{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T03:02:02Z","timestamp":1789614122193,"version":"build-2803163510"},"reference-count":57,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T00:00:00Z","timestamp":1789603200000},"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. Genet."],"abstract":"<jats:sec>\n                    <jats:title>Introduction<\/jats:title>\n                    <jats:p>Drought stress adversely affects photosynthesis, redox homeostasis and metabolic signalling in plants. Exogenous application of hormonal and nonhormonal elicitors, have emerged as promising strategies to improve plant resilience to changing climatic conditions.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      In the current study, the effect of foliar application of gibberellic acid (GA\n                      <jats:sub>3<\/jats:sub>\n                      ) and 6\u2013benzylaminopurine (6\u2013BAP) individually and in combination was assessed at different doses (0.2 and 0.4 mM) in pigeonpea (\n                      <jats:italic>Cajanus cajan<\/jats:italic>\n                      ) under drought stress through integrated physiological, metabolomic and transcriptomic approaches.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>Growth, photosynthetic performance and metabolic homeostasis were severely affected by drought stress. The combined application of 0.4 mM GA3+6-BAP resulted in 20.6%, 29.7% and 33.2% increase in the maximum quantum efficiency of PSII (Fv\/Fm), \u0394pH, and \u0394\u03a8, respectively, compared to untreated drought-stressed plants, which shows that the stability of PSII and the energy regulation of the thylakoids improved with combined phytohormone application. Untargeted metabolite profiling revealed that drought reduced the abundance of metabolites belonging to carbohydrates, lipids and organoheterocyclic compounds respectively. While GA3+6-BAP led to the restoration of metabolites involved in carbon metabolism and antioxidative metabolism wherein highest accumulation of flavonoids, phenolics and terpenoids was noticed. Hormonal application induced 24 unique metabolic signatures including quercetin 3-(2-glucosylrhamnoside), camellianin A, 3-O-p-coumaroylquinic acid, byakangelicin. Besides, transcriptomic profiling demonstrated treatment-associated changes in the abundance of transcripts involved in photosynthesis, redox regulation and calcium-based signaling, suggesting coordinated stress signaling.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusion<\/jats:title>\n                    <jats:p>Overall, 0.4 mM GA3+ 6-BAP treatment proved to be the most effective among other treatments for the induction of drought stress tolerance in pigeonpea by dynamic transcriptional and metabolic regulatory mechanisms.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fgene.2026.1895564","type":"journal-article","created":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T02:11:11Z","timestamp":1789611071000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Dose-dependent gibberellic acid and benzylaminopurine priming coordinates transcriptional and metabolic reprogramming, enhancing photochemical efficiency to confer drought tolerance in pigeonpea"],"prefix":"10.3389","volume":"17","author":[{"given":"Rekha","family":"Rani","sequence":"first","affiliation":[{"name":"Biofuels and Climate Change Mitigation Laboratory, Department of Life Sciences, Chhatrapati Shahu Ji Maharaj (CSJM) University","place":["Kanpur, India"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shivam","family":"Kumar","sequence":"additional","affiliation":[{"name":"Biofuels and Climate Change Mitigation Laboratory, Department of Life Sciences, Chhatrapati Shahu Ji Maharaj (CSJM) University","place":["Kanpur, India"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Akhil","family":"Singh","sequence":"additional","affiliation":[{"name":"Biofuels and Climate Change Mitigation Laboratory, Department of Life Sciences, Chhatrapati Shahu Ji Maharaj (CSJM) University","place":["Kanpur, India"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rachapudi Venkata","family":"Sreeharsha","sequence":"additional","affiliation":[{"name":"Biofuels and Climate Change Mitigation Laboratory, Department of Life Sciences, Chhatrapati Shahu Ji Maharaj (CSJM) University","place":["Kanpur, India"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,9,17]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1126\/science.1118642","article-title":"Integration of plant responses to environmentally activated phytohormonal signals","volume":"311","author":"Achard","year":"2006","journal-title":"Science"},{"key":"B2","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/j.plantsci.2012.07.014","article-title":"Flavonoids as antioxidants in plants: location and functional significance","volume":"196","author":"Agati","year":"2012","journal-title":"Plant Science"},{"key":"B3","doi-asserted-by":"publisher","first-page":"1214845","DOI":"10.3389\/fmicb.2023.1214845","article-title":"Recent advances in PGPR-Mediated resilience toward interactive effects of drought and salt stress in plants","volume":"14","author":"Al-Turki","year":"2023","journal-title":"Front. Microbiology"},{"key":"B4","doi-asserted-by":"publisher","first-page":"1607","DOI":"10.1093\/jxb\/erh196","article-title":"Applications of chlorophyll fluorescence can improve crop production strategies: an examination of future possibilities","volume":"55","author":"Baker","year":"2004","journal-title":"J. Experimental Botany"},{"key":"B5","doi-asserted-by":"publisher","first-page":"16975","DOI":"10.1038\/s41598-020-73489-z","article-title":"Plant growth promoting rhizobacteria alleviates drought stress in potato in response to suppressive oxidative stress and antioxidant enzymes activities","volume":"10","author":"Batool","year":"2020","journal-title":"Sci. Rep."},{"key":"B6","doi-asserted-by":"publisher","first-page":"248","DOI":"10.1006\/abio.1976.9999","article-title":"A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding","volume":"72","author":"Bradford","year":"1976","journal-title":"Anal. Biochemistry"},{"key":"B7","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1242\/jeb.089938","article-title":"The role of gibberellin signalling in plant responses to abiotic stress","volume":"217","author":"Colebrook","year":"2014","journal-title":"J. Experimental Biology"},{"key":"B8","doi-asserted-by":"publisher","first-page":"4999","DOI":"10.1093\/jxb\/erv132","article-title":"Regulation of chloroplast development and function by cytokinin","volume":"66","author":"Cortleven","year":"2015","journal-title":"J. Experimental Botany"},{"key":"B9","doi-asserted-by":"publisher","first-page":"2650","DOI":"10.3390\/plants11192650","article-title":"Plant development and crop yield: the role of gibberellins","volume":"11","author":"Castro-Camba","year":"2022","journal-title":"Plants"},{"key":"B10","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s00299-021-02759-5","article-title":"Climate change regulated abiotic stress mechanisms in plants: a comprehensive review","volume":"41","author":"Chaudhry","year":"2022","journal-title":"Plant Cell Rep."},{"key":"B11","doi-asserted-by":"publisher","first-page":"i884","DOI":"10.1093\/bioinformatics\/bty560","article-title":"fastp: an ultra-fast all-in-one FASTQ preprocessor","volume":"34","author":"Chen","year":"2018","journal-title":"Bioinformatics"},{"key":"B12","first-page":"165","article-title":"Breeding for abiotic stresses in pigeonpea","volume":"24","author":"Choudhary","year":"2011","journal-title":"J. Food Legumes"},{"key":"B13","doi-asserted-by":"publisher","first-page":"R365","DOI":"10.1016\/j.cub.2011.03.013","article-title":"Hormone signalling crosstalk in plant growth regulation","volume":"21","author":"Depuydt","year":"2011","journal-title":"Curr. Biology"},{"key":"B14","doi-asserted-by":"publisher","first-page":"46","DOI":"10.1016\/j.plaphy.2022.02.004","article-title":"Change in the photochemical and structural organization of thylakoids from pea (Pisum sativum) under salt stress","volume":"177","author":"Dhokne","year":"2022","journal-title":"Plant Physiology Biochem."},{"key":"B15","doi-asserted-by":"publisher","first-page":"1077","DOI":"10.1093\/jxb\/ery437","article-title":"The metabolic response to drought","volume":"70","author":"F\u00e0bregas","year":"2019","journal-title":"J. Experimental Botany"},{"key":"B16","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1055\/s-2004-820867","article-title":"Diffusive and metabolic limitations to photosynthesis under drought and salinity in C3 plants","volume":"6","author":"Flexas","year":"2004","journal-title":"Plant Biology"},{"key":"B17","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1186\/s43897-021-00006-9","article-title":"Leaf senescence: progression, regulation, and application","volume":"1","author":"Guo","year":"2021","journal-title":"Mol. Hortic."},{"key":"B18","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1042\/BJ20120245","article-title":"Gibberellin biosynthesis and its regulation","volume":"444","author":"Hedden","year":"2012","journal-title":"Biochem. J."},{"key":"B19","first-page":"420","volume-title":"Methods in Carbohydrate Chemistry","author":"Hedge","year":"1962"},{"key":"B20","doi-asserted-by":"crossref","first-page":"1332","DOI":"10.1139\/b79-163","article-title":"A method for the extraction of chlorophyll from leaf tissue without maceration","volume":"57","author":"Hiscox","year":"1979","journal-title":"Can. Journal Botany"},{"key":"B21","doi-asserted-by":"publisher","first-page":"D309","DOI":"10.1093\/nar\/gky1085","article-title":"eggNOG 5.0: a hierarchical, functionally and phylogenetically annotated orthology resource based on 5090 organisms and 2502 viruses","volume":"47","author":"Huerta-Cepas","year":"2019","journal-title":"Nucleic Acids Research"},{"key":"B22","doi-asserted-by":"publisher","first-page":"799318","DOI":"10.3389\/fpls.2021.799318","article-title":"Phytohormones trigger drought tolerance in crop plants: outlook and future perspectives","volume":"12","author":"Iqbal","year":"2022","journal-title":"Front. Plant Sci."},{"key":"B23","doi-asserted-by":"publisher","first-page":"102","DOI":"10.1007\/s11738-016-2113-y","article-title":"Chlorophyll a fluorescence as a tool to monitor physiological status of plants under abiotic stress conditions","volume":"38","author":"Kalaji","year":"2016","journal-title":"Acta Physiol. Plant."},{"key":"B24","doi-asserted-by":"publisher","first-page":"719","DOI":"10.3389\/fpls.2017.00719","article-title":"Chloroplast ATP synthase modulation of the thylakoid proton motive force: implications for photosystem I and photosystem II photoprotection","volume":"8","author":"Kanazawa","year":"2017","journal-title":"Front. Plant Sci."},{"key":"B25","doi-asserted-by":"publisher","first-page":"dev149344","DOI":"10.1242\/dev.149344","article-title":"Cytokinin signaling in plant development","volume":"145","author":"Kieber","year":"2018","journal-title":"Development"},{"key":"B26","doi-asserted-by":"publisher","first-page":"907","DOI":"10.1038\/s41587-019-0201-4","article-title":"Graph-based genome alignment and genotyping with HISAT2 and HISAT-genotype","volume":"37","author":"Kim","year":"2019","journal-title":"Nat. Biotechnology"},{"key":"B27","doi-asserted-by":"publisher","first-page":"1250","DOI":"10.1105\/tpc.16.00027","article-title":"Vascular cell induction culture system using Arabidopsis leaves (VISUAL) reveals the sequential differentiation of sieve element-like cells","volume":"28","author":"Kondo","year":"2016","journal-title":"Plant Cell"},{"key":"B28","doi-asserted-by":"publisher","first-page":"1658","DOI":"10.1093\/bioinformatics\/btl158","article-title":"Cd-hit: a fast program for clustering and comparing large sets of protein or nucleotide sequences","volume":"22","author":"Li","year":"2006","journal-title":"Bioinformatics"},{"key":"B29","doi-asserted-by":"publisher","first-page":"402","DOI":"10.1006\/meth.2001.1262","article-title":"Analysis of relative gene expression data using real-time quantitative PCR and the 2\u2212 \u0394\u0394CT method","volume":"25","author":"Livak","year":"2001","journal-title":"Methods"},{"key":"B30","doi-asserted-by":"publisher","first-page":"659","DOI":"10.1093\/jxb\/51.345.659","article-title":"Chlorophyll fluorescence\u2014a practical guide","volume":"51","author":"Maxwell","year":"2000","journal-title":"J. Experimental Botany"},{"key":"B31","doi-asserted-by":"publisher","first-page":"417","DOI":"10.1038\/nmeth.4197","article-title":"Salmon provides fast and bias-aware quantification of transcript expression","volume":"14","author":"Patro","year":"2017","journal-title":"Nat. Methods"},{"key":"B32","doi-asserted-by":"publisher","first-page":"290","DOI":"10.1016\/j.pbi.2011.02.001","article-title":"Hormone balance and abiotic stress tolerance in crop plants","volume":"14","author":"Peleg","year":"2011","journal-title":"Curr. Opinion Plant Biology"},{"key":"B33","doi-asserted-by":"publisher","first-page":"869","DOI":"10.1093\/jxb\/erq340","article-title":"Photosynthesis and drought: can we make metabolic connections from available data?","volume":"62","author":"Pinheiro","year":"2011","journal-title":"J. Experimental Botany"},{"key":"B34","doi-asserted-by":"publisher","first-page":"1808","DOI":"10.3390\/plants13131808","article-title":"Impacts of drought on photosynthesis in major food crops and the related mechanisms of plant responses to drought","volume":"13","author":"Qiao","year":"2024","journal-title":"Plants"},{"key":"B35","doi-asserted-by":"publisher","first-page":"90","DOI":"10.1007\/s13205-019-1626-7","article-title":"Application of silicon nanoparticles in agriculture","volume":"9","author":"Rastogi","year":"2019","journal-title":"3 Biotech."},{"key":"B36","doi-asserted-by":"publisher","first-page":"19631","DOI":"10.1073\/pnas.0709453104","article-title":"Delayed leaf senescence induces extreme drought tolerance in a flowering plant","volume":"104","author":"Rivero","year":"2007","journal-title":"Proc. Natl. Acad. Sci."},{"key":"B58","doi-asserted-by":"publisher","first-page":"172","DOI":"10.1016\/j.jphotobiol.2015.08.008","article-title":"Differential responses in photosynthesis, growth and biomass yields in two mulberry genotypes grown under elevated CO2 atmosphere","volume":"151","author":"Sekhar","year":"2015","journal-title":"J. Photochem. Photobiol. B: Biol."},{"key":"B37","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1007\/BF00024185","article-title":"Continuous recording of photochemical and non-photochemical chlorophyll fluorescence quenching with a new type of modulation fluorometer","volume":"10","author":"Schreiber","year":"1986","journal-title":"Photosynth. Research"},{"key":"B38","doi-asserted-by":"publisher","first-page":"1237295","DOI":"10.3389\/fpls.2023.1237295","article-title":"Phytohormonal modulation of the drought stress in soybean: outlook, research progress, and cross-talk","volume":"14","author":"Shaffique","year":"2023","journal-title":"Front. Plant Science"},{"key":"B39","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1093\/plphys\/kiae044","article-title":"Highlights in gibberellin research: a tale of the dwarf and the slender","volume":"195","author":"Shani","year":"2024","journal-title":"Plant Physiol."},{"key":"B40","doi-asserted-by":"publisher","first-page":"6760","DOI":"10.1007\/s00344-023-11010-2","article-title":"Unraveling the plant growth-promoting mechanisms of Stenotrophomonas sp. CV83 for drought stress tolerance and growth enhancement in chickpea","volume":"42","author":"Sharma","year":"2023","journal-title":"J. Plant Growth Regul."},{"key":"B41","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1016\/j.proenv.2015.07.231","article-title":"Dynamics of vegetative and reproductive growth patterns in Pigeonpea (Cajanus cajan L.) grown under elevated CO2","volume":"29","author":"Sreeharsha","year":"2015","journal-title":"Procedia Environ. Sci."},{"key":"B42","doi-asserted-by":"publisher","first-page":"82","DOI":"10.1016\/j.plantsci.2014.11.012","article-title":"Delayed flowering is associated with lack of photosynthetic acclimation in Pigeon pea (Cajanus cajan L.) grown under elevated CO2","volume":"231","author":"Sreeharsha","year":"2015","journal-title":"Plant Sci."},{"key":"B43","doi-asserted-by":"publisher","first-page":"425","DOI":"10.1007\/s11120-018-0586-9","article-title":"Mitigation of drought-induced oxidative damage by enhanced carbon assimilation and an efficient antioxidative metabolism under high CO2 environment in pigeonpea (Cajanus cajan L.)","volume":"139","author":"Sreeharsha","year":"2019","journal-title":"Photosynth. Research"},{"key":"B57","doi-asserted-by":"publisher","first-page":"28129","DOI":"10.1007\/s13399-023-05011-0","article-title":"Catalytic hydrothermal liquefaction of Pongamia pinnata (L.) to produce bio-oil and biochar within a biorefinery framework","volume":"15","author":"Sreeharsha","year":"2025","journal-title":"Biomass Conv. Bioref."},{"key":"B44","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1038\/s41586-018-0009-2","article-title":"A small peptide modulates stomatal control via abscisic acid in long-distance signalling","volume":"556","author":"Takahashi","year":"2018","journal-title":"Nature"},{"key":"B45","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1016\/j.trac.2008.01.008","article-title":"LC-MS-based methodology for global metabolite profiling in metabonomics\/metabolomics","volume":"27","author":"Theodoridis","year":"2008","journal-title":"TrAC Trends Anal. Chem."},{"key":"B46","doi-asserted-by":"publisher","first-page":"1008","DOI":"10.1104\/pp.104.039578","article-title":"DELLA proteins and gibberellin-regulated seed germination and floral development in Arabidopsis","volume":"135","author":"Tyler","year":"2004","journal-title":"Plant Physiol."},{"key":"B47","doi-asserted-by":"publisher","first-page":"80","DOI":"10.1007\/s11738-021-03245-x","article-title":"Growth, seed yield and nutritional characteristics of pigeonpea grown under elevated CO2 atmosphere","volume":"43","author":"Unnikrishnan","year":"2021","journal-title":"Acta Physiol. Plant."},{"key":"B48","doi-asserted-by":"publisher","first-page":"483","DOI":"10.1007\/s12298-024-01434-9","article-title":"Flowering onset time is regulated by microRNA-mediated trehalose-6-phosphate signaling in Cajanus cajan L. under elevated CO2","volume":"30","author":"Unnikrishnan","year":"2024","journal-title":"Physiol. Mol. Biol. Plants"},{"key":"B49","doi-asserted-by":"publisher","first-page":"1620","DOI":"10.3390\/plants11131620","article-title":"Plants\u2019 physio-biochemical and phyto-hormonal responses to alleviate the adverse effects of drought stress: a comprehensive review","volume":"11","author":"Wahab","year":"2022","journal-title":"Plants"},{"key":"B50","doi-asserted-by":"publisher","first-page":"107723","DOI":"10.1016\/j.plaphy.2023.107723","article-title":"Integrative transcriptome and metabolome analysis reveals the mechanism of exogenous melatonin alleviating drought stress in maize roots","volume":"199","author":"Wang","year":"2023","journal-title":"Plant Physiology Biochem."},{"key":"B51","doi-asserted-by":"publisher","first-page":"100678","DOI":"10.1016\/j.xplc.2023.100678","article-title":"Calcium signalling in plant mineral nutrition: from uptake to transport","volume":"4","author":"Wang","year":"2023","journal-title":"Plant Commun."},{"key":"B52","doi-asserted-by":"publisher","first-page":"162","DOI":"10.1016\/j.cj.2016.01.010","article-title":"Phytohormones and their metabolic engineering for abiotic stress tolerance in crop plants","volume":"4","author":"Wani","year":"2016","journal-title":"Crop Journal"},{"key":"B54","doi-asserted-by":"publisher","first-page":"421","DOI":"10.3390\/cimb47060421","article-title":"Progress in transcriptomics and metabolomics in plant responses to abiotic stresses","volume":"47","author":"Yu","year":"2025","journal-title":"Curr. Issues Mol. Biol."},{"key":"B55","doi-asserted-by":"publisher","first-page":"1161","DOI":"10.1111\/nph.18087","article-title":"Plant responses to multifactorial stress combination","volume":"234","author":"Zandalinas","year":"2022","journal-title":"New Phytol."},{"key":"B56","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1007\/s13593-025-01030-5","article-title":"Yield gaps in soybean: causes and pathways for increasing yield in smallholder farming from Northeast China","volume":"45","author":"Zhao","year":"2025","journal-title":"Agron. Sustain. Dev."}],"container-title":["Frontiers in Genetics"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fgene.2026.1895564\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T02:11:13Z","timestamp":1789611073000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fgene.2026.1895564\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,17]]},"references-count":57,"alternative-id":["10.3389\/fgene.2026.1895564"],"URL":"https:\/\/doi.org\/10.3389\/fgene.2026.1895564","relation":{},"ISSN":["1664-8021"],"issn-type":[{"value":"1664-8021","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,17]]},"article-number":"1895564"}}