journal-title":"Mol. Plant"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1111\/j.1365-3040.2005.01324.x","article-title":"Effects of moderate heat stress on photosynthesis: Importance of thylakoid reactions, rubisco deactivation, reactive oxygen species, and thermotolerance provided by isoprene","volume":"28","author":"Sharkey","year":"2005","journal-title":"Plant Cell Environ."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1023\/A:1022355525907","article-title":"Chlorophyllase activity and chlorophyll content in wild and mutant plants of Arabidopsis thaliana","volume":"46","author":"Todorov","year":"2003","journal-title":"Biol. Plant."},{"key":"ref_96","doi-asserted-by":"crossref","unstructured":"Jahan, M.S., Shu, S., Wang, Y., Chen, Z., He, M., Tao, M., Sun, J., and Guo, S. (2019). Melatonin alleviates heat-induced damage of tomato seedlings by balancing redox homeostasis and modulating polyamine and nitric oxide biosynthesis. BMC Plant Biol., 19.","DOI":"10.1186\/s12870-019-1992-7"},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1016\/j.jplph.2019.01.007","article-title":"Dynamic changes in the starch-sugar interconversion within plant source and sink tissues promote a better abiotic stress response","volume":"234","author":"Dong","year":"2019","journal-title":"J. Plant Physiol."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1016\/j.jcs.2012.04.002","article-title":"Effects of environmental factors on cereal starch biosynthesis and composition","volume":"56","author":"Thitisaksakul","year":"2012","journal-title":"J. Cereal Sci."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"543","DOI":"10.1093\/jxb\/erz296","article-title":"Heat stress effects on source\u2013sink relationships and metabolome dynamics in wheat","volume":"71","author":"Abdelrahman","year":"2020","journal-title":"J. Exp. Bot."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"12466","DOI":"10.1111\/jpi.12466","article-title":"Melatonin redirects carbohydrates metabolism during sugar starvation in plant cells","volume":"64","author":"Borek","year":"2018","journal-title":"J. Pineal Res."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"E23279","DOI":"10.4161\/psb.23279","article-title":"Exogenous melatonin affects photosynthesis in Characeae Charaaustralis","volume":"8","author":"Murch","year":"2013","journal-title":"Plant Signal. Behav."},{"key":"ref_102","first-page":"1663","article-title":"Melatonin application to Pisum sativum L. seeds positively influences the function of the photosynthetic apparatus in growing seedlings during paraquat-induced oxidative stress","volume":"7","author":"Reiter","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1146\/annurev.arplant.57.032905.105441","article-title":"Sugar sensing and signaling in plants: Conserved and novel mechanisms","volume":"57","author":"Rolland","year":"2006","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"2230","DOI":"10.3389\/fpls.2017.02230","article-title":"UDP-glucose: A potential signaling molecule in plants?","volume":"8","year":"2018","journal-title":"Front. Plant Sci."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"442","DOI":"10.21273\/JASHS.125.4.442","article-title":"Carbohydrate accumulation in relation to heat stress tolerance in two creeping bentgrass cultivar","volume":"125","author":"Liu","year":"2000","journal-title":"J. Am. Soc. Hortic. Sci."},{"key":"ref_106","doi-asserted-by":"crossref","unstructured":"Khan, M.N., Mukherjee, S., Al-Huqail, A.A., Basahi, R.A., Ali, H.M., Al-Munqedhi, B.M.A., Siddiqui, M.H., and Kalaji, H.M. (2021). Exogenous Potassium (K+) Positively regulates Na+\/H+ antiport system, carbohydrate metabolism, and ascorbate\u2014Glutathione cycle in H2S-dependent manner in NaCl-stressed tomato seedling roots. Plants, 10.","DOI":"10.3390\/plants10050948"},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1016\/j.postharvbio.2011.05.008","article-title":"Low temperature induced changes in activity and protein levels of the enzymes associated to conversion of starch to sucrose in banana fruit","volume":"62","author":"Soares","year":"2011","journal-title":"Postharvest Biol. Technol."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1111\/jpi.12258","article-title":"Unveiling the mechanism of melatonin impacts on maize seedling growth: Sugar metabolism as a case","volume":"59","author":"Zhao","year":"2015","journal-title":"J. Pineal Res."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1104\/pp.109.2.637","article-title":"Effect of high temperature on plant growth and carbohydrate metabolism in potato","volume":"109","author":"Lafta","year":"1995","journal-title":"Plant Physiol."},{"key":"ref_110","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."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/10\/9\/1778\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T06:52:27Z","timestamp":1760165547000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/10\/9\/1778"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,8,26]]},"references-count":110,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2021,9]]}},"alternative-id":["plants10091778"],"URL":"https:\/\/doi.org\/10.3390\/plants10091778","relation":{"has-preprint":[{"id-type":"doi","id":"10.20944\/preprints202106.0412.v1","asserted-by":"object"}]},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,8,26]]}}}