{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,30]],"date-time":"2026-10-30T22:13:47Z","timestamp":1793398427364,"version":"4.5.0"},"reference-count":48,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T00:00:00Z","timestamp":1785715200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"<jats:sec>\n                    <jats:title>Introduction<\/jats:title>\n                    <jats:p>Soil salinization constrains plant growth, and enhancing salt tolerance is of great significance.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      Here,\n                      <jats:italic>Ginkgo biloba<\/jats:italic>\n                      seedlings were treated with control (CK), salt stress (S), or salt stress + melatonin (S+M). Integrating physiology, transcriptomics, and weighted gene co-expression network analysis (WGCNA), we explored how melatonin enhances salt tolerance.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results and Discussion<\/jats:title>\n                    <jats:p>\n                      Salt stress reduced chlorophyll content, suggesting potential impairment of photosynthetic capacity, appeared to increase membrane lipid peroxidation, and led to substantial proline accumulation. Melatonin treatment appeared to up-regulate the MAPK gene\n                      <jats:italic>MYC2<\/jats:italic>\n                      and down-regulate\n                      <jats:italic>PP2C<\/jats:italic>\n                      and\n                      <jats:italic>RbohD<\/jats:italic>\n                      , suggesting a potential alleviation of oxidative stress. WGCNA identified MEblue (MAPK-enriched) positively correlated with salt stress and proline, and MEgreen (photosynthesis-enriched) positively correlated with chlorophyll and catalase (CAT) activity but negatively correlated with salt stress. Melatonin appeared to help restore MEgreen expression, potentially reducing oxidative damage and appearing to help stabilize chlorophyll. Melatonin also appeared to help maintain photosystem II (PSII) genes (\n                      <jats:italic>Lhcb1<\/jats:italic>\n                      ,\n                      <jats:italic>Lhcb4<\/jats:italic>\n                      ) and reduced flavonoid accumulation. These findings provide insights into the model of melatonin-mediated salt stress responses in gymnosperms. This work may support salt-tolerant\n                      <jats:italic>Ginkgo biloba<\/jats:italic>\n                      breeding and the application of melatonin in forestry.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fpls.2026.1881953","type":"journal-article","created":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T04:27:43Z","timestamp":1785731263000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":1,"title":["Physiological and transcriptomic responses of Ginkgo biloba to salt stress and exogenous melatonin"],"prefix":"10.3389","volume":"17","author":[{"given":"Xinyu","family":"Yang","sequence":"first","affiliation":[{"name":"Key Laboratory of National Forestry and Grassland Administration for Warm Temperate Forest Ecosystem Conservation and Restoration, Forestry College of Shandong Agricultural University","place":["Tai\u2019an, Shandong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shurong","family":"Peng","sequence":"additional","affiliation":[{"name":"Key Laboratory of National Forestry and Grassland Administration for Warm Temperate Forest Ecosystem Conservation and Restoration, Forestry College of Shandong Agricultural University","place":["Tai\u2019an, Shandong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yanxian","family":"Zhao","sequence":"additional","affiliation":[{"name":"Tai\u2019an 88th Hospital","place":["Tai\u2019an, Shandong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dongmei","family":"Shi","sequence":"additional","affiliation":[{"name":"Weifang Engineering Vocational College","place":["Weifang, Shandong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fusheng","family":"Wu","sequence":"additional","affiliation":[{"name":"Shandong Provincial Center for Forest and Grassland Germplasm Resources","place":["Jinan, Shandong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Limin","family":"Sun","sequence":"additional","affiliation":[{"name":"Key Laboratory of National Forestry and Grassland Administration for Warm Temperate Forest Ecosystem Conservation and Restoration, Forestry College of Shandong Agricultural University","place":["Tai\u2019an, Shandong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qian","family":"Zhang","sequence":"additional","affiliation":[{"name":"Shandong Academy of Forestry Sciences","place":["Jinan, Shandong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,8,3]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.3390\/plants15010171","article-title":"Flavonoids in plant salt stress responses: Biosynthesis, regulation, functions, and signaling networks","volume":"15","author":"Akhtar","year":"2026","journal-title":"Plants."},{"key":"B2","doi-asserted-by":"publisher","first-page":"1138","DOI":"10.1111\/nph.13550","article-title":"Hydrogen peroxide produced by NADPH oxidases increases proline accumulation during salt or mannitol stress in Arabidopsis thaliana","volume":"208","author":"Ben Rejeb","year":"2015","journal-title":"New Phytol."},{"key":"B3","doi-asserted-by":"publisher","first-page":"379","DOI":"10.1104\/pp.112.202408","article-title":"Unique drought resistance functions of the highly ABA-induced clade A protein phosphatase 2Cs","volume":"160","author":"Bhaskara","year":"2012","journal-title":"Plant Physiol."},{"key":"B4","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1002\/PPP3.7","article-title":"An evolutionary and cultural biography of Ginkgo biloba","volume":"1","author":"Crane","year":"2019","journal-title":"Plants People Planet"},{"key":"B5","doi-asserted-by":"publisher","DOI":"10.1016\/j.jplph.2022.153862","article-title":"Overexpression of SmMYC2 enhances salt resistance in Arabidopsis thaliana and Salvia miltiorrhiza hairy roots","volume":"280","author":"Deng","year":"2023","journal-title":"J. Plant Physiol."},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1016\/j.jphotobiol.2024.113004","article-title":"The mechanisms of photoinhibition and repair in plants under high light conditions and interplay with abiotic stressors","volume":"259","author":"Didaran","year":"2024","journal-title":"J. Photochem. Photobiol. B. Biol."},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.1016\/j.indcrop.2025.121492","article-title":"Targeted-metabolome and transcriptome analysis revealed the mechanisms by which jasmonate and flavonoids regulate Ginkgo biloba adaptation to high-dose NaCl stress","volume":"234","author":"Feng","year":"2025","journal-title":"Ind. Crops Prod."},{"key":"B8","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/978-981-10-9044-8_18","article-title":"Ion homeostasis and antioxidant defense toward salt tolerance in plants","volume-title":"Plant Nutrients and Abiotic Stress Tolerance","author":"Garc\u00eda-Caparr\u00f3s","year":"2018"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.3390\/ijms25042096","article-title":"IbMYC2 contributes to salt and drought stress tolerance via modulating anthocyanin accumulation and ROS-scavenging system in sweet potato","volume":"25","author":"Hu","year":"2024","journal-title":"Int. J. Mol. Sci."},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.1016\/j.plaphy.2024.108507","article-title":"Stress salinity in plants: New strategies to cope with in the foreseeable scenario","volume":"208","author":"Hualpa-Ramirez","year":"2024","journal-title":"Plant Physiol. Biochem."},{"key":"B11","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1016\/j.plaphy.2021.01.029","article-title":"Recent progress in understanding salinity tolerance in plants: Story of Na+\/K+ balance and beyond","volume":"160","author":"Hussain","year":"2021","journal-title":"Plant Physiol. Biochem."},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1016\/j.envexpbot.2024.105942","article-title":"Melatonin: Discovery, biosynthesis, phytohormones crosstalk, and roles in agricultural crops under abiotic stress conditions","volume":"226","author":"Jindal","year":"2024","journal-title":"Environ. Exp. Bot."},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.3390\/antiox11081534","article-title":"Role of plasma membrane NADPH oxidase in response to salt stress in cucumber seedlings","volume":"11","author":"Kaba\u0142a","year":"2022","journal-title":"Antioxidants."},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.1016\/j.scienta.2023.112664","article-title":"Foliar treatment with melatonin modulates photosynthetic and antioxidant responses in Silybum marianum L. under salt stress","volume":"325","author":"Kang","year":"2024","journal-title":"Sci. Hortic."},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkae909","article-title":"KEGG: biological systems database as a model of the real world","volume":"53","author":"Kanehisa","year":"2025","journal-title":"Nucleic Acids Res."},{"key":"B16","doi-asserted-by":"publisher","first-page":"1040","DOI":"10.1111\/nph.17376","article-title":"Under salt stress guard cells rewire ion transport and abscisic acid signaling","volume":"231","author":"Karimi","year":"2021","journal-title":"New Phytol."},{"key":"B17","doi-asserted-by":"publisher","first-page":"2587","DOI":"10.1021\/ja01543a060","article-title":"Isolation of melatonin, the pineal gland factor that lightens melanocytes","volume":"80","author":"Lerner","year":"1958","journal-title":"J. Am. Chem. Soc"},{"key":"B18","doi-asserted-by":"publisher","first-page":"591","DOI":"10.1042\/BST0110591","article-title":"Determinations of total carotenoids and chlorophylls a and b of leaf extracts in different solvents","volume":"11","author":"Lichtenthaler","year":"1983","journal-title":"Biochem. Soc Trans."},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.3390\/ijms22041543","article-title":"Plant mitogen-activated protein kinase cascades in environmental stresses","volume":"22","author":"Lin","year":"2021","journal-title":"Int. J. Mol. Sci."},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.3390\/horticulturae10111221","article-title":"Plant responses and adaptations to salt stress: A review","volume":"10","author":"Liu","year":"2024","journal-title":"Horticulturae"},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.1016\/j.indcrop.2022.115576","article-title":"Identification and characterization of thirteen gene families involved in flavonoid biosynthesis in Ginkgo biloba","volume":"188","author":"Liu","year":"2022","journal-title":"Ind. Crops Prod."},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.1016\/j.jep.2022.115645","article-title":"Leaves, seeds and exocarp of Ginkgo biloba L. (Ginkgoaceae): A comprehensive review of traditional uses, phytochemistry, pharmacology, resource utilization and toxicity","volume":"298","author":"Liu","year":"2022","journal-title":"J. Ethnopharmacol."},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.1016\/j.jhazmat.2025.138036","article-title":"Boron-induced phenylpropanoid metabolism, Na+\/K+ homeostasis and antioxidant defense mechanisms in salt-stressed Soybean seedlings","volume":"491","author":"Lu","year":"2025","journal-title":"J. Hazard. Mater."},{"key":"B24","doi-asserted-by":"publisher","first-page":"538","DOI":"10.1016\/j.molp.2025.12.009","article-title":"Plant salt-tolerance mechanisms: Classic signaling pathways, emerging frontiers, and future perspectives","volume":"19","author":"Ma","year":"2026","journal-title":"Mol. Plant"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2026.1746328","article-title":"Evaluation of the photosynthetic response of Ginkgo biloba as an urban tree to air pollution, soil salinity, and excess humidity","volume":"17","author":"Matsuura","year":"2026","journal-title":"Front. Plant Sci."},{"key":"B26","doi-asserted-by":"publisher","DOI":"10.3390\/molecules25225359","article-title":"Role of melatonin in plant tolerance to soil stressors: Salinity, pH and heavy metals","volume":"25","author":"Moustafa-Farag","year":"2020","journal-title":"Molecules."},{"key":"B27","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1186\/s12284-025-00825-0","article-title":"Melatonin enhances antioxidant defense systems and stress tolerance in plants under variable environmental conditions","volume":"18","author":"Muhammad","year":"2025","journal-title":"Rice."},{"key":"B28","doi-asserted-by":"publisher","first-page":"1446","DOI":"10.21273\/HORTSCI15058-20","article-title":"Cell membrane stability and relative cell injury in response to heat stress during early and late seedling stages of diverse carrot (Daucus carota L.) germplasm","volume":"55","author":"Nijabat","year":"2020","journal-title":"HortScience"},{"key":"B29","doi-asserted-by":"publisher","first-page":"7396","DOI":"10.3390\/molecules20047396","article-title":"Phytomelatonin: Assisting plants to survive and thrive","volume":"20","author":"Reiter","year":"2015","journal-title":"Molecules."},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.1111\/ppl.70402","article-title":"Unveiling the plant growth regulators crosstalk in agricultural crop response to salinity-stress: A concise review","volume":"177","author":"Shafi","year":"2025","journal-title":"Physiol. Plant"},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.1016\/j.jplph.2026.154743","article-title":"The crucial roles of MAPK pathways in sophisticated salt stress responses in plants","volume":"319","author":"Shi","year":"2026","journal-title":"J. Plant Physiol."},{"key":"B32","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1016\/j.sajb.2023.05.034","article-title":"Linking the role of melatonin in plant stress acclimatization","volume":"159","author":"Singh","year":"2023","journal-title":"S. Afr. J. Bot."},{"key":"B33","doi-asserted-by":"publisher","first-page":"961","DOI":"10.1007\/s00299-023-03013-w","article-title":"Abscisic acid in plants under abiotic stress: Crosstalk with major phytohormones","volume":"42","author":"Singh","year":"2023","journal-title":"Plant Cell Rep."},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.3390\/plants13071025","article-title":"Changes in photosystem II complex and physiological activities in Pea and Maize plants in response to salt stress","volume":"13","author":"Stefanov","year":"2024","journal-title":"Plants."},{"key":"B35","doi-asserted-by":"publisher","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":"B36","doi-asserted-by":"publisher","first-page":"178","DOI":"10.1016\/j.tplants.2008.01.005","article-title":"How do environmental stresses accelerate photoinhibition","volume":"13","author":"Takahashi","year":"2008","journal-title":"Trends Plant Sci."},{"key":"B37","doi-asserted-by":"publisher","first-page":"5604","DOI":"10.1111\/pce.15544","article-title":"Molecular insights into salt stress adaptation in plants","volume":"48","author":"Tibesigwa","year":"2025","journal-title":"Plant Cell Environ."},{"key":"B38","doi-asserted-by":"publisher","first-page":"2560","DOI":"10.1111\/febs.15157","article-title":"A bHLH transcription factor, MYC2, imparts salt intolerance by regulating proline biosynthesis in Arabidopsis","volume":"287","author":"Verma","year":"2020","journal-title":"FEBS J."},{"key":"B39","doi-asserted-by":"publisher","first-page":"804","DOI":"10.1016\/j.hpj.2023.12.004","article-title":"The critical role of CmCIPK1-CmRbohD1\/D2 complexes in generating H2O2 signals for enhancing salt tolerance in pumpkin","volume":"11","author":"Wei","year":"2025","journal-title":"Hortic. Plant J."},{"key":"B40","doi-asserted-by":"publisher","first-page":"470","DOI":"10.1186\/s12870-024-05187-1","article-title":"Melatonin\u00a0enhances resistance to Botryosphaeria dothidea in pear by promoting jasmonic acid and phlorizin biosynthesis","volume":"24","author":"Xu","year":"2024","journal-title":"BMC Plant Biol."},{"key":"B41","doi-asserted-by":"publisher","first-page":"689","DOI":"10.1007\/s10725-024-01138-w","article-title":"Integrative analyses of transcriptome, microRNA-seq and metabolome reveal insights into exogenous melatonin-mediated salt tolerance during seed germination of maize","volume":"103","author":"Xu","year":"2024","journal-title":"Plant Growth Regul."},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2025.1658091","article-title":"Integrated physiological and transcriptomic analyses elucidate the molecular mechanisms of exogenous melatonin-mediated salt tolerance in pomegranate (Punica granatum L.)","volume":"16","author":"Yang","year":"2025","journal-title":"Front. Plant Sci."},{"key":"B43","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1016\/j.plaphy.2022.05.033","article-title":"Exogenous melatonin stimulated isoflavone biosynthesis in NaCl-stressed germinating soybean (Glycine max L.)","volume":"185","author":"Yin","year":"2022","journal-title":"Plant Physiol. Biochem."},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.1016\/j.xinn.2020.100017","article-title":"Mechanisms of plant responses and adaptation to soil salinity","volume":"1","author":"Zhao","year":"2020","journal-title":"Innovation"},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.3390\/horticulturae10010089","article-title":"Effects of exogenous melatonin on the growth and physiological characteristics of Ginkgo biloba L. under salinity stress conditions","volume":"10","author":"Zhou","year":"2024","journal-title":"Horticulturae"},{"key":"B46","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1016\/j.jgg.2023.08.007","article-title":"Insights into plant salt stress signaling and tolerance","volume":"51","author":"Zhou","year":"2024","journal-title":"J. Genet. Genomics"},{"key":"B47","doi-asserted-by":"publisher","DOI":"10.3390\/ijms252010940","article-title":"Understanding of plant salt tolerance mechanisms and application to molecular breeding","volume":"25","author":"Zhou","year":"2024","journal-title":"Int. J. Mol. Sci."},{"key":"B48","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1134\/S1021443724603689","article-title":"Mechanism of melatonin promotion of flavonoid biosynthesis in Ginkgo biloba","volume":"71","author":"Zu","year":"2024","journal-title":"Russ. J. Plant Physiol."}],"container-title":["Frontiers in Plant Science"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1881953\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T04:27:44Z","timestamp":1785731264000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1881953\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,3]]},"references-count":48,"alternative-id":["10.3389\/fpls.2026.1881953"],"URL":"https:\/\/doi.org\/10.3389\/fpls.2026.1881953","relation":{},"ISSN":["1664-462X"],"issn-type":[{"value":"1664-462X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,3]]},"article-number":"1881953"}}