{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T23:31:18Z","timestamp":1788910278378,"version":"build-2803163510"},"reference-count":78,"publisher":"Springer Science and Business Media LLC","issue":"6","license":[{"start":{"date-parts":[[2025,10,29]],"date-time":"2025-10-29T00:00:00Z","timestamp":1761696000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2025,10,29]],"date-time":"2025-10-29T00:00:00Z","timestamp":1761696000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"Universidad de C\u00f3rdoba"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Planta"],"published-print":{"date-parts":[[2025,12]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:sec>\n                    <jats:title>Main conclusion<\/jats:title>\n                    <jats:p>\n                      Inoculation with\n                      <jats:italic>Pseudomonas simiae<\/jats:italic>\n                      WCS417 improves cucumber growth under Fe deficiency conditions and induces iron-deficiency responses, making it a promising candidate for sustainable biofertilization strategies in dicot plants.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Abstract<\/jats:title>\n                    <jats:p>\n                      Iron (Fe) deficiency poses a significant agronomic challenge in calcareous soils, particularly affecting dicot plants. Conventional production methods rely heavily on high-yielding varieties and the application of substantial amounts of agrochemicals, leading to considerable environmental concerns. In this context, leveraging the potential of beneficial rhizosphere microorganisms as biofertilizers represents a highly promising and environmentally sound alternative to chemical fertilizers. This study aims to investigate the efficacy of the nonpathogenic strain\n                      <jats:italic>Pseudomonas simiae<\/jats:italic>\n                      WCS417 in eliciting Fe deficiency responses in cucumber plants, along with its impacts on plant growth and Fe chlorosis. Conducted under hydroponic conditions, our experiments reveal compelling outcomes. Root inoculation of cucumber plants with\n                      <jats:italic>P. simiae<\/jats:italic>\n                      significantly enhances plant growth while concurrently mitigating Fe chlorosis symptoms over successive cultivation days. The inoculation with this bacterium induces acidification in the subapical zone of cucumber roots, facilitating Fe solubility in the rhizosphere. Additionally,\n                      <jats:italic>P. simiae<\/jats:italic>\n                      triggers the upregulation of Fe-related genes in inoculated plants, even under Fe sufficiency. In conclusion,\n                      <jats:italic>P. simiae<\/jats:italic>\n                      emerges as a potent enhancer of Fe deficiency responses in cucumber plants. Its ability to promote growth, enhance Fe solubility through rhizosphere acidification, and alleviate Fe chlorosis underscores its potential as an effective biofertilizer for a sustainable Fe nutrition of dicot plants.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1007\/s00425-025-04844-5","type":"journal-article","created":{"date-parts":[[2025,10,29]],"date-time":"2025-10-29T05:34:47Z","timestamp":1761716087000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["The inoculation with Pseudomonas simiae WCS417 strain promotes growth and the induction of iron-deficiency responses in cucumber plants (Cucumis sativus L.)"],"prefix":"10.1007","volume":"262","author":[{"given":"Miguel A.","family":"Aparicio","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Francisco J.","family":"Ruiz-Castilla","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jos\u00e9","family":"Ramos","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Francisco J.","family":"Romera","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5069-6435","authenticated-orcid":false,"given":"Carlos","family":"Lucena","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,10,29]]},"reference":[{"key":"4844_CR1","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1016\/j.copbio.2021.06.015","volume":"70","author":"D Abedini","year":"2021","unstructured":"Abedini D, Jaupitre S, Bouwmeester H, Dong L (2021) Metabolic interactions in beneficial microbe recruitment by plants. Curr Opin Biotechnol 70:241\u2013247. https:\/\/doi.org\/10.1016\/j.copbio.2021.06.015","journal-title":"Curr Opin Biotechnol"},{"key":"4844_CR2","doi-asserted-by":"publisher","first-page":"e12609","DOI":"10.1016\/j.heliyon.2022.e12609","volume":"8","author":"MMA Allouzi","year":"2022","unstructured":"Allouzi MMA, Allouzi SMA, Keng ZX et al (2022) Liquid biofertilizers as a sustainable solution for agriculture. Heliyon 8:e12609. https:\/\/doi.org\/10.1016\/j.heliyon.2022.e12609","journal-title":"Heliyon"},{"key":"4844_CR3","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1016\/j.mib.2017.03.011","volume":"37","author":"FD Andreote","year":"2017","unstructured":"Andreote FD, Pereira E Silva MDC (2017) Microbial communities associated with plants: learning from nature to apply it in agriculture. Curr Opin Microbiol 37:29\u201334. https:\/\/doi.org\/10.1016\/j.mib.2017.03.011","journal-title":"Curr Opin Microbiol"},{"key":"4844_CR4","doi-asserted-by":"publisher","DOI":"10.1007\/s00425-023-04079-2","volume":"257","author":"MA Aparicio","year":"2023","unstructured":"Aparicio MA, Lucena C, Garc\u00eda MJ et al (2023) The nonpathogenic strain of Fusarium oxysporum FO12 induces Fe deficiency responses in cucumber (Cucumis sativus L.) plants. Planta 257:50. https:\/\/doi.org\/10.1007\/s00425-023-04079-2","journal-title":"Planta"},{"key":"4844_CR5","doi-asserted-by":"publisher","DOI":"10.3390\/plants14020264","volume":"14","author":"MA Aparicio","year":"2025","unstructured":"Aparicio MA, Ruiz-Castilla FJ, Ramos J et al (2025) Pseudomonas simiae WCS417 strain enhances tomato (Solanum lycopersicum L.) plant growth under alkaline conditions. Plants 14:264. https:\/\/doi.org\/10.3390\/plants14020264","journal-title":"Plants"},{"key":"4844_CR6","first-page":"704","volume":"7","author":"M Ashraf","year":"2013","unstructured":"Ashraf M, Asif M, Zaheer A et al (2013) Plant growth promoting rhizobacteria and sustainable agriculture: a review. Afr J Microbiol Res 7:704\u2013709","journal-title":"Afr J Microbiol Res"},{"key":"4844_CR7","doi-asserted-by":"publisher","first-page":"1473","DOI":"10.3389\/fpls.2018.01473","volume":"9","author":"R Backer","year":"2018","unstructured":"Backer R, Rokem JS, Ilangumaran G et al (2018) Plant growth-promoting rhizobacteria: context, mechanisms of action and roadmap to commercialization of biostimulants for sustainable agriculture. Front Plant Sci 9:1473. https:\/\/doi.org\/10.3389\/fpls.2018.01473","journal-title":"Front Plant Sci"},{"key":"4844_CR8","doi-asserted-by":"publisher","first-page":"642","DOI":"10.1016\/j.copbio.2009.09.014","volume":"20","author":"DV Badri","year":"2009","unstructured":"Badri DV, Weir TL, Van Der Lelie D, Vivanco JM (2009) Rhizosphere chemical dialogues: plant\u2013microbe interactions. Curr Opin Biotechnol 20:642\u2013650. https:\/\/doi.org\/10.1016\/j.copbio.2009.09.014","journal-title":"Curr Opin Biotechnol"},{"key":"4844_CR9","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1146\/annurev.arplant.57.032905.105159","volume":"57","author":"HP Bais","year":"2006","unstructured":"Bais HP, Weir TL, Perry LG et al (2006) The role of root exudates in rhizosphere interactions with plants and other organisms. Annu Rev Plant Biol 57:233\u2013266. https:\/\/doi.org\/10.1146\/annurev.arplant.57.032905.105159","journal-title":"Annu Rev Plant Biol"},{"issue":"8","key":"4844_CR10","doi-asserted-by":"publisher","first-page":"478","DOI":"10.1016\/j.tplants.2012.04.001","volume":"17","author":"RL Berendsen","year":"2012","unstructured":"Berendsen RL, Pieterse CM, Bakker PA (2012) The rhizosphere microbiome and plant health. Trends Plant Sci 17(8):478\u2013486","journal-title":"Trends Plant Sci"},{"key":"4844_CR11","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1023\/A:1026290508166","volume":"256","author":"C Bertin","year":"2003","unstructured":"Bertin C, Yang X, Weston LA (2003) The role of root exudates and allelochemicals in the rhizosphere. Plant Soil 256:67\u201383. https:\/\/doi.org\/10.1023\/A:1026290508166","journal-title":"Plant Soil"},{"key":"4844_CR12","doi-asserted-by":"publisher","first-page":"1409","DOI":"10.1038\/s41467-018-03851-3","volume":"9","author":"R Bhosale","year":"2018","unstructured":"Bhosale R, Giri J, Pandey BK et al (2018) A mechanistic framework for auxin dependent Arabidopsis root hair elongation to low external phosphate. Nat Commun 9:1409. https:\/\/doi.org\/10.1038\/s41467-018-03851-3","journal-title":"Nat Commun"},{"key":"4844_CR13","doi-asserted-by":"publisher","first-page":"124","DOI":"10.1016\/j.tplants.2014.11.004","volume":"20","author":"T Brumbarova","year":"2015","unstructured":"Brumbarova T, Bauer P, Ivanov R (2015) Molecular mechanisms governing Arabidopsis iron uptake. Trends Plant Sci 20:124\u2013133. https:\/\/doi.org\/10.1016\/j.tplants.2014.11.004","journal-title":"Trends Plant Sci"},{"key":"4844_CR14","doi-asserted-by":"publisher","first-page":"392","DOI":"10.1016\/j.chom.2015.01.011","volume":"17","author":"D Bulgarelli","year":"2015","unstructured":"Bulgarelli D, Garrido-Oter R, M\u00fcnch PC et al (2015) Structure and function of the bacterial root microbiota in wild and domesticated barley. Cell Host Microbe 17:392\u2013403. https:\/\/doi.org\/10.1016\/j.chom.2015.01.011","journal-title":"Cell Host Microbe"},{"key":"4844_CR15","doi-asserted-by":"publisher","first-page":"428","DOI":"10.1007\/s00114-002-0347-6","volume":"89","author":"A Carrillo","year":"2002","unstructured":"Carrillo A, Li C, Bashan Y (2002) Increased acidification in the rhizosphere of cactus seedlings induced by Azospirillum brasilense. Naturwissenschaften 89:428\u2013432. https:\/\/doi.org\/10.1007\/s00114-002-0347-6","journal-title":"Naturwissenschaften"},{"key":"4844_CR16","doi-asserted-by":"publisher","first-page":"42","DOI":"10.1016\/j.apsoil.2019.06.018","volume":"144","author":"A De Santiago","year":"2019","unstructured":"De Santiago A, Perea-Torres F, Avil\u00e9s M et al (2019) Shifts in microbial community structure influence the availability of Fe and other micronutrients to lupin (Lupinus albus L.). Appl Soil Ecol 144:42\u201350. https:\/\/doi.org\/10.1016\/j.apsoil.2019.06.018","journal-title":"Appl Soil Ecol"},{"key":"4844_CR17","doi-asserted-by":"publisher","first-page":"500","DOI":"10.1016\/j.resmic.2012.08.006","volume":"163","author":"B Drogue","year":"2012","unstructured":"Drogue B, Dor\u00e9 H, Borland S et al (2012) Which specificity in cooperation between phytostimulating rhizobacteria and plants? Res Microbiol 163:500\u2013510. https:\/\/doi.org\/10.1016\/j.resmic.2012.08.006","journal-title":"Res Microbiol"},{"key":"4844_CR18","doi-asserted-by":"publisher","first-page":"277","DOI":"10.1007\/BF01974309","volume":"99","author":"BJ Duijff","year":"1993","unstructured":"Duijff BJ, Meijer JW, Bakker PAHM, Schippers B (1993) Siderophore-mediated competition for iron and induced resistance in the suppression of fusarium wilt of carnation by fluorescent Pseudomonas spp. Neth J Plant Pathol 99:277\u2013289. https:\/\/doi.org\/10.1007\/BF01974309","journal-title":"Neth J Plant Pathol"},{"key":"4844_CR19","doi-asserted-by":"publisher","first-page":"E911","DOI":"10.1073\/pnas.1414592112","volume":"112","author":"J Edwards","year":"2015","unstructured":"Edwards J, Johnson C, Santos-Medell\u00edn C et al (2015) Structure, variation, and assembly of the root-associated microbiomes of rice. Proc Natl Acad Sci USA 112:E911\u2013E920. https:\/\/doi.org\/10.1073\/pnas.1414592112","journal-title":"Proc Natl Acad Sci USA"},{"key":"4844_CR21","doi-asserted-by":"publisher","first-page":"1387","DOI":"10.1126\/science.1112665","volume":"309","author":"J Gans","year":"2005","unstructured":"Gans J, Wolinsky M, Dunbar J (2005) Computational improvements reveal great bacterial diversity and high metal toxicity in soil. Science 309:1387\u20131390. https:\/\/doi.org\/10.1126\/science.1112665","journal-title":"Science"},{"key":"4844_CR22","doi-asserted-by":"publisher","first-page":"1","DOI":"10.6064\/2012\/963401","volume":"2012","author":"BR Glick","year":"2012","unstructured":"Glick BR (2012) Plant growth-promoting bacteria: mechanisms and applications. Scientifica 2012:1\u201315. https:\/\/doi.org\/10.6064\/2012\/963401","journal-title":"Scientifica"},{"key":"4844_CR23","doi-asserted-by":"publisher","first-page":"e14065","DOI":"10.1371\/journal.pone.0014065","volume":"5","author":"NCM Gomes","year":"2010","unstructured":"Gomes NCM, Cleary DFR, Pinto FN et al (2010) Taking root: enduring effect of rhizosphere bacterial colonization in mangroves. PLoS ONE 5:e14065. https:\/\/doi.org\/10.1371\/journal.pone.0014065","journal-title":"PLoS ONE"},{"key":"4844_CR25","doi-asserted-by":"publisher","first-page":"395","DOI":"10.1016\/j.soilbio.2004.08.030","volume":"37","author":"EJ Gray","year":"2005","unstructured":"Gray EJ, Smith DL (2005) Intracellular and extracellular PGPR: commonalities and distinctions in the plant\u2013bacterium signaling processes. Soil Biol Biochem 37:395\u2013412. https:\/\/doi.org\/10.1016\/j.soilbio.2004.08.030","journal-title":"Soil Biol Biochem"},{"key":"4844_CR26","doi-asserted-by":"publisher","first-page":"579","DOI":"10.1007\/s13213-010-0117-1","volume":"60","author":"R Hayat","year":"2010","unstructured":"Hayat R, Ali S, Amara U et al (2010) Soil beneficial bacteria and their role in plant growth promotion: a review. Ann Microbiol 60:579\u2013598. https:\/\/doi.org\/10.1007\/s13213-010-0117-1","journal-title":"Ann Microbiol"},{"key":"4844_CR27","doi-asserted-by":"publisher","first-page":"990","DOI":"10.1080\/01904167.2014.881857","volume":"37","author":"M Ipek","year":"2014","unstructured":"Ipek M, Pirlak L, Esitken A et al (2014) Plant growth-promoting rhizobacteria (PGPR) increase yield, growth and nutrition of strawberry under high-calcareous soil conditions. J Plant Nutr 37:990\u20131001. https:\/\/doi.org\/10.1080\/01904167.2014.881857","journal-title":"J Plant Nutr"},{"key":"4844_CR28","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1093\/mp\/ssr065","volume":"5","author":"R Ivanov","year":"2012","unstructured":"Ivanov R, Brumbarova T, Bauer P (2012) Fitting into the harsh reality: regulation of iron-deficiency responses in dicotyledonous plants. Mol Plant 5:27\u201342. https:\/\/doi.org\/10.1093\/mp\/ssr065","journal-title":"Mol Plant"},{"key":"4844_CR29","doi-asserted-by":"publisher","unstructured":"Khan MN, Mobin M, Abbas ZK, Alamri SA (2018) Fertilizers and their contaminants in soils, surface and groundwater. In: DellaSala DA, Goldstein MI (eds) The encyclopedia of the anthropocene, vol 5. Elsevier, Oxford, pp 225\u2013240. https:\/\/doi.org\/10.1016\/B978-0-12-809665-9.09888-8","DOI":"10.1016\/B978-0-12-809665-9.09888-8"},{"key":"4844_CR30","doi-asserted-by":"publisher","first-page":"e13898","DOI":"10.1111\/ppl.13898","volume":"175","author":"N Khatun","year":"2023","unstructured":"Khatun N, Shinozawa A, Takahashi K et al (2023) Abscisic acid-mediated sugar responses are essential for vegetative desiccation tolerance in the liverwort Marchantia polymorpha. Physiol Plant 175:e13898. https:\/\/doi.org\/10.1111\/ppl.13898","journal-title":"Physiol Plant"},{"key":"4844_CR31","first-page":"301","volume":"44","author":"ED King","year":"1954","unstructured":"King ED, Ward MK, Raney DE (1954) Two simple media for the demonstration of pyocyanin and fluorescin. J Lab Clin Med 44:301\u2013307","journal-title":"J Lab Clin Med"},{"key":"4844_CR32","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1146\/annurev-arplant-042811-105522","volume":"63","author":"T Kobayashi","year":"2012","unstructured":"Kobayashi T, Nishizawa NK (2012) Iron uptake, translocation and regulation in higher plants. Annu Rev Plant Biol 63:131\u2013152. https:\/\/doi.org\/10.1146\/annurev-arplant-042811-105522","journal-title":"Annu Rev Plant Biol"},{"key":"4844_CR33","doi-asserted-by":"crossref","unstructured":"Koo B-.J, Adriano DC, Bolan NS, Barton CD (2005) Root exudates and microorganisms. In: Hillel D (ed) Encyclopedia of soils in the environment, vol 3. Elsevier, Amsterdam, pp 421\u2013428","DOI":"10.1016\/B0-12-348530-4\/00461-6"},{"key":"4844_CR34","doi-asserted-by":"publisher","first-page":"337","DOI":"10.1016\/j.jare.2020.04.014","volume":"24","author":"A Kumar","year":"2020","unstructured":"Kumar A, Dubey A (2020) Rhizosphere microbiome: engineering bacterial competitiveness for enhancing crop production. J Adv Res 24:337\u2013352. https:\/\/doi.org\/10.1016\/j.jare.2020.04.014","journal-title":"J Adv Res"},{"key":"4844_CR35","doi-asserted-by":"publisher","first-page":"1127","DOI":"10.1038\/ismej.2009.61","volume":"3","author":"M Kyselkov\u00e1","year":"2009","unstructured":"Kyselkov\u00e1 M, Kopeck\u00fd J, Frapolli M et al (2009) Comparison of rhizobacterial community composition in soil suppressive or conducive to tobacco black root rot disease. ISME J 3:1127\u20131138. https:\/\/doi.org\/10.1038\/ismej.2009.61","journal-title":"ISME J"},{"key":"4844_CR36","doi-asserted-by":"publisher","first-page":"4145","DOI":"10.1093\/jxb\/erl189","volume":"57","author":"C Lucena","year":"2006","unstructured":"Lucena C, Waters BM, Romera FJ et al (2006) Ethylene could influence ferric reductase, iron transporter, and H+-ATPase gene expression by affecting FER (or FER-like) gene activity. J Exp Bot 57:4145\u20134154. https:\/\/doi.org\/10.1093\/jxb\/erl189","journal-title":"J Exp Bot"},{"key":"4844_CR37","doi-asserted-by":"publisher","first-page":"1002","DOI":"10.1071\/FP07136","volume":"34","author":"C Lucena","year":"2007","unstructured":"Lucena C, Romera FJ, Rojas CL et al (2007) Bicarbonate blocks the expression of several genes involved in the physiological responses to Fe deficiency of strategy i plants. Funct Plant Biol 34:1002. https:\/\/doi.org\/10.1071\/FP07136","journal-title":"Funct Plant Biol"},{"key":"4844_CR38","doi-asserted-by":"publisher","first-page":"1056","DOI":"10.3389\/fpls.2015.01056","volume":"6","author":"C Lucena","year":"2015","unstructured":"Lucena C, Romera FJ, Garc\u00eda MJ et al (2015) Ethylene participates in the regulation of Fe deficiency responses in strategy i plants and in rice. Front Plant Sci 6:1056. https:\/\/doi.org\/10.3389\/fpls.2015.01056","journal-title":"Front Plant Sci"},{"key":"4844_CR39","doi-asserted-by":"publisher","first-page":"148","DOI":"10.3390\/agronomy8080148","volume":"8","author":"C Lucena","year":"2018","unstructured":"Lucena C, Porras R, Romera FJ et al (2018) Similarities and differences in the acquisition of Fe and P by dicot plants. Agronomy 8:148. https:\/\/doi.org\/10.3390\/agronomy8080148","journal-title":"Agronomy"},{"key":"4844_CR40","doi-asserted-by":"publisher","first-page":"541","DOI":"10.1146\/annurev.micro.62.081307.162918","volume":"63","author":"B Lugtenberg","year":"2009","unstructured":"Lugtenberg B, Kamilova F (2009) Plant-growth-promoting rhizobacteria. Annu Rev Microbiol 63:541\u2013556. https:\/\/doi.org\/10.1146\/annurev.micro.62.081307.162918","journal-title":"Annu Rev Microbiol"},{"key":"4844_CR41","doi-asserted-by":"publisher","unstructured":"Marschner H (2012) Mineral nutrition of higher plants, 3rd edn. Academic Press, San Diego. https:\/\/doi.org\/10.1016\/C2009-0-63043-9","DOI":"10.1016\/C2009-0-63043-9"},{"key":"4844_CR42","doi-asserted-by":"publisher","first-page":"2691","DOI":"10.1111\/pce.13016","volume":"40","author":"A Mart\u00ednez-Medina","year":"2017","unstructured":"Mart\u00ednez-Medina A, Van Wees SCM, Pieterse CMJ (2017) Airborne signals from Trichoderma fungi stimulate iron uptake responses in roots resulting in priming of jasmonic acid-dependent defences in shoots of Arabidopsis thaliana and Solanum lycopersicum. Plant Cell Environ 40:2691\u20132705. https:\/\/doi.org\/10.1111\/pce.13016","journal-title":"Plant Cell Environ"},{"key":"4844_CR43","doi-asserted-by":"publisher","first-page":"433","DOI":"10.1007\/s003740050021","volume":"30","author":"J Masalha","year":"2000","unstructured":"Masalha J, Kosegarten H, Elmaci \u00d6, Mengel K (2000) The central role of microbial activity for iron acquisition in maize and sunflower. Biol Fertil Soils 30:433\u2013439. https:\/\/doi.org\/10.1007\/s003740050021","journal-title":"Biol Fertil Soils"},{"key":"4844_CR44","doi-asserted-by":"publisher","first-page":"458","DOI":"10.1016\/j.tim.2009.07.004","volume":"17","author":"C Masson-Boivin","year":"2009","unstructured":"Masson-Boivin C, Giraud E, Perret X, Batut J (2009) Establishing nitrogen-fixing symbiosis with legumes: how many rhizobium recipes? Trends Microbiol 17:458\u2013466. https:\/\/doi.org\/10.1016\/j.tim.2009.07.004","journal-title":"Trends Microbiol"},{"key":"4844_CR45","doi-asserted-by":"publisher","first-page":"226","DOI":"10.1134\/S0026261706020184","volume":"75","author":"OV Merzaeva","year":"2006","unstructured":"Merzaeva OV, Shirokikh IG (2006) Colonization of plant rhizosphere by actinomycetes of different genera. Microbiology 75:226\u2013230. https:\/\/doi.org\/10.1134\/S0026261706020184","journal-title":"Microbiology"},{"key":"4844_CR46","doi-asserted-by":"publisher","first-page":"106","DOI":"10.3389\/fsufs.2020.00106","volume":"4","author":"LA Msimbira","year":"2020","unstructured":"Msimbira LA, Smith DL (2020) The roles of plant growth promoting microbes in enhancing plant tolerance to acidity and alkalinity stresses. Front Sustain Food Syst 4:106. https:\/\/doi.org\/10.3389\/fsufs.2020.00106","journal-title":"Front Sustain Food Syst"},{"key":"4844_CR47","doi-asserted-by":"publisher","first-page":"1224","DOI":"10.3389\/fpls.2015.01224","volume":"6","author":"G Neumann","year":"2016","unstructured":"Neumann G (2016) The role of ethylene in plant adaptations for phosphate acquisition in soils\u2014a review. Front Plant Sci 6:1224. https:\/\/doi.org\/10.3389\/fpls.2015.01224","journal-title":"Front Plant Sci"},{"key":"4844_CR48","unstructured":"Odum EP, Barrett GW (2005) Fundamentals of ecology, 5th edn. Brooks\/Cole, Belmont"},{"key":"4844_CR49","doi-asserted-by":"publisher","first-page":"763","DOI":"10.1038\/nrmicro1987","volume":"6","author":"M Parniske","year":"2008","unstructured":"Parniske M (2008) Arbuscular mycorrhiza: the mother of plant root endosymbioses. Nat Rev Microbiol 6:763\u2013775. https:\/\/doi.org\/10.1038\/nrmicro1987","journal-title":"Nat Rev Microbiol"},{"key":"4844_CR50","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1007\/s11104-020-04786-9","volume":"461","author":"CMJ Pieterse","year":"2021","unstructured":"Pieterse CMJ, Berendsen RL, De Jonge R et al (2021) Pseudomonas simiae WCS417: star track of a model beneficial rhizobacterium. Plant Soil 461:245\u2013263. https:\/\/doi.org\/10.1007\/s11104-020-04786-9","journal-title":"Plant Soil"},{"key":"4844_CR51","doi-asserted-by":"publisher","first-page":"3608","DOI":"10.1099\/mic.0.2007\/009381-0","volume":"153","author":"JF Pothier","year":"2007","unstructured":"Pothier JF, Wisniewski-Dy\u00e9 F, Weiss-Gayet M et al (2007) Promoter-trap identification of wheat seed extract-induced genes in the plant-growth-promoting rhizobacterium Azospirillum brasilense Sp245. Microbiology 153:3608\u20133622. https:\/\/doi.org\/10.1099\/mic.0.2007\/009381-0","journal-title":"Microbiology"},{"key":"4844_CR52","doi-asserted-by":"publisher","first-page":"406","DOI":"10.1016\/j.plaphy.2020.07.045","volume":"155","author":"S Rahimi","year":"2020","unstructured":"Rahimi S, Talebi M, Baninasab B et al (2020) The role of plant growth-promoting rhizobacteria (PGPR) in improving iron acquisition by altering physiological and molecular responses in quince seedlings. Plant Physiol Biochem 155:406\u2013415. https:\/\/doi.org\/10.1016\/j.plaphy.2020.07.045","journal-title":"Plant Physiol Biochem"},{"key":"4844_CR53","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1007\/s11104-009-9895-2","volume":"321","author":"AE Richardson","year":"2009","unstructured":"Richardson AE, Barea J-M, McNeill AM, Prigent-Combaret C (2009) Acquisition of phosphorus and nitrogen in the rhizosphere and plant growth promotion by microorganisms. Plant Soil 321:305\u2013339. https:\/\/doi.org\/10.1007\/s11104-009-9895-2","journal-title":"Plant Soil"},{"key":"4844_CR54","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1007\/s11104-007-9458-3","volume":"302","author":"LFW Roesch","year":"2008","unstructured":"Roesch LFW, Camargo FAO, Bento FM, Triplett EW (2008) Biodiversity of diazotrophic bacteria within the soil, root and stem of field-grown maize. Plant Soil 302:91\u2013104. https:\/\/doi.org\/10.1007\/s11104-007-9458-3","journal-title":"Plant Soil"},{"key":"4844_CR55","doi-asserted-by":"publisher","first-page":"287","DOI":"10.3389\/fpls.2019.00287","volume":"10","author":"FJ Romera","year":"2019","unstructured":"Romera FJ, Garc\u00eda MJ, Lucena C et al (2019) Induced systemic resistance (ISR) and Fe deficiency responses in dicot plants. Front Plant Sci 10:287. https:\/\/doi.org\/10.3389\/fpls.2019.00287","journal-title":"Front Plant Sci"},{"key":"4844_CR56","doi-asserted-by":"publisher","first-page":"354","DOI":"10.1111\/j.1399-3054.1981.tb04512.x","volume":"53","author":"V R\u00f6mheld","year":"1981","unstructured":"R\u00f6mheld V, Marschner H (1981) Iron deficiency stress induced morphological and physiological changes in root tips of sunflower. Physiol Plant 53:354\u2013360. https:\/\/doi.org\/10.1111\/j.1399-3054.1981.tb04512.x","journal-title":"Physiol Plant"},{"key":"4844_CR57","doi-asserted-by":"publisher","unstructured":"Ryan J, Ibrikci H, Delgado A, et al (2012) Significance of phosphorus for agriculture and the environment in the West Asia and North Africa region. In: Sparks DL (ed) Advances in agronomy, vol 114. Elsevier, Amsterdam, pp 91\u2013153. https:\/\/doi.org\/10.1016\/B978-0-12-394275-3.00004-3","DOI":"10.1016\/B978-0-12-394275-3.00004-3"},{"key":"4844_CR58","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1007\/s10725-010-9479-4","volume":"62","author":"V Sandhya","year":"2010","unstructured":"Sandhya V, Ali SkZ, Grover M et al (2010) Effect of plant growth promoting Pseudomonas spp. on compatible solutes, antioxidant status and plant growth of maize under drought stress. Plant Growth Regul 62:21\u201330. https:\/\/doi.org\/10.1007\/s10725-010-9479-4","journal-title":"Plant Growth Regul"},{"key":"4844_CR59","doi-asserted-by":"publisher","first-page":"5729","DOI":"10.3390\/ijms25115729","volume":"25","author":"J Sevillano-Cano","year":"2024","unstructured":"Sevillano-Cano J, Garc\u00eda MJ, C\u00f3rdoba-Galv\u00e1n C et al (2024) Exploring the role of Debaryomyces hansenii as biofertilizer in iron-deficient environments to enhance plant nutrition and crop production sustainability. Int J Mol Sci 25:5729. https:\/\/doi.org\/10.3390\/ijms25115729","journal-title":"Int J Mol Sci"},{"key":"4844_CR60","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1080\/17429145.2010.545147","volume":"6","author":"A ShaikZ","year":"2011","unstructured":"ShaikZ A, Sandhya V, Grover M et al (2011) Effect of inoculation with a thermotolerant plant growth promoting Pseudomonas putida strain AKMP7 on growth of wheat (Triticum spp.) under heat stress. J Plant Interact 6:239\u2013246. https:\/\/doi.org\/10.1080\/17429145.2010.545147","journal-title":"J Plant Interact"},{"key":"4844_CR61","doi-asserted-by":"publisher","first-page":"77","DOI":"10.1078\/0944-5013-00182","volume":"158","author":"A Sharma","year":"2003","unstructured":"Sharma A, Johri BN (2003) Combat of iron-deprivation through a plant growth promoting fluorescent Pseudomonas strain GRP3A in mung bean (Vigna radiata L. Wilzeck). Microbiol Res 158:77\u201381. https:\/\/doi.org\/10.1078\/0944-5013-00182","journal-title":"Microbiol Res"},{"key":"4844_CR62","doi-asserted-by":"publisher","first-page":"1181","DOI":"10.1093\/jxb\/ers342","volume":"64","author":"J Shen","year":"2013","unstructured":"Shen J, Li C, Mi G et al (2013) Maximizing root\/rhizosphere efficiency to improve crop productivity and nutrient use efficiency in intensive agriculture of China. J Exp Bot 64:1181\u20131192. https:\/\/doi.org\/10.1093\/jxb\/ers342","journal-title":"J Exp Bot"},{"key":"4844_CR63","doi-asserted-by":"publisher","first-page":"796","DOI":"10.3389\/fpls.2015.00796","volume":"6","author":"L Song","year":"2015","unstructured":"Song L, Liu D (2015) Ethylene and plant responses to phosphate deficiency. Front Plant Sci 6:796. https:\/\/doi.org\/10.3389\/fpls.2015.00796","journal-title":"Front Plant Sci"},{"key":"4844_CR64","doi-asserted-by":"publisher","first-page":"e2891","DOI":"10.7717\/peerj.2891","volume":"5","author":"MG Stetter","year":"2017","unstructured":"Stetter MG, Benz M, Ludewig U (2017) Increased root hair density by loss of WRKY6 in Arabidopsis thaliana. PeerJ 5:e2891. https:\/\/doi.org\/10.7717\/peerj.2891","journal-title":"PeerJ"},{"key":"4844_CR65","doi-asserted-by":"publisher","first-page":"E5213","DOI":"10.1073\/pnas.1722335115","volume":"115","author":"IA Stringlis","year":"2018","unstructured":"Stringlis IA, Yu K, Feussner K et al (2018) MYB72-dependent coumarin exudation shapes root microbiome assembly to promote plant health. Proc Natl Acad Sci USA 115:E5213\u2013E5222. https:\/\/doi.org\/10.1073\/pnas.1722335115","journal-title":"Proc Natl Acad Sci USA"},{"key":"4844_CR66","first-page":"2","volume":"13","author":"D Torres","year":"2004","unstructured":"Torres D, Capote T (2004) Agroqu\u00edmicos un problema ambiental global: uso del an\u00e1lisis qu\u00edmico como herramienta para el monitoreo ambiental. Rev Ecosistemas 13:2\u20136","journal-title":"Rev Ecosistemas"},{"key":"4844_CR67","doi-asserted-by":"publisher","first-page":"675","DOI":"10.1104\/pp.15.01537","volume":"171","author":"P Trapet","year":"2016","unstructured":"Trapet P, Avoscan L, Klinguer A et al (2016) The Pseudomonas fluorescens siderophore pyoverdine weakens Arabidopsis thaliana defense in favor of growth in iron-deficient conditions. Plant Physiol 171:675\u2013693. https:\/\/doi.org\/10.1104\/pp.15.01537","journal-title":"Plant Physiol"},{"key":"4844_CR68","doi-asserted-by":"publisher","first-page":"243","DOI":"10.1007\/s10658-007-9165-1","volume":"119","author":"LC Van Loon","year":"2007","unstructured":"Van Loon LC (2007) Plant responses to plant growth-promoting rhizobacteria. Eur J Plant Pathol 119:243\u2013254. https:\/\/doi.org\/10.1007\/s10658-007-9165-1","journal-title":"Eur J Plant Pathol"},{"key":"4844_CR69","doi-asserted-by":"publisher","first-page":"909","DOI":"10.3389\/fpls.2019.00909","volume":"10","author":"EH Verbon","year":"2019","unstructured":"Verbon EH, Trapet PL, Kruijs S et al (2019) Rhizobacteria-mediated activation of the Fe deficiency response in Arabidopsis roots: impact on Fe status and signaling. Front Plant Sci 10:909. https:\/\/doi.org\/10.3389\/fpls.2019.00909","journal-title":"Front Plant Sci"},{"key":"4844_CR70","doi-asserted-by":"publisher","first-page":"530","DOI":"10.1016\/j.pbi.2008.06.013","volume":"11","author":"EL Walker","year":"2008","unstructured":"Walker EL, Connolly EL (2008) Time to pump iron: iron-deficiency-signaling mechanisms of higher plants. Curr Opin Plant Biol 11:530\u2013535. https:\/\/doi.org\/10.1016\/j.pbi.2008.06.013","journal-title":"Curr Opin Plant Biol"},{"key":"4844_CR71","doi-asserted-by":"publisher","first-page":"293","DOI":"10.1016\/j.plaphy.2007.03.011","volume":"45","author":"BM Waters","year":"2007","unstructured":"Waters BM, Lucena C, Romera FJ et al (2007) Ethylene involvement in the regulation of the H+-ATPase CsHA1 gene and of the new isolated ferric reductase CsFRO1 and iron transporter CsIRT1 genes in cucumber plants. Plant Physiol Biochem 45:293\u2013301. https:\/\/doi.org\/10.1016\/j.plaphy.2007.03.011","journal-title":"Plant Physiol Biochem"},{"key":"4844_CR72","doi-asserted-by":"publisher","first-page":"1493","DOI":"10.3389\/fpls.2018.01493","volume":"9","author":"A Williams","year":"2018","unstructured":"Williams A, P\u00e9triacq P, Beerling DJ et al (2018) Impacts of atmospheric CO2 and soil nutritional value on plant responses to rhizosphere colonization by soil bacteria. Front Plant Sci 9:1493. https:\/\/doi.org\/10.3389\/fpls.2018.01493","journal-title":"Front Plant Sci"},{"issue":"3","key":"4844_CR73","doi-asserted-by":"publisher","first-page":"386","DOI":"10.3390\/plants11030386","volume":"11","author":"Y Yu","year":"2022","unstructured":"Yu Y, Gui Y, Li Z, Jiang C et al (2022) Induced systemic resistance for improving plant immunity by beneficial microbes. Plants 11(3):386","journal-title":"Plants"},{"key":"4844_CR74","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1016\/j.tplants.2008.02.003","volume":"13","author":"JS Yuan","year":"2008","unstructured":"Yuan JS, Galbraith DW, Dai SY et al (2008) Plant systems biology comes of age. Trends Plant Sci 13:165\u2013171. https:\/\/doi.org\/10.1016\/j.tplants.2008.02.003","journal-title":"Trends Plant Sci"},{"issue":"1","key":"4844_CR75","doi-asserted-by":"publisher","first-page":"304","DOI":"10.1104\/pp.112.212597","volume":"162","author":"C Zamioudis","year":"2013","unstructured":"Zamioudis C, Mastranesti P, Dhonukshe P et al (2013) Unraveling root developmental programs initiated by beneficial Pseudomonas spp. bacteria. Plant Physiol 162(1):304\u2013318","journal-title":"Plant Physiol"},{"key":"4844_CR76","doi-asserted-by":"publisher","first-page":"368","DOI":"10.1111\/nph.12980","volume":"204","author":"C Zamioudis","year":"2014","unstructured":"Zamioudis C, Hanson J, Pieterse CMJ (2014) \u03b2-Glucosidase BGLU42 is a MYB72-dependent key regulator of rhizobacteria-induced systemic resistance and modulates iron deficiency responses in Arabidopsis roots. New Phytol 204:368\u2013379. https:\/\/doi.org\/10.1111\/nph.12980","journal-title":"New Phytol"},{"key":"4844_CR77","doi-asserted-by":"publisher","first-page":"309","DOI":"10.1111\/tpj.12995","volume":"84","author":"C Zamioudis","year":"2015","unstructured":"Zamioudis C, Korteland J, Van Pelt JA et al (2015) Rhizobacterial volatiles and photosynthesis-related signals coordinate MYB72 expression in Arabidopsis roots during onset of induced systemic resistance and iron-deficiency responses. Plant J 84:309\u2013322. https:\/\/doi.org\/10.1111\/tpj.12995","journal-title":"Plant J"},{"key":"4844_CR78","doi-asserted-by":"publisher","first-page":"568","DOI":"10.1111\/j.1365-313X.2009.03803.x","volume":"58","author":"H Zhang","year":"2009","unstructured":"Zhang H, Sun Y, Xie X et al (2009) A soil bacterium regulates plant acquisition of iron via deficiency-inducible mechanisms. Plant J 58:568\u2013577. https:\/\/doi.org\/10.1111\/j.1365-313X.2009.03803.x","journal-title":"Plant J"},{"key":"4844_CR79","doi-asserted-by":"publisher","first-page":"162","DOI":"10.1016\/j.plaphy.2016.04.025","volume":"105","author":"C Zhou","year":"2016","unstructured":"Zhou C, Guo J, Zhu L et al (2016) Paenibacillus polymyxa BFKC01 enhances plant iron absorption via improved root systems and activated iron acquisition mechanisms. Plant Physiol Biochem 105:162\u2013173. https:\/\/doi.org\/10.1016\/j.plaphy.2016.04.025","journal-title":"Plant Physiol Biochem"},{"key":"4844_CR80","doi-asserted-by":"publisher","first-page":"723","DOI":"10.1111\/j.1574-6976.2008.00123.x","volume":"32","author":"I Zilber-Rosenberg","year":"2008","unstructured":"Zilber-Rosenberg I, Rosenberg E (2008) Role of microorganisms in the evolution of animals and plants: the hologenome theory of evolution. FEMS Microbiol Rev 32:723\u2013735. https:\/\/doi.org\/10.1111\/j.1574-6976.2008.00123.x","journal-title":"FEMS Microbiol Rev"}],"container-title":["Planta"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00425-025-04844-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00425-025-04844-5\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00425-025-04844-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,11,15]],"date-time":"2025-11-15T10:40:42Z","timestamp":1763203242000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00425-025-04844-5"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,10,29]]},"references-count":78,"journal-issue":{"issue":"6","published-print":{"date-parts":[[2025,12]]}},"alternative-id":["4844"],"URL":"https:\/\/doi.org\/10.1007\/s00425-025-04844-5","relation":{},"ISSN":["0032-0935","1432-2048"],"issn-type":[{"value":"0032-0935","type":"print"},{"value":"1432-2048","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,10,29]]},"assertion":[{"value":"8 March 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"9 October 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 October 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no conflict of interest.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Conflict of interest"}}],"article-number":"136"}}