{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,18]],"date-time":"2026-10-18T08:47:01Z","timestamp":1792313221640,"version":"4.4.0"},"reference-count":111,"publisher":"MDPI AG","issue":"8","license":[{"start":{"date-parts":[[2018,7,28]],"date-time":"2018-07-28T00:00:00Z","timestamp":1532736000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["81560622"],"award-info":[{"award-number":["81560622"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Molecules"],"abstract":"<jats:p>Plant hormone candidate melatonin has been widely studied in plants under various stress conditions, such as heat, cold, salt, drought, heavy metal, and pathogen attack. Under stress, melatonin usually accumulates sharply by modulating its biosynthesis and metabolic pathways. Beginning from the precursor tryptophan, four consecutive enzymes mediate the biosynthesis of tryptamine or 5-hydroxytryptophan, serotonin, N-acetylserotonin or 5-methoxytryptamine, and melatonin. Then, the compound is catabolized into 2-hydroxymelatonin, cyclic-3-hydroxymelatonin, and N1-acetyl-N2-formyl-5-methoxyknuramine through 2-oxoglutarate-dependent dioxygenase catalysis or reaction with reactive oxygen species. As an ancient and powerful antioxidant, melatonin directly scavenges ROS induced by various stress conditions. Furthermore, it confreres stress tolerance by activating the plant\u2019s antioxidant system, alleviating photosynthesis inhibition, modulating transcription factors that are involved with stress resisting, and chelating and promoting the transport of heavy metals. Melatonin is even proven to defense against pathogen attacks for the plant by activating other stress-relevant hormones, like salicylic acid, ethylene, and jasmonic acid. Intriguingly, other precursors and metabolite molecules involved with melatonin also can increase stress tolerance for plant except for unconfirmed 5-methoxytryptamine, cyclic-3-hydroxymelatonin, and N1-acetyl-N2-formyl-5-methoxyknuramine. Therefore, the precursors and metabolites locating at the whole biosynthesis and catabolism pathway of melatonin could contribute to plant stress resistance, thus providing a new perspective for promoting plant stress tolerance.<\/jats:p>","DOI":"10.3390\/molecules23081887","type":"journal-article","created":{"date-parts":[[2018,7,30]],"date-time":"2018-07-30T11:55:08Z","timestamp":1532951708000},"page":"1887","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":143,"title":["The Role of Phyto-Melatonin and Related Metabolites in Response to Stress"],"prefix":"10.3390","volume":"23","author":[{"given":"Yang","family":"Yu","sequence":"first","affiliation":[{"name":"State Key Laboratory for Conservation and Utilization of Bio-resources in Yunnan, Yunnan University, Kunming 650091, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yan","family":"Lv","sequence":"additional","affiliation":[{"name":"School of Agriculture, Yunnan University, Kunming 650504, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yana","family":"Shi","sequence":"additional","affiliation":[{"name":"Institute of Medicinal Plants, Yunnan Academy of Agricultural Sciences, Kunming 650205, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tao","family":"Li","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Conservation and Utilization of Bio-resources in Yunnan, Yunnan University, Kunming 650091, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yanchun","family":"Chen","sequence":"additional","affiliation":[{"name":"School of Agriculture, Yunnan University, Kunming 650504, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7792-5138","authenticated-orcid":false,"given":"Dake","family":"Zhao","sequence":"additional","affiliation":[{"name":"Biocontrol Engineering Research Center of Plant Disease &amp; Pest, Yunnan University, Kunming 650504, China"},{"name":"Biocontrol Engineering Research Center of Crop Disease &amp; Pest, Yunnan University, Kunming 650504, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7792-5138","authenticated-orcid":false,"given":"Zhiwei","family":"Zhao","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Conservation and Utilization of Bio-resources in Yunnan, Yunnan University, Kunming 650091, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2018,7,28]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"198","DOI":"10.3389\/fpls.2016.00198","article-title":"Melatonin in plants\u2014Diversity of levels and multiplicity of functions","volume":"7","author":"Hardeland","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"657","DOI":"10.1093\/jxb\/eru332","article-title":"Melatonin promotes ripening and improves quality of tomato fruit during postharvest life","volume":"66","author":"Sun","year":"2015","journal-title":"J. Exp. Bot."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1016\/j.plaphy.2015.06.011","article-title":"Melatonin in Arabidopsis thaliana acts as plant growth regulator at low concentrations and preserves seed viability at high concentrations","volume":"94","author":"Gomez","year":"2015","journal-title":"Plant Physiol. Biochem."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1111\/jpi.12267","article-title":"Melatonin: An ancient molecule that makes oxygen metabolically tolerable","volume":"59","author":"Manchester","year":"2015","journal-title":"J. Pineal Res."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"65","DOI":"10.2307\/3869675","article-title":"Evidence for chilling-induced oxidative stress in maize seedlings and a regulatory role for hydrogen peroxide","volume":"6","author":"Prasad","year":"1994","journal-title":"Plant Cell"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1111\/j.1399-3054.1997.tb04780.x","article-title":"Hydrogen peroxide- and glutathione-associated mechanisms of acclimatory stress tolerance and signalling","volume":"100","author":"Foyer","year":"1997","journal-title":"Physiol. Plant."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Reiter, R., Tan, D., Rosales-Corral, S., Galano, A., Zhou, X., and Xu, B. (2018). Mitochondria: Central organelles for melatonin\u2032s antioxidant and anti-aging actions. Molecules, 23.","DOI":"10.3390\/molecules23020509"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1111\/j.1600-079X.1998.tb00361.x","article-title":"The interaction of melatonin and its precursors with aluminium, cadmium, copper, iron, lead, and zinc: An adsorptive voltammetric study","volume":"24","author":"Janice","year":"1998","journal-title":"J. Pineal Res."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1093\/jxb\/eru336","article-title":"Roles of melatonin in abiotic stress resistance in plants","volume":"66","author":"Zhang","year":"2015","journal-title":"J. Exp. Bot."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Hern\u00e1ndez-Ruiz, J., and Arnao, M. (2018). Relationship of melatonin and salicylic acid in biotic\/abiotic plant stress responses. Agronomy, 8.","DOI":"10.3390\/agronomy8040033"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"18886","DOI":"10.3390\/molecules201018886","article-title":"Melatonin as a potent and inducible endogenous antioxidant: Synthesis and metabolism","volume":"20","author":"Tan","year":"2015","journal-title":"Molecules"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"304","DOI":"10.1111\/j.1600-079X.2010.00841.x","article-title":"Molecular cloning of a plant N-acetylserotonin methyltransferase and its expression characteristics in rice","volume":"50","author":"Kang","year":"2011","journal-title":"J. Pineal Res."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Hardeland, R. (2017). Taxon- and site-specific melatonin catabolism. Molecules, 22.","DOI":"10.3390\/molecules22112015"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1007\/s11738-015-1850-7","article-title":"The levels of melatonin and its metabolites in conditioned corn (Zea mays L.) and cucumber (Cucumis sativus L.) seeds during storage","volume":"37","author":"Szewczyk","year":"2015","journal-title":"Acta Physiol. Plant."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1347","DOI":"10.1016\/j.biotechadv.2016.10.002","article-title":"Serotonin: An ancient molecule and an important regulator of plant processes","volume":"34","author":"Erland","year":"2016","journal-title":"Biotechnol. Adv."},{"key":"ref_16","first-page":"e1538","article-title":"Beyond a neurotransmitter: The role of serotonin in plants","volume":"4","author":"Erland","year":"2017","journal-title":"Neurotransmitter"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"238","DOI":"10.1111\/jpi.12115","article-title":"Role of melatonin in alleviating cold stress in Arabidopsis thaliana","volume":"56","author":"Bajwa","year":"2014","journal-title":"J. Pineal Res."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1111\/jpi.12155","article-title":"The cysteine2\/histidine2-type transcription factor ZINC FINGER OF ARABIDOPSIS THALIANA 6-activated C-REPEAT-BINDING FACTOR pathway is essential for melatonin-mediated freezing stress resistance in Arabidopsis","volume":"57","author":"Haitao","year":"2014","journal-title":"J. Pineal Res."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1016\/j.scienta.2017.03.029","article-title":"Exogenous melatonin ameliorates cold-induced damage in tomato plants","volume":"219","author":"Ding","year":"2017","journal-title":"Sci. Hortic."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Ding, F., Wang, M., Liu, B., and Zhang, S. (2017). Exogenous melatonin mitigates photoinhibition by accelerating non-photochemical quenching in tomato seedlings exposed to moderate light during chilling. Front. Plant Sci., 8.","DOI":"10.3389\/fpls.2017.00244"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Han, Q.-H., Huang, B., Ding, C.-B., Zhang, Z.-W., Chen, Y.-E., Hu, C., Zhou, L.-J., Huang, Y., Liao, J.-Q., and Yuan, S. (2017). Effects of melatonin on anti-oxidative systems and photosystem ll in cold-stressed rice seedlings. Front. Plant Sci., 8.","DOI":"10.3389\/fpls.2017.00785"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"659","DOI":"10.1016\/j.foodchem.2017.10.008","article-title":"Melatonin treatment reduces chilling injury in peach fruit through its regulation of membrane fatty acid contents and phenolic metabolism","volume":"245","author":"Gao","year":"2018","journal-title":"Food Chem."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"40858","DOI":"10.1038\/srep40858","article-title":"Local melatonin application induces cold tolerance in distant organs of Citrullus lanatus L. via long distance transport","volume":"7","author":"Li","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"e12453","DOI":"10.1111\/jpi.12453","article-title":"Melatonin alleviates low PS I-limited carbon assimilation under elevated CO2 and enhances the cold tolerance of offspring in chlorophyll b-deficient mutant wheat","volume":"64","author":"Li","year":"2018","journal-title":"J. Pineal Res."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"4998","DOI":"10.1038\/s41598-017-05267-3","article-title":"Melatonin alleviates chilling stress in cucumber seedlings by up-regulation of CsZat12 and modulation of polyamine and abscisic acid metabolism","volume":"7","author":"Zhao","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1111\/jpi.12219","article-title":"Melatonin induces class A1 heat-shock factors (HSFA1s) and their possible involvement of thermotolerance in Arabidopsis","volume":"58","author":"Shi","year":"2015","journal-title":"J. Pineal Res."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1111\/jpi.12359","article-title":"Melatonin enhances thermotolerance by promoting cellular protein protection in tomato plants","volume":"61","author":"Xu","year":"2016","journal-title":"J. Pineal Res."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Qi, Z.-Y., Wang, K.-X., Yan, M.-Y., Kanwar, K.M., Li, D.-Y., Wijaya, L., Alyemeni, N.M., Ahmad, P., and Zhou, J. (2018). Melatonin alleviates high temperature-induced pollen abortion in Solanum lycopersicum. Molecules, 23.","DOI":"10.3390\/molecules23020386"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"36","DOI":"10.1016\/j.envexpbot.2017.02.012","article-title":"Melatonin suppression of heat-induced leaf senescence involves changes in abscisic acid and cytokinin biosynthesis and signaling pathways in perennial ryegrass (Lolium perenne L.)","volume":"138","author":"Zhang","year":"2017","journal-title":"Environ. Exp. Bot."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Alam, M.N., Zhang, L., Yang, L., Islam, M.R., Liu, Y., Luo, H., Yang, P., Wang, Q., and Chan, Z. (2018). Transcriptomic profiling of tall fescue in response to heat stress and improved thermotolerance by melatonin and 24-epibrassinolide. BMC Genom., 19.","DOI":"10.1186\/s12864-018-4588-y"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1016\/j.freeradbiomed.2017.04.009","article-title":"The AtrbohF-dependent regulation of ROS signaling is required for melatonin-induced salinity tolerance in Arabidopsis","volume":"108","author":"Chen","year":"2017","journal-title":"Free Radic. Biol. Med."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"41236","DOI":"10.1038\/srep41236","article-title":"Chloroplastic biosynthesis of melatonin and its involvement in protection of plants from salt stress","volume":"7","author":"Zheng","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1007\/s11099-015-0140-3","article-title":"Exogenous melatonin improves growth and photosynthetic capacity of cucumber under salinity-induced stress","volume":"54","author":"Wang","year":"2016","journal-title":"Photosynthetica"},{"key":"ref_34","first-page":"295","article-title":"Exogenous melatonin confers salt stress tolerance to watermelon by improving photosynthesis and redox homeostasis","volume":"8","author":"Li","year":"2017","journal-title":"Front. Plant Sci."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1016\/j.freeradbiomed.2017.02.042","article-title":"Melatonin and nitric oxide regulate sunflower seedling growth under salt stress accompanying differential expression of Cu\/Zn SOD and Mn SOD","volume":"106","author":"Arora","year":"2017","journal-title":"Free Radic. Biol. Med."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Chen, Y.-E., Mao, J.-J., Sun, L.-Q., Huang, B., Ding, C.-B., Gu, Y., Liao, J.-Q., Hu, C., Zhang, Z.-W., and Yuan, S. (2018). Exogenous melatonin enhances salt stress tolerance in maize seedlings by improving antioxidant and photosynthetic capacity. Physiol. Plant.","DOI":"10.1111\/ppl.12737"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"408","DOI":"10.1111\/jpi.12180","article-title":"Overexpression of MzASMT improves melatonin production and enhances drought tolerance in transgenic Arabidopsis thaliana plants","volume":"57","author":"Zuo","year":"2014","journal-title":"J. Pineal Res."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"e12429","DOI":"10.1111\/jpi.12429","article-title":"Plant mitochondria synthesize melatonin and enhance the tolerance of plants to drought stress","volume":"63","author":"Wang","year":"2017","journal-title":"J. Pineal Res."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"2033","DOI":"10.1080\/15548627.2015.1098798","article-title":"Tomato HsfA1a plays a critical role in plant drought tolerance by activating ATG genes and inducing autophagy","volume":"11","author":"Wang","year":"2015","journal-title":"Autophagy"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1007\/s11738-015-2045-y","article-title":"Melatonin increased maize (Zea mays L.) seedling drought tolerance by alleviating drought-induced photosynthetic inhibition and oxidative damage","volume":"38","author":"Ye","year":"2016","journal-title":"Acta Physiol. Plant."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"286","DOI":"10.1111\/jac.12201","article-title":"Melatonin may exert a protective role against drought stress in maize","volume":"203","author":"Bonet","year":"2017","journal-title":"J. Agron. Crop Sci."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1016\/j.plaphy.2017.06.014","article-title":"Beneficial effects of melatonin in overcoming drought stress in wheat seedlings","volume":"118","author":"Cui","year":"2017","journal-title":"Plant Physiol. Biochem."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"e12401","DOI":"10.1111\/jpi.12401","article-title":"Melatonin systemically ameliorates drought stress-induced damage in Medicago sativa plants by modulating nitro-oxidative homeostasis and proline metabolism","volume":"62","author":"Antoniou","year":"2017","journal-title":"J. Pineal Res."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"470","DOI":"10.1111\/jpi.12232","article-title":"Coordinated regulation of melatonin synthesis and degradation genes in rice leaves in response to cadmium treatment","volume":"58","author":"Byeon","year":"2015","journal-title":"J. Pineal Res."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"e12441","DOI":"10.1111\/jpi.12441","article-title":"Cadmium-induced melatonin synthesis in rice requires light, hydrogen peroxide, and nitric oxide: Key regulatory roles for tryptophan decarboxylase and caffeic acid O-methyltransferase","volume":"63","author":"Lee","year":"2017","journal-title":"J. Pineal Res."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"e12392","DOI":"10.1111\/jpi.12392","article-title":"Overexpression of rice serotonin N-acetyltransferase 1 in transgenic rice plants confers resistance to cadmium and senescence and increases grain yield","volume":"62","author":"Lee","year":"2017","journal-title":"J. Pineal Res."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"e12387","DOI":"10.1111\/jpi.12387","article-title":"Hsfa1a upregulates melatonin biosynthesis to confer cadmium tolerance in tomato plants","volume":"62","author":"Cai","year":"2017","journal-title":"J. Pineal Res."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"601","DOI":"10.3389\/fpls.2015.00601","article-title":"Melatonin mitigates cadmium phytotoxicity through modulation of phytochelatins biosynthesis, vacuolar sequestration, and antioxidant potential in Solanum lycopersicum L.","volume":"6","author":"Hasan","year":"2015","journal-title":"Front. Plant Sci."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1016\/j.plantsci.2017.05.001","article-title":"Melatonin confers plant tolerance against cadmium stress via the decrease of cadmium accumulation and reestablishment of microRNA-mediated redox homeostasis","volume":"261","author":"Gu","year":"2017","journal-title":"Plant Sci."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1016\/j.jplph.2017.11.003","article-title":"Melatonin pretreatment improves vanadium stress tolerance of watermelon seedlings by reducing vanadium concentration in the leaves and regulating melatonin biosynthesis and antioxidant-related gene expression","volume":"220","author":"Nawaz","year":"2018","journal-title":"J. Plant Physiol."},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Zuo, Z., Sun, L., Wang, T., Miao, P., Zhu, X., Liu, S., Song, F., Mao, H., and Li, X. (2017). Melatonin improves the photosynthetic carbon assimilation and antioxidant capacity in wheat exposed to Nano-ZnO stress. Molecules, 22.","DOI":"10.3390\/molecules22101727"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1111\/jpi.12165","article-title":"Melatonin as a signal molecule triggering defense responses against pathogen attack in Arabidopsis and tobacco","volume":"57","author":"Yool","year":"2014","journal-title":"J. Pineal Res."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"102","DOI":"10.1111\/jpi.12244","article-title":"Melatonin induces nitric oxide and the potential mechanisms relate to innate immunity against bacterial pathogen infection in Arabidopsis","volume":"59","author":"Shi","year":"2015","journal-title":"J. Pineal Res."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"e12379","DOI":"10.1111\/jpi.12379","article-title":"Melatonin is required for H2O2- and NO-mediated defense signaling through MAPKKK3 and OXI1 in Arabidopsis thaliana","volume":"62","author":"Yool","year":"2017","journal-title":"J. Pineal Res."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Zhang, S., Liu, S., Zhang, J., Reiter, R.J., Wang, Y., Qiu, D., Luo, X., Khalid, A.R., Wang, H., and Feng, L. (2018). Synergistic anti-oomycete effect of melatonin with a biofungicide against oomycetic black shank disease. J. Pineal Res., e12492.","DOI":"10.1111\/jpi.12492"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"e12367","DOI":"10.1111\/jpi.12367","article-title":"Identification, transcriptional and functional analysis of heat-shock protein 90s in banana (Musa acuminata L.) highlight their novel role in melatonin-mediated plant response to Fusarium wilt","volume":"62","author":"Wei","year":"2017","journal-title":"J. Pineal Res."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Zhang, S., Zheng, X., Reiter, R.J., Feng, S., Wang, Y., Liu, S., Jin, L., Li, Z., Datla, R., and Ren, M. (2017). Melatonin attenuates potato late blight by disrupting cell growth, stress tolerance, fungicide susceptibility and homeostasis of gene expression in Phytophthora infestans. Front. Plant Sci., 8.","DOI":"10.3389\/fpls.2017.01993"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"e12454","DOI":"10.1111\/jpi.12454","article-title":"Rav transcription factors are essential for disease resistance against cassava bacterial blight via activation of melatonin biosynthesis genes","volume":"64","author":"Wei","year":"2018","journal-title":"J. Pineal Res."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Zhang, H., Liu, X., Chen, T., Ji, Y., Shi, K., Wang, L., Zheng, X., and Kong, J. (2018). Melatonin in apples and juice: Inhibition of browning and microorganism growth in apple juice. Molecules, 23.","DOI":"10.3390\/molecules23030521"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s10327-014-0560-0","article-title":"Tryptamine and sakuranetin accumulation in sekiguchi lesions associated with the light-enhanced resistance of the lesion mimic mutant of rice to Magnaporthe oryzae","volume":"81","author":"Ueno","year":"2015","journal-title":"J. Gen. Plant Pathol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"714","DOI":"10.1111\/ppl.12218","article-title":"Salt stress-induced seedling growth inhibition coincides with differential distribution of serotonin and melatonin in sunflower seedling roots and cotyledons","volume":"152","author":"Mukherjee","year":"2014","journal-title":"Physiol. Plant."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"390","DOI":"10.1007\/BF02147983","article-title":"Protection of vicia faba equina against x-rays by serotonin","volume":"20","author":"Lozeron","year":"1964","journal-title":"Experientia"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1111\/jpi.12314","article-title":"Mitogen-activated protein kinase pathways are required for melatonin-mediated defense responses in plants","volume":"60","author":"Lee","year":"2016","journal-title":"J. Pineal Res."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1111\/jpi.12347","article-title":"2-hydroxymelatonin promotes the resistance of rice plant to multiple simultaneous abiotic stresses (combined cold and drought)","volume":"61","author":"Lee","year":"2016","journal-title":"J. Pineal Res."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"426","DOI":"10.1111\/jpi.12364","article-title":"Melatonin biosynthesis in plants: Multiple pathways catalyze tryptophan to melatonin in the cytoplasm or chloroplasts","volume":"61","author":"Back","year":"2016","journal-title":"J. Pineal Res."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"371","DOI":"10.1111\/jpi.12080","article-title":"Molecular cloning and functional analysis of serotonin N-acetyltransferase from the cyanobacterium Synechocystis sp. PCC 6803","volume":"55","author":"Byeon","year":"2013","journal-title":"J. Pineal Res."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1111\/jpi.12289","article-title":"Cloning and functional characterization of the Arabidopsis N-acetylserotonin O-methyltransferase responsible for melatonin synthesis","volume":"60","author":"Byeon","year":"2016","journal-title":"J. Pineal Res."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1007\/BF00017782","article-title":"Tryptophan decarboxylase from catharanthus roseus cell suspension cultures: Purification, molecular and kinetic data of the homogenous protein","volume":"3","author":"Mollenschott","year":"1984","journal-title":"Plant Mol. Biol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"11308","DOI":"10.1074\/jbc.M109.091371","article-title":"Sekiguchi lesion gene encodes a cytochrome p450 monooxygenase that catalyzes conversion of tryptamine to serotonin in rice","volume":"285","author":"Fujiwara","year":"2010","journal-title":"J. Biol. Chem."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1007\/s00425-007-0614-z","article-title":"Characterization of rice tryptophan decarboxylases and their direct involvement in serotonin biosynthesis in transgenic rice","volume":"227","author":"Kang","year":"2007","journal-title":"Planta"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"1197","DOI":"10.1007\/s00425-009-1015-2","article-title":"Induction of serotonin biosynthesis is uncoupled from the coordinated induction of tryptophan biosynthesis in pepper fruits (Capsicum annuum) upon pathogen infection","volume":"230","author":"Park","year":"2009","journal-title":"Planta"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1111\/jpi.12088","article-title":"Functional analyses of three ASMT gene family members in rice plants","volume":"55","author":"Park","year":"2013","journal-title":"J. Pineal Res."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1111\/jpi.12011","article-title":"Molecular cloning of rice serotonin N-acetyltransferase, the penultimate gene in plant melatonin biosynthesis","volume":"55","author":"Kang","year":"2013","journal-title":"J. Pineal Res."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1111\/jpi.12160","article-title":"Caffeic acid O-methyltransferase is involved in the synthesis of melatonin by methylating N-acetylserotonin in Arabidopsis","volume":"57","author":"Byeon","year":"2014","journal-title":"J. Pineal Res."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"1724","DOI":"10.1096\/fj.06-7745com","article-title":"Novel rhythms of N1-acetyl-N2-formyl-5-methoxykynuramine and its precursor melatonin in water hyacinth: Importance for phytoremediation","volume":"21","author":"Tan","year":"2007","journal-title":"FASEB J."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1111\/jpi.12220","article-title":"Molecular cloning of melatonin 2-hydroxylase responsible for 2-hydroxymelatonin production in rice (Oryza sativa)","volume":"58","author":"Byeon","year":"2015","journal-title":"J. Pineal Res."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"470","DOI":"10.1111\/jpi.12361","article-title":"Molecular cloning of melatonin 3-hydroxylase and its production of cyclic 3-hydroxymelatonin in rice (Oryza sativa)","volume":"61","author":"Lee","year":"2016","journal-title":"J. Pineal Res."},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Shi, H., Chen, K., Wei, Y., and He, C. (2016). Fundamental issues of melatonin-mediated stress signaling in plants. Front. Plant Sci., 7.","DOI":"10.3389\/fpls.2016.01124"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1111\/jpi.12111","article-title":"Melatonin synthesis in rice seedlings in vivo is enhanced at high temperatures and under dark conditions due to increased serotonin N-acetyltransferase and N-acetylserotonin methyltransferase activities","volume":"56","author":"Byeon","year":"2013","journal-title":"J. Pineal Res."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1111\/j.1600-079X.2011.00930.x","article-title":"Tryptamine 5-hydroxylase-deficient sekiguchi rice induces synthesis of 5-hydroxytryptophan and N-acetyltryptamine but decreases melatonin biosynthesis during senescence process of detached leaves","volume":"52","author":"Park","year":"2012","journal-title":"J. Pineal Res."},{"key":"ref_81","first-page":"176","article-title":"Enhanced production of melatonin by ectopic overexpression of human serotonin N-acetyltransferase plays a role in cold resistance in transgenic rice seedlings","volume":"49","author":"Kang","year":"2010","journal-title":"J. Pineal Res."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"1387","DOI":"10.1007\/s00253-010-2994-4","article-title":"Production of serotonin by dual expression of tryptophan decarboxylase and tryptamine 5-hydroxylase in Escherichia coli","volume":"89","author":"Park","year":"2011","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1111\/jpi.12105","article-title":"Changes in melatonin levels in transgenic \u2018Micro-Tom\u2019 tomato overexpressing ovine AANAT and ovine HIOMT genes","volume":"56","author":"Wang","year":"2014","journal-title":"J. Pineal Res."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1111\/jpi.12214","article-title":"Arabidopsis serotonin N-acetyltransferase knockout mutant plants exhibit decreased melatonin and salicylic acid levels resulting in susceptibility to an avirulent pathogen","volume":"58","author":"Lee","year":"2015","journal-title":"J. Pineal Res."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"348","DOI":"10.1111\/jpi.12317","article-title":"Low melatonin production by suppression of either serotonin N-acetyltransferase or N-acetylserotonin methyltransferase in rice causes seedling growth retardation with yield penalty, abiotic stress susceptibility, and enhanced coleoptile growth under anoxic conditions","volume":"60","author":"Byeon","year":"2016","journal-title":"J. Pineal Res."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"12212","DOI":"10.1038\/s41598-017-12566-2","article-title":"Overexpression of ovine AANAT and HIOMT genes in switchgrass leads to improved growth performance and salt-tolerance","volume":"7","author":"Huang","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"856","DOI":"10.1111\/tpj.13299","article-title":"Reactive oxygen species, abiotic stress and stress combination","volume":"90","author":"Choudhury","year":"2017","journal-title":"Plant J."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"20","DOI":"10.14348\/molcells.2016.2324","article-title":"Thiol-based peroxidases and ascorbate peroxidases: Why plants rely on multiple peroxidase systems in the photosynthesizing chloroplast?","volume":"39","author":"Dietz","year":"2016","journal-title":"Mol. Cells"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"1704","DOI":"10.1104\/pp.16.00359","article-title":"Lack of GLYCOLATE OXIDASE1, but not GLYCOLATE OXIDASE2, attenuates the photorespiratory phenotype of CATALASE2-deficient Arabidopsis","volume":"171","author":"Kerchev","year":"2016","journal-title":"Plant Physiol."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"120","DOI":"10.1111\/jpi.12246","article-title":"Comparative physiological and proteomic analyses reveal the actions of melatonin in the reduction of oxidative stress in bermuda grass (Cynodon dactylon (L). Pers.)","volume":"59","author":"Shi","year":"2015","journal-title":"J. Pineal Res."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"1241","DOI":"10.1093\/jxb\/ert430","article-title":"Ros homeostasis in halophytes in the context of salinity stress tolerance","volume":"65","author":"Bose","year":"2014","journal-title":"J. Exp. Bot."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1034\/j.1600-079X.2003.00037.x","article-title":"Mechanistic and comparative studies of melatonin and classic antioxidants in terms of their interactions with the ABTS cation radical","volume":"34","author":"Tan","year":"2003","journal-title":"J. Pineal Res."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"472","DOI":"10.1093\/bja\/aes577","article-title":"Antioxidants that protect mitochondria reduce interleukin-6 and oxidative stress, improve mitochondrial function, and reduce biochemical markers of organ dysfunction in a rat model of acute sepsis","volume":"110","author":"Lowes","year":"2013","journal-title":"Br. J. Anaesth."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"1393","DOI":"10.1211\/0022357011777747","article-title":"Individual and synergistic antioxidative actions of melatonin: Studies with vitamin E, vitamin c, glutathione and desferrrioxamine (desferoxamine) in rat liver homogenates","volume":"53","author":"Gitto","year":"2001","journal-title":"J. Pharm. Pharmacol."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1111\/jpi.12132","article-title":"A review of metal-catalyzed molecular damage: Protection by melatonin","volume":"56","author":"Romero","year":"2014","journal-title":"J. Pineal Res."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1046\/j.1600-079X.2003.00092.x","article-title":"Regulation of antioxidant enzymes: A significant role for melatonin","volume":"36","author":"Rodriguez","year":"2004","journal-title":"J. Pineal Res."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1111\/j.1600-079X.2011.00966.x","article-title":"Delayed senescence of apple leaves by exogenous melatonin treatment: Toward regulating the ascorbate\u2013glutathione cycle","volume":"53","author":"Wang","year":"2012","journal-title":"J. Pineal Res."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1146\/annurev.arplant.59.032607.092759","article-title":"Chlorophyll fluorescence: A probe of photosynthesis in vivo","volume":"59","author":"Baker","year":"2008","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.envexpbot.2017.10.010","article-title":"Interactive effects of melatonin and cytokinin on alleviating drought-induced leaf senescence in creeping bentgrass (Agrostis stolonifera)","volume":"145","author":"Ma","year":"2018","journal-title":"Environ. Exp. Bot."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"334","DOI":"10.1111\/jpi.12262","article-title":"Melatonin induces the transcripts of CBF\/DREB1s and their involvement in both abiotic and biotic stresses in Arabidopsis","volume":"59","author":"Shi","year":"2015","journal-title":"J. Pineal Res."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"698","DOI":"10.1007\/s002990000206","article-title":"Tryptophan is a precursor for melatonin and serotonin biosynthesis in in vitro regenerated St. John\u2019s wort (Hypericum perforatum L. cv. Anthos) plants","volume":"19","author":"Murch","year":"2000","journal-title":"Plant Cell Rep."},{"key":"ref_102","first-page":"921","article-title":"Tryptophan biosynthesis and metabolism: Biochemical and molecular genetics","volume":"7","author":"Radwanski","year":"1995","journal-title":"Plant Cell"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1046\/j.1365-313X.2003.01875.x","article-title":"Increased tryptophan decarboxylase and monoamine oxidase activities induce sekiguchi lesion formation in rice infected with magnaporthe grisea","volume":"36","author":"Ueno","year":"2003","journal-title":"Plant J."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1016\/0006-3002(55)90356-5","article-title":"The effects of 5-hydroxytryptamine, indole-3-acetic acid, and some other substances, on pigment effusion, sodium uptake, and potassium efflux, by slices of red beetroot in vitro","volume":"17","author":"Pickles","year":"1955","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"8535","DOI":"10.1021\/acs.jpcb.5b04920","article-title":"N-acetylserotonin and 6-hydroxymelatonin against oxidative stress: Implications for the overall protection exerted by melatonin","volume":"119","author":"Galano","year":"2015","journal-title":"J. Phys. Chem."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1016\/j.pbi.2010.04.006","article-title":"Comparing signaling mechanisms engaged in pattern-triggered and effector-triggered immunity","volume":"13","author":"Tsuda","year":"2010","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1146\/annurev-arplant-050213-040012","article-title":"Effector-triggered immunity: From pathogen perception to robust defense","volume":"66","author":"Cui","year":"2015","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"344","DOI":"10.1093\/pcp\/pcr181","article-title":"Increasing nitric oxide content in Arabidopsis thaliana by expressing rat neuronal nitric oxide synthase resulted in enhanced stress tolerance","volume":"53","author":"Shi","year":"2012","journal-title":"Plant Cell Physiol."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"3386","DOI":"10.1105\/tpc.104.026609","article-title":"Phosphorylation of 1-aminocyclopropane-1-carboxylic acid synthase by MPK6, a stress-responsive mitogen-activated protein kinase, induces ethylene biosynthesis in Arabidopsis","volume":"16","author":"Liu","year":"2004","journal-title":"Plant Cell"},{"key":"ref_110","doi-asserted-by":"crossref","unstructured":"Mandal Mihir, K., Suren, H., Ward, B., Boroujerdi, A., and Kousik, C. (2018). Differential roles of melatonin in plant-host resistance and pathogen suppression in cucurbits. J. Pineal Res., e12505.","DOI":"10.1111\/jpi.12505"},{"key":"ref_111","doi-asserted-by":"crossref","unstructured":"Weeda, S., Zhang, N., Zhao, X., Ndip, G., Guo, Y., Buck, G.A., Fu, C., and Ren, S. (2014). Arabidopsis transcriptome analysis reveals key roles of melatonin in plant defense systems. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0093462"}],"container-title":["Molecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1420-3049\/23\/8\/1887\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T15:14:56Z","timestamp":1760195696000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1420-3049\/23\/8\/1887"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2018,7,28]]},"references-count":111,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2018,8]]}},"alternative-id":["molecules23081887"],"URL":"https:\/\/doi.org\/10.3390\/molecules23081887","relation":{},"ISSN":["1420-3049"],"issn-type":[{"value":"1420-3049","type":"electronic"}],"subject":[],"published":{"date-parts":[[2018,7,28]]}}}