{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T17:56:00Z","timestamp":1791482160544,"version":"4.3.4"},"reference-count":39,"publisher":"Public Library of Science (PLoS)","issue":"10","license":[{"start":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T00:00:00Z","timestamp":1791417600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS One"],"abstract":"<jats:p>\n                    Biostimulant-induced responses in plants are dynamic, involving rapid signaling events followed by slower, downstream transcriptional and metabolic adjustments. To characterize the temporal molecular responses of maize to two biostimulant formulations, including fulvic acid (BLG) and mixture of fulvic acid and Quillaja extract (BLGQ), RNA sequencing was performed on leaf tissues collected at two time points representing the early (3 h) and later (27 h) transcriptional responses following biostimulant application. Differential gene expression, functional enrichment, and transcription factor analyses were subsequently conducted to compare the two treatments with untreated controls. Early responses revealed rapid activation of hormone signaling, ion transport, proteostasis, and transcription factor (TF) networks, with BLGQ inducing broader and stronger transcriptional changes compared to BLG. Notably, TF families such as\n                    <jats:italic>ERF<\/jats:italic>\n                    ,\n                    <jats:italic>NAC<\/jats:italic>\n                    ,\n                    <jats:italic>C2H2<\/jats:italic>\n                    ,\n                    <jats:italic>WRKY<\/jats:italic>\n                    , and\n                    <jats:italic>MYB-related<\/jats:italic>\n                    were prominently upregulated, suggesting early transcriptional activation of genes associated withstress, growth, and developmental pathways. Later responses showed that BLG predominantly reinforced broad metabolic, redox, and transcriptional regulatory networks, whereas BLGQ emphasized signaling integration, vesicle trafficking, and secondary metabolism. Differential transcription factor modulation further highlighted stimulus- and time-specific strategies: BLG sustained upregulation of\n                    <jats:italic>MYB-related<\/jats:italic>\n                    ,\n                    <jats:italic>NAC<\/jats:italic>\n                    ,\n                    <jats:italic>GRAS<\/jats:italic>\n                    , and\n                    <jats:italic>ERF<\/jats:italic>\n                    TFs, while BLGQ selectively balanced upregulation and downregulation of\n                    <jats:italic>NF-YB<\/jats:italic>\n                    ,\n                    <jats:italic>HSF<\/jats:italic>\n                    , and\n                    <jats:italic>bZIP<\/jats:italic>\n                    families to fine-tune signaling and specialized pathways. These transcriptomic results suggest that maize responses to biostimulants are highly dynamic and stimulus-specific, with BLG response associated with broader transcriptional changes in genes involved in general adaptive processes, whereas BLGQ response associated with more specific signaling and regulatory pathways. Our results provide transcriptomic insight into the temporal layers of biostimulant action and underline the importance of time-resolved analyses for understanding plant growth and stress adaptation.\n                  <\/jats:p>","DOI":"10.1371\/journal.pone.0360081","type":"journal-article","created":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T17:28:37Z","timestamp":1791480517000},"page":"e0360081","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":0,"title":["Temporal and stimulus-specific transcriptional responses of maize to a biostimulant mixture of quillaja extract and fulvic acid"],"prefix":"10.1371","volume":"21","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5289-1835","authenticated-orcid":true,"given":"Mohsen","family":"Hesami","sequence":"first","affiliation":[{"name":"University of Guelph","place":["Guelph, ON, Canada"],"department":["Department of Plant Agriculture"]},{"name":"BioLiNE\u00ae Corporation","place":["Alvinston, ON, Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mohammad","family":"Rahbari","sequence":"additional","affiliation":[{"name":"BioLiNE\u00ae Corporation","place":["Alvinston, ON, Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ron A.","family":"Salzman","sequence":"additional","affiliation":[{"name":"Texas A&M University","place":["Texas, United States of America"],"department":["Department of Biochemistry and Biophysics"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2026,10,8]]},"reference":[{"key":"pone.0360081.ref001","first-page":"100032","article-title":"A review on drought stress in plants: Implications, mitigation and the role of plant growth promoting rhizobacteria","volume":"5","author":"O Ahluwalia","year":"2021","journal-title":"Resour Environ Sustain"},{"key":"pone.0360081.ref002","doi-asserted-by":"crossref","first-page":"103634","DOI":"10.1016\/j.agsy.2023.103634","article-title":"Challenges and potential pathways towards sustainable agriculture within the European Green Deal","volume":"207","author":"C Boix-Fayos","year":"2023","journal-title":"Agric Syst"},{"issue":"3","key":"pone.0360081.ref003","doi-asserted-by":"crossref","first-page":"659","DOI":"10.1007\/s00344-023-11144-3","article-title":"Biostimulants: The futuristic sustainable approach for alleviating crop productivity and abiotic stress tolerance","volume":"43","author":"R Johnson","year":"2023","journal-title":"J Plant Growth Regul"},{"key":"pone.0360081.ref004","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1016\/j.scienta.2015.09.013","article-title":"Humic and fulvic acids as biostimulants in horticulture","volume":"196","author":"LP Canellas","year":"2015","journal-title":"Sci Hortic"},{"key":"pone.0360081.ref005","doi-asserted-by":"crossref","first-page":"101210","DOI":"10.1016\/j.stress.2025.101210","article-title":"Biochemical insights into fulvic acid-driven drought tolerance in canola","volume":"19","author":"M Rahbari","year":"2026","journal-title":"Plant Stress"},{"issue":"1","key":"pone.0360081.ref006","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1146\/annurev-food-032818-122010","article-title":"Quillaja saponin characteristics and functional properties","volume":"10","author":"CL Reichert","year":"2019","journal-title":"Annu Rev Food Sci Technol"},{"key":"pone.0360081.ref007","doi-asserted-by":"crossref","first-page":"1439747","DOI":"10.3389\/fpls.2024.1439747","article-title":"Integrative transcriptome and metabolome analysis reveals the mechanism of fulvic acid alleviating drought stress in oat","volume":"15","author":"S Zhu","year":"2024","journal-title":"Front Plant Sci"},{"key":"pone.0360081.ref008","doi-asserted-by":"crossref","first-page":"983772","DOI":"10.3389\/fpls.2022.983772","article-title":"A dual-omics approach for profiling plant responses to biostimulant applications under controlled and field conditions","volume":"13","author":"A Baghdadi","year":"2022","journal-title":"Front Plant Sci"},{"key":"pone.0360081.ref009","doi-asserted-by":"crossref","first-page":"583888","DOI":"10.3389\/fgene.2021.583888","article-title":"Transcriptomics of biostimulation of plants under abiotic stress","volume":"12","author":"S Gonz\u00e1lez-Morales","year":"2021","journal-title":"Front Genet"},{"issue":"2","key":"pone.0360081.ref010","doi-asserted-by":"crossref","first-page":"1609","DOI":"10.1007\/s42729-024-01687-z","article-title":"Maize grain yield and quality improvement through biostimulant application: A systematic review","volume":"24","author":"A Ocwa","year":"2024","journal-title":"J Soil Sci Plant Nutr"},{"issue":"6","key":"pone.0360081.ref011","doi-asserted-by":"crossref","DOI":"10.1111\/ppl.70007","article-title":"Antioxidant activity and comparative RNA-seq analysis support mitigating effects of an algae-based biostimulant on drought stress in tomato plants","volume":"176","author":"P Cerruti","year":"2024","journal-title":"Physiol Plant"},{"issue":"8","key":"pone.0360081.ref012","doi-asserted-by":"crossref","first-page":"1096","DOI":"10.3390\/biom11081096","article-title":"Recent advances in the molecular effects of biostimulants in plants: An overview","volume":"11","author":"M Baltazar","year":"2021","journal-title":"Biomolecules"},{"issue":"10","key":"pone.0360081.ref013","doi-asserted-by":"crossref","first-page":"616","DOI":"10.3390\/agronomy9100616","article-title":"Biostimulants for plant growth promotion and sustainable management of phytoparasitic nematodes in vegetable crops","volume":"9","author":"T D\u2019Addabbo","year":"2019","journal-title":"Agronomy"},{"key":"pone.0360081.ref014","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1002\/9781119357254.ch5","article-title":"Auxin.","volume-title":"The Chemical Biology of Plant Biostimulants","author":"S Raggi","year":"2020"},{"issue":"1","key":"pone.0360081.ref015","doi-asserted-by":"crossref","first-page":"10710","DOI":"10.1038\/s41598-024-61576-4","article-title":"Integrated multi-omic approach reveals the effect of a Graminaceae-derived biostimulant and its lighter fraction on salt-stressed lettuce plants","volume":"14","author":"S Monterisi","year":"2024","journal-title":"Sci Rep"},{"issue":"3","key":"pone.0360081.ref016","doi-asserted-by":"crossref","first-page":"596","DOI":"10.1111\/jipb.13873","article-title":"Decoding small peptides: Regulators of plant growth and stress resilience","volume":"67","author":"F Xiao","year":"2025","journal-title":"J Integr Plant Biol"},{"issue":"3","key":"pone.0360081.ref017","doi-asserted-by":"crossref","first-page":"189","DOI":"10.3390\/horticulturae8030189","article-title":"Biostimulants on crops: Their impact under abiotic stress conditions","volume":"8","author":"G Franzoni","year":"2022","journal-title":"Horticulturae"},{"issue":"6","key":"pone.0360081.ref018","doi-asserted-by":"crossref","first-page":"403","DOI":"10.3390\/metabo11060403","article-title":"A metabolic choreography of maize plants treated with a humic substance-based biostimulant under normal and starved conditions","volume":"11","author":"K Othibeng","year":"2021","journal-title":"Metabolites"},{"key":"pone.0360081.ref019","doi-asserted-by":"crossref","first-page":"100373","DOI":"10.1016\/j.cpb.2024.100373","article-title":"Impact of a biostimulant enriched in betalain degradation products on ROS signaling, proline accumulation, and phytohormone homeostasis","volume":"39","author":"N Gatti","year":"2024","journal-title":"Curr Plant Biol"},{"issue":"1","key":"pone.0360081.ref020","doi-asserted-by":"crossref","first-page":"1777","DOI":"10.1186\/s12870-025-07810-1","article-title":"Genotype-specific transcriptomic response to drought stress in potato cultivars modulated by microbial biostimulants","volume":"25","author":"R Tamburino","year":"2025","journal-title":"BMC Plant Biol"},{"issue":"22","key":"pone.0360081.ref021","doi-asserted-by":"crossref","first-page":"13958","DOI":"10.3390\/ijms232213958","article-title":"Enhanced resistance to Sclerotinia sclerotiorum in Brassica rapa by activating host immunity through exogenous Verticillium dahliae Aspf2-like protein (VDAL) treatment","volume":"23","author":"S Jiang","year":"2022","journal-title":"Int J Mol Sci"},{"key":"pone.0360081.ref022","doi-asserted-by":"crossref","first-page":"100408","DOI":"10.1016\/j.plgene.2023.100408","article-title":"Molecular and ionomic responses of Solanum lycopersicum L. (cv. Micro-Tom) plants treated with a novel calcium-based plant biostimulant","volume":"34","author":"G Bertoldo","year":"2023","journal-title":"Plant Gene"},{"key":"pone.0360081.ref023","doi-asserted-by":"crossref","unstructured":"Trejo-T\u00e9llez LI, G\u00f3mez-Trejo LF, G\u00f3mez-Merino FC. Biostimulant Effects and Concentration Patterns of Beneficial Elements in Plants. In: Pandey S, Tripathi DK, Singh VP, Sharma S, Chauhan DK, editors. Beneficial Chemical Elements of Plants. 2023. p. 58\u2013102.","DOI":"10.1002\/9781119691419.ch4"},{"issue":"5","key":"pone.0360081.ref024","doi-asserted-by":"crossref","first-page":"1127","DOI":"10.1105\/tpc.18.00723","article-title":"The bZIP1 transcription factor regulates lipid remodeling and contributes to ER stress management in Chlamydomonas reinhardtii","volume":"31","author":"Y Yamaoka","year":"2019","journal-title":"Plant Cell"},{"key":"pone.0360081.ref025","doi-asserted-by":"crossref","first-page":"1132561","DOI":"10.3389\/fgene.2023.1132561","article-title":"Rice-Magnaporthe transcriptomics reveals host defense activation induced by red seaweed-biostimulant in rice plants","volume":"14","author":"SN Banakar","year":"2023","journal-title":"Front Genet"},{"issue":"2","key":"pone.0360081.ref026","doi-asserted-by":"crossref","DOI":"10.1111\/tpj.70382","article-title":"Genomics control of biostimulant-induced stress tolerance and crop yield enhancement","volume":"123","author":"T Gechev","year":"2025","journal-title":"Plant J"},{"key":"pone.0360081.ref027","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1007\/s11104-023-06156-7","article-title":"Plant physiological and molecular responses triggered by humic based biostimulants - A way forward to sustainable agriculture","volume":"492","author":"P Rathor","year":"2023","journal-title":"Plant Soil"},{"issue":"1","key":"pone.0360081.ref028","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1186\/s12864-020-06815-4","article-title":"Fulvic acid ameliorates drought stress-induced damage in tea plants by regulating the ascorbate metabolism and flavonoids biosynthesis","volume":"21","author":"J Sun","year":"2020","journal-title":"BMC Genomics"},{"key":"pone.0360081.ref029","doi-asserted-by":"crossref","first-page":"1248648","DOI":"10.3389\/fpls.2023.1248648","article-title":"The calcium connection: Exploring the intricacies of calcium signaling in plant-microbe interactions","volume":"14","author":"NP Negi","year":"2023","journal-title":"Front Plant Sci"},{"issue":"4","key":"pone.0360081.ref030","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1080\/07352680091139240","article-title":"Calcium Signaling: Linking Environmental signals to cellular functions","volume":"19","author":"S Pandey","year":"2000","journal-title":"Crit Rev Plant Sci"},{"issue":"9","key":"pone.0360081.ref031","doi-asserted-by":"crossref","first-page":"5043","DOI":"10.3390\/ijms25095043","article-title":"Molecular mechanisms of CBL-CIPK signaling pathway in plant abiotic stress tolerance and hormone crosstalk","volume":"25","author":"C Kaya","year":"2024","journal-title":"Int J Mol Sci"},{"issue":"6","key":"pone.0360081.ref032","doi-asserted-by":"crossref","first-page":"620","DOI":"10.1016\/j.pbi.2008.10.010","article-title":"Phosphoinositides in plants: Novel functions in membrane trafficking","volume":"11","author":"JM Thole","year":"2008","journal-title":"Curr Opin Plant Biol"},{"issue":"4","key":"pone.0360081.ref033","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1016\/j.tplants.2020.11.010","article-title":"Coordination of phospholipid-based signaling and membrane trafficking in plant immunity","volume":"26","author":"J Xing","year":"2021","journal-title":"Trends Plant Sci"},{"key":"pone.0360081.ref034","doi-asserted-by":"crossref","first-page":"1535943","DOI":"10.3389\/fpls.2025.1535943","article-title":"Dynamic molecular regulation of salt stress responses in maize (Zea mays L.) seedlings","volume":"16","author":"A Maimaiti","year":"2025","journal-title":"Front Plant Sci"},{"issue":"4","key":"pone.0360081.ref035","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1080\/07352689.2017.1402852","article-title":"Biosynthesis and regulation of phenylpropanoids in plants","volume":"36","author":"Y Deng","year":"2017","journal-title":"Crit Rev Plant Sci"},{"issue":"5","key":"pone.0360081.ref036","doi-asserted-by":"crossref","first-page":"454","DOI":"10.3390\/antiox9050454","article-title":"Role of Cytochrome P450 Enzymes in plant stress response","volume":"9","author":"BA Pandian","year":"2020","journal-title":"Antioxidants"},{"key":"pone.0360081.ref037","doi-asserted-by":"crossref","first-page":"582267","DOI":"10.3389\/fpls.2020.582267","article-title":"Humic acid enhances the growth of tomato promoted by endophytic bacterial strains through the activation of hormone-, growth-, and transcription-related processes","volume":"11","author":"N Galambos","year":"2020","journal-title":"Front Plant Sci"},{"issue":"10","key":"pone.0360081.ref038","doi-asserted-by":"crossref","first-page":"2537","DOI":"10.3390\/cells10102537","article-title":"Biostimulants for the regulation of reactive oxygen species metabolism in plants under abiotic stress","volume":"10","author":"M Hasanuzzaman","year":"2021","journal-title":"Cells"},{"issue":"3","key":"pone.0360081.ref039","doi-asserted-by":"crossref","first-page":"257","DOI":"10.3390\/horticulturae8030257","article-title":"Exogenous melatonin protects lime plants from drought stress-induced damage by maintaining cell membrane structure, detoxifying ROS and regulating antioxidant systems","volume":"8","author":"M Jafari","year":"2022","journal-title":"Horticulturae"}],"container-title":["PLOS One"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0360081","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T17:28:43Z","timestamp":1791480523000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0360081"}},"subtitle":[],"editor":[{"given":"Mohammad Sadegh","family":"Taghizadeh","sequence":"first","affiliation":[{"name":"Shiraz University","place":["IRAN, ISLAMIC REPUBLIC OF"]}],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,10,8]]},"references-count":39,"journal-issue":{"issue":"10","published-online":{"date-parts":[[2026,10,8]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pone.0360081","relation":{},"ISSN":["1932-6203"],"issn-type":[{"value":"1932-6203","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,8]]}}}