{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T19:06:50Z","timestamp":1789499210397,"version":"build-2803163510"},"reference-count":61,"publisher":"MDPI AG","issue":"18","license":[{"start":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T00:00:00Z","timestamp":1789430400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100011785","name":"Guangxi Natural Science Foundation","doi-asserted-by":"publisher","award":["2023GXNSFBA026122"],"award-info":[{"award-number":["2023GXNSFBA026122"]}],"id":[{"id":"10.13039\/501100011785","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100011785","name":"Guangxi Natural Science Foundation","doi-asserted-by":"publisher","award":["2025GXNSFAA069090"],"award-info":[{"award-number":["2025GXNSFAA069090"]}],"id":[{"id":"10.13039\/501100011785","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32360067"],"award-info":[{"award-number":["32360067"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"award":["32360067"],"award-info":[{"award-number":["32360067"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]},{"name":"Guangxi Young Talent Project"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Phosphate (Pi) starvation severely limits plant growth, and the transcription factor PHR1 is a central regulator of the phosphate starvation response (PSR). However, its role in modulating the secreted proteome remains poorly understood. Here, we performed comparative secretome profiling of wild-type (WT) and phr1 mutant Arabidopsis seedlings under Pi-sufficient (+P) and Pi-deficient (\u2212P) conditions using LC\u2013MS\/MS. We quantified 2077 proteins from culture media across four biological replicates. Pi starvation induced 394 differentially abundant secreted proteins in WT, while the phr1 mutation severely attenuated this response, particularly down-regulating 213 proteins under \u2013P conditions. PHR1 was essential for the starvation-induced secretion of phosphate-acquisition enzymes, including GDPD6, PAP1, and RNS1, and its role in the promotion of extracellular phosphatases was confirmed by acid phosphatase activity assays. Additionally, PHR1 modulated redox-related and defense-related secreted proteins, linking nutrient stress with oxidative and immune signaling. Integrative transcriptomic analysis revealed partial concordance between protein and mRNA levels, suggesting both transcriptional and post-transcriptional regulation. Collectively, our findings establish PHR1 as a vital regulator of the Pi-starvation-induced secretome and provide new insights into plant adaptation to phosphorus limitation through coordinated protein secretion.<\/jats:p>","DOI":"10.3390\/plants15182826","type":"journal-article","created":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T18:23:53Z","timestamp":1789496633000},"page":"2826","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Comparative Secretome Profiling of Arabidopsis Seedlings Under Pi Starvation Unveils the Regulatory Role of PHR1"],"prefix":"10.3390","volume":"15","author":[{"given":"Ran","family":"Xu","sequence":"first","affiliation":[{"name":"Guangxi Sugarcane Bio-Breeding Laboratory, State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources, College of Life Science and Technology, Guangxi University, Nanning 530004, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"}]},{"given":"Jizhen","family":"Xie","sequence":"additional","affiliation":[{"name":"Guangxi Sugarcane Bio-Breeding Laboratory, State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources, College of Life Science and Technology, Guangxi University, Nanning 530004, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"}]},{"given":"Xueqing","family":"Duan","sequence":"additional","affiliation":[{"name":"Guangxi Sugarcane Bio-Breeding Laboratory, State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources, College of Life Science and Technology, Guangxi University, Nanning 530004, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"validation","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"}]},{"given":"Piao","family":"Liu","sequence":"additional","affiliation":[{"name":"Guangxi Sugarcane Bio-Breeding Laboratory, State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources, College of Life Science and Technology, Guangxi University, Nanning 530004, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"validation","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"}]},{"given":"Xingyi","family":"Zheng","sequence":"additional","affiliation":[{"name":"Guangxi Sugarcane Bio-Breeding Laboratory, State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources, College of Life Science and Technology, Guangxi University, Nanning 530004, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-6829-228X","authenticated-orcid":false,"given":"Xiaoyang","family":"Li","sequence":"additional","affiliation":[{"name":"Guangxi Sugarcane Bio-Breeding Laboratory, State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources, College of Life Science and Technology, Guangxi University, Nanning 530004, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"validation","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"}]},{"given":"Sheng","family":"Jiang","sequence":"additional","affiliation":[{"name":"Guangxi Sugarcane Bio-Breeding Laboratory, State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources, College of Life Science and Technology, Guangxi University, Nanning 530004, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"data-curation","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"}]},{"given":"Wenjun","family":"Wang","sequence":"additional","affiliation":[{"name":"Cloud-Clone Co., Ltd., Wuhan 430056, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"formal-analysis","vocabulary":"credit"}]},{"given":"Xixi","family":"Cai","sequence":"additional","affiliation":[{"name":"Guangxi Sugarcane Bio-Breeding Laboratory, State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources, College of Life Science and Technology, Guangxi University, Nanning 530004, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"resources","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"}]},{"given":"Shengchao","family":"Ge","sequence":"additional","affiliation":[{"name":"Guangxi Sugarcane Bio-Breeding Laboratory, State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources, College of Life Science and Technology, Guangxi University, Nanning 530004, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"resources","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4166-9056","authenticated-orcid":false,"given":"Yunfeng","family":"Liu","sequence":"additional","affiliation":[{"name":"Guangxi Sugarcane Bio-Breeding Laboratory, State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources, College of Life Science and Technology, Guangxi University, Nanning 530004, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4406-3946","authenticated-orcid":false,"given":"Yuan","family":"Su","sequence":"additional","affiliation":[{"name":"Guangxi Sugarcane Bio-Breeding Laboratory, State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources, College of Life Science and Technology, Guangxi University, Nanning 530004, China"},{"name":"Guangxi Technology Innovation Center for Microbial Resources Development and Utilization, Nanning 530004, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,15]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1504","DOI":"10.1093\/plcell\/koad326","article-title":"Milestones in understanding transport, sensing, and signaling of the plant nutrient phosphorus","volume":"36","author":"Yang","year":"2024","journal-title":"Plant Cell"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"6417","DOI":"10.1093\/jxb\/erad326","article-title":"Phosphate starvation: Response mechanisms and solutions","volume":"74","author":"Madison","year":"2023","journal-title":"J. Exp. Bot."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"100064","DOI":"10.1016\/j.ncrops.2024.100064","article-title":"Molecular mechanisms underlying plant responses to low phosphate stress and potential applications in crop improvement","volume":"2","author":"Hu","year":"2025","journal-title":"New Crops"},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Sega, P., and Pacak, A. (2019). Plant PHR transcription factors: Put on a map. Genes, 10.","DOI":"10.3390\/genes10121018"},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Bustos, R., Castrillo, G., Linhares, F., Puga, M.I., Rubio, V., P\u00e9rez-P\u00e9rez, J., Solano, R., Leyva, A., and Paz-Ares, J. (2010). A central regulatory system largely controls transcriptional activation and repression responses to phosphate starvation in Arabidopsis. PLoS Genet., 6.","DOI":"10.1371\/journal.pgen.1001102"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"14947","DOI":"10.1073\/pnas.1404654111","article-title":"SPX1 is a phosphate-dependent inhibitor of PHOSPHATE STARVATION RESPONSE 1 in Arabidopsis","volume":"111","author":"Puga","year":"2014","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"2156","DOI":"10.1093\/jxb\/erag002","article-title":"Phosphate starvation response 1 (PHR1): A versatile master regulator shaping plant resilience beyond phosphate deprivation","volume":"77","author":"Mangalakkadan","year":"2026","journal-title":"J. Exp. Bot."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"2122","DOI":"10.1101\/gad.204401","article-title":"A conserved MYB transcription factor involved in phosphate starvation signaling both in vascular plants and in unicellular algae","volume":"15","author":"Rubio","year":"2001","journal-title":"Genes Dev."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"4317","DOI":"10.1002\/pmic.200800292","article-title":"Proteomic analysis of alterations in the secretome of Arabidopsis thaliana suspension cells subjected to nutritional phosphate deficiency","volume":"8","author":"Tran","year":"2008","journal-title":"Proteomics"},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"He, Y., Hu, J., Jiang, L., Wang, S., Li, L., Zhang, X., Zhao, G., He, S., Yang, Y., and Hong, G. (2026). Phosphate starvation primes plant immunity through an ABA-OsbZIP12-phytoalexin signaling network. Plant Commun., 102038.","DOI":"10.1016\/j.xplc.2026.102038"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1038\/nature21417","article-title":"Root microbiota drive direct integration of phosphate stress and immunity","volume":"543","author":"Castrillo","year":"2017","journal-title":"Nature"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"EMBJ2021109102","DOI":"10.15252\/embj.2021109102","article-title":"Plant immunity suppression via PHR1-RALF-FERONIA shapes the root microbiome to alleviate phosphate starvation","volume":"41","author":"Tang","year":"2022","journal-title":"EMBO J."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"100011","DOI":"10.1016\/j.ncrops.2024.100011","article-title":"Uncovering the function of peptides: Bridging hormone signaling, microbial interactions, and root development in plants","volume":"1","author":"Zhang","year":"2024","journal-title":"New Crops"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1023","DOI":"10.1038\/s41587-021-01156-3","article-title":"SignalP 6.0 predicts all five types of signal peptides using protein language models","volume":"40","author":"Teufel","year":"2022","journal-title":"Nat. Biotechnol."},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Alexandersson, E., Ashfaq, A., Resj\u00f6, S., and Andreasson, E. (2013). Plant secretome proteomics. Front. Plant Sci., 4.","DOI":"10.3389\/fpls.2013.00009"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"kiag585","DOI":"10.1093\/plphys\/kiag585","article-title":"Phospholipid homeostasis and gametophyte development require the glycerophosphodiester phosphodiesterases GDPD5 and GDPD6","volume":"202","author":"Siebers","year":"2026","journal-title":"Plant Physiol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"382","DOI":"10.1093\/jxb\/erab409","article-title":"Arabidopsis PAP17 is a dual-localized purple acid phosphatase up-regulated during phosphate deprivation, senescence, and oxidative stress","volume":"73","author":"Ghahremani","year":"2022","journal-title":"J. Exp. Bot."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1232","DOI":"10.1104\/pp.010835","article-title":"Phosphite, an analog of phosphate, suppresses the coordinated expression of genes under phosphate starvation","volume":"129","author":"Varadarajan","year":"2002","journal-title":"Plant Physiol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"673","DOI":"10.1046\/j.1365-313X.1994.6050673.x","article-title":"The Arabidopsis ribonuclease gene RNS1 is tightly controlled in response to phosphate limitation","volume":"6","author":"Bariola","year":"1994","journal-title":"Plant J."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"485","DOI":"10.1111\/mpp.12769","article-title":"Contribution of cell wall peroxidase- and NADPH oxidase-derived reactive oxygen species to Alternaria brassicicola-induced oxidative burst in Arabidopsis","volume":"20","author":"Gullner","year":"2019","journal-title":"Mol. Plant Pathol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1063","DOI":"10.1105\/tpc.19.00525","article-title":"CRK2 and C-terminal phosphorylation of NADPH oxidase RBOHD regulate reactive oxygen species production in Arabidopsis","volume":"32","author":"Kimura","year":"2020","journal-title":"Plant Cell"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1094\/MPMI-8-0435","article-title":"Expression of defense-related and putative signaling genes during tolerant and susceptible interactions of Arabidopsis with Xanthomonas campestris pv. campestris","volume":"8","author":"Buell","year":"1995","journal-title":"Mol. Plant-Microbe Interact."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"3721","DOI":"10.1105\/tpc.106.044180","article-title":"Phytotoxicity and innate immune responses induced by Nep1-like proteins","volume":"18","author":"Qutob","year":"2006","journal-title":"Plant Cell"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"999","DOI":"10.1104\/pp.103.021683","article-title":"Systemic gene expression in Arabidopsis during an incompatible interaction with Alternaria brassicicola","volume":"132","author":"Schenk","year":"2003","journal-title":"Plant Physiol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"709","DOI":"10.1016\/j.jprot.2009.10.005","article-title":"Proteomics of the response of Arabidopsis thaliana to infection with Alternaria brassicicola","volume":"73","author":"Mukherjee","year":"2010","journal-title":"J. Proteom."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Castillo, N., Pastor, V., Ch\u00e1vez, \u00c1., Arr\u00f3, M., Boronat, A., Flors, V., Ferrer, A., and Altabella, T. (2019). Inactivation of UDP-glucose sterol glucosyltransferases enhances Arabidopsis resistance to Botrytis cinerea. Front. Plant Sci., 10.","DOI":"10.3389\/fpls.2019.01162"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"608","DOI":"10.1094\/MPMI.2002.15.6.608","article-title":"NHL25 and NHL3, two NDR1\/HIN1-like genes in Arabidopsis thaliana with potential role(s) in plant defense","volume":"15","author":"Varet","year":"2002","journal-title":"Mol. Plant-Microbe Interact."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Launay, A., Jolivet, S., Cl\u00e9ment, G., Zarattini, M., Dellero, Y., Le Hir, R., Jossier, M., Hodges, M., Expert, D., and Fagard, M. (2022). DspA\/E-triggered non-host resistance against E. amylovora depends on the Arabidopsis GLYCOLATE OXIDASE 2 gene. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23084224"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"436","DOI":"10.1104\/pp.108.121038","article-title":"Global analysis of Arabidopsis gene Eexpression uncovers a complex array of changes impacting pathogen response and cell cycle during geminivirus infection","volume":"148","author":"Sozzani","year":"2008","journal-title":"Plant Physiol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1093\/pcp\/pcx180","article-title":"A phi-class glutathione S-transferase gene for Verticillium wilt resistance in Gossypium arboreum identified in a genome-wide association study","volume":"59","author":"Gong","year":"2018","journal-title":"Plant Cell Physiol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"2018218","DOI":"10.1080\/15592324.2021.2018218","article-title":"Preinvasive nonhost resistance of Arabidopsis against melanized appressorium-mediated entry of multiple nonadapted Colletotrichum fungi","volume":"17","author":"Irieda","year":"2022","journal-title":"Plant Signal. Behav."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"540","DOI":"10.1111\/tpj.14237","article-title":"An Arabidopsis berberine bridge enzyme-like protein specifically oxidizes cellulose oligomers and plays a role in immunity","volume":"98","author":"Locci","year":"2019","journal-title":"Plant J."},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Ali, M.A., Abbas, A., Azeem, F., Shahzadi, M., and Bohlmann, H. (2020). The Arabidopsis GPI-anchored LTPg5 encoded by At3g22600 has a role in resistance against a diverse range of pathogens. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21051774"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"6955","DOI":"10.1093\/jxb\/erac342","article-title":"Characterization of the PHOSPHATE RESPONSE 2-dependent and -independent Pi-starvation response secretome in rice","volume":"73","author":"Du","year":"2022","journal-title":"J. Exp. Bot."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"2968","DOI":"10.1021\/pr901168v","article-title":"The mucilage proteome of Maize (Zea mays L.) primary roots","volume":"9","author":"Ma","year":"2010","journal-title":"J. Proteome Res."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"4748","DOI":"10.1021\/pr400614n","article-title":"Effect of phosphorus levels on the protein profiles of secreted protein and root surface protein of rice","volume":"12","author":"Shinano","year":"2013","journal-title":"J. Proteome Res."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1007\/s11104-006-9048-9","article-title":"Extracellular proteomes of Arabidopsis thaliana and Brassica napus roots: Analysis and comparison by MudPIT and LC-MS\/MS","volume":"286","author":"Basu","year":"2006","journal-title":"Plant Soil"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"773","DOI":"10.1104\/pp.106.091637","article-title":"Extracellular proteins in Pea root tip and border cell exudates","volume":"143","author":"Wen","year":"2007","journal-title":"Plant Physiol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"728","DOI":"10.1104\/pp.16.01253","article-title":"Extracellular vesicles isolated from the leaf apoplast carry stress-response proteins","volume":"173","author":"Rutter","year":"2017","journal-title":"Plant Physiol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"691","DOI":"10.1016\/j.molp.2026.01.013","article-title":"Extracellular vesicles in plants","volume":"19","author":"Liu","year":"2026","journal-title":"Mol. Plant."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1093\/jxb\/eraa482","article-title":"Recent insights into the metabolic adaptations of phosphorus-deprived plants","volume":"72","author":"Dissanayaka","year":"2021","journal-title":"J. Exp. Bot."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1007\/s00299-021-02773-7","article-title":"Purple acid phosphatases: Roles in phosphate utilization and new emerging functions","volume":"41","author":"Bhadouria","year":"2022","journal-title":"Plant Cell Rep."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"105241","DOI":"10.1016\/j.envexpbot.2023.105241","article-title":"WRKY transcription factors regulate phosphate uptake in plants","volume":"208","author":"Tang","year":"2023","journal-title":"Environ. Exp. Bot."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"499","DOI":"10.1104\/pp.15.01336","article-title":"Arabidopsis PHL2 and PHR1 act redundantly as the key components of the central regulatory system controlling transcriptional responses to phosphate starvation","volume":"170","author":"Sun","year":"2016","journal-title":"Plant Physiol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1299","DOI":"10.1078\/0176-1617-00541","article-title":"Oxidative stress during phosphate deficiency in roots of bean plants (Phaseolus vulgaris L.)","volume":"158","author":"Juszczuk","year":"2001","journal-title":"J. Plant Physiol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1111\/j.1744-7909.2007.00358.x","article-title":"Oxidative stress and antioxidant responses in young leaves of Mulberry plants grown under nitrogen, phosphorus or potassium deficiency","volume":"49","author":"Tewari","year":"2007","journal-title":"J. Integr. Plant Biol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"e70264","DOI":"10.1111\/ppl.70264","article-title":"Hydrogen peroxide positively regulates phosphate starvation responses in rice","volume":"177","author":"Yang","year":"2025","journal-title":"Physiol. Plant."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1038\/s41580-022-00499-2","article-title":"Reactive oxygen species signalling in plant stress responses","volume":"23","author":"Mittler","year":"2022","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1146\/annurev-arplant-042817-040322","article-title":"Reactive oxygen species in plant signaling","volume":"69","author":"Waszczak","year":"2018","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Guo, Z., Yan, X., and Liang, J. (2026). Proteome\u2013transcriptome discordance in rice under drought is modulated by post-translational modifications with functional consequences for photosynthesis and energy metabolism. Plants, 15.","DOI":"10.3390\/plants15101559"},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Liu, S., Shi, H., Yan, L., Jiang, C., Zhao, H., Lu, H., Li, H., Wang, S., and Huang, Z. (2025). Identification of candidate genes and proteins for tasseling stage drought tolerance through integrated transcriptomic and proteomic analysis approach in maize. BMC Plant Biol., 25.","DOI":"10.1186\/s12870-025-07264-5"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"154863","DOI":"10.1016\/j.jplph.2026.154863","article-title":"An integrated transcriptomic and proteomics analyses reveal the impact of drought stress on Pisum sativum","volume":"325","author":"Pandey","year":"2026","journal-title":"J. Plant Physiol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"799","DOI":"10.1002\/pmic.200900514","article-title":"Plant secretome: Unlocking secrets of the secreted proteins","volume":"10","author":"Agrawal","year":"2010","journal-title":"Proteomics"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"2429","DOI":"10.1016\/j.bbapap.2013.03.024","article-title":"Plant secretome\u2014From cellular process to biological activity","volume":"1834","author":"Krause","year":"2013","journal-title":"Biochim. Biophys. Acta Proteins Proteom."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"775","DOI":"10.1111\/j.1365-313X.2010.04375.x","article-title":"Dissection of local and systemic transcriptional responses to phosphate starvation in Arabidopsis","volume":"64","author":"Thibaud","year":"2010","journal-title":"Plant J."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"470","DOI":"10.1038\/s41564-018-0129-3","article-title":"Dynamic root exudate chemistry and microbial substrate preferences drive patterns in rhizosphere microbial community assembly","volume":"3","author":"Zhalnina","year":"2018","journal-title":"Nat. Microbiol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/j.tplants.2017.09.003","article-title":"Feed your friends: Do plant exudates shape the root microbiome?","volume":"23","author":"Sasse","year":"2018","journal-title":"Trends Plant Sci."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"fuaf008","DOI":"10.1093\/femsre\/fuaf008","article-title":"Plant exudates-driven microbiome recruitment and assembly facilitates plant health management","volume":"49","author":"Yang","year":"2025","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"4077","DOI":"10.1111\/nph.71174","article-title":"Evaluation of combined root exudate and rhizosphere microbiota sampling approaches to elucidate plant\u2013soil\u2013microbe interactions","volume":"250","author":"Browne","year":"2026","journal-title":"New Phytol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"4321","DOI":"10.1093\/jxb\/eraa179","article-title":"Purple acid phosphatase 10c encodes a major acid phosphatase that regulates plant growth under phosphate-deficient conditions in rice","volume":"71","author":"Deng","year":"2020","journal-title":"J. Exp. Bot."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"D543","DOI":"10.1093\/nar\/gkae1011","article-title":"The PRIDE database at 20 years: 2025 update","volume":"53","author":"Bandla","year":"2025","journal-title":"Nucleic Acids Res."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/18\/2826\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T18:43:49Z","timestamp":1789497829000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/18\/2826"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,15]]},"references-count":61,"journal-issue":{"issue":"18","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["plants15182826"],"URL":"https:\/\/doi.org\/10.3390\/plants15182826","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,15]]}}}