{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,13]],"date-time":"2026-10-13T22:45:51Z","timestamp":1791931551092,"version":"4.3.4"},"reference-count":92,"publisher":"MDPI AG","issue":"6","license":[{"start":{"date-parts":[[2023,3,7]],"date-time":"2023-03-07T00:00:00Z","timestamp":1678147200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Mitacs, Canada","award":["(IT08347)"],"award-info":[{"award-number":["(IT08347)"]}]},{"name":"cadian Seaplants Limited","award":["(IT08347)"],"award-info":[{"award-number":["(IT08347)"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Abiotic stresses, including salinity stress, affect numerous crops, causing yield reduction, and, as a result, important economic losses. Extracts from the brown alga Ascophyllum nodosum (ANE), and compounds secreted by the Pseudomonas protegens strain, CHA0, can mitigate these effects by inducing tolerance against salt stress. However, the influence of ANE on P. protegens CHA0 secretion, and the combined effects of these two biostimulants on plant growth, are not known. Fucoidan, alginate, and mannitol are abundant components of brown algae and of ANE. Reported here are the effects of a commercial formulation of ANE, fucoidan, alginate, and mannitol, on pea (Pisum sativum), and on the plant growth-promoting activity of P. protegens CHA0. In most situations, ANE and fucoidan increased indole-3-acetic acid (IAA) and siderophore production, phosphate solubilization, and hydrogen cyanide (HCN) production by P. protegens CHA0. Colonization of pea roots by P. protegens CHA0 was found to be increased mostly by ANE and fucoidan in normal conditions and under salt stress. Applications of P. protegens CHA0 combined with ANE, or with fucoidan, alginate, and mannitol, generally augmented root and shoot growth in normal and salinity stress conditions. Real-time quantitative PCR analyses of P. protegens revealed that, in many instances, ANE and fucoidan enhanced the expression of several genes involved in chemotaxis (cheW and WspR), pyoverdine production (pvdS), and HCN production (hcnA), but gene expression patterns overlapped only occasionally those of growth-promoting parameters. Overall, the increased colonization and the enhanced activities of P. protegens CHA0 in the presence of ANE and its components mitigated salinity stress in pea. Among treatments, ANE and fucoidan were found responsible for most of the increased activities of P. protegens CHA0 and the improved plant growth.<\/jats:p>","DOI":"10.3390\/plants12061208","type":"journal-article","created":{"date-parts":[[2023,3,7]],"date-time":"2023-03-07T02:07:38Z","timestamp":1678154858000},"page":"1208","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":19,"title":["A Plant Biostimulant from Ascophyllum nodosum Potentiates Plant Growth Promotion and Stress Protection Activity of Pseudomonas protegens CHA0"],"prefix":"10.3390","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5480-0677","authenticated-orcid":false,"given":"Jai Singh","family":"Patel","sequence":"first","affiliation":[{"name":"Department of Plant Food and Environmental Sciences, Dalhousie University, Halifax, NS B3H 4R2, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vinodkumar","family":"Selvaraj","sequence":"additional","affiliation":[{"name":"Department of Plant Food and Environmental Sciences, Dalhousie University, Halifax, NS B3H 4R2, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7452-5916","authenticated-orcid":false,"given":"Prashant","family":"More","sequence":"additional","affiliation":[{"name":"Department of Plant Food and Environmental Sciences, Dalhousie University, Halifax, NS B3H 4R2, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3467-2089","authenticated-orcid":false,"given":"Ramin","family":"Bahmani","sequence":"additional","affiliation":[{"name":"Department of Plant Food and Environmental Sciences, Dalhousie University, Halifax, NS B3H 4R2, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2527-6551","authenticated-orcid":false,"given":"Tudor","family":"Borza","sequence":"additional","affiliation":[{"name":"Department of Plant Food and Environmental Sciences, Dalhousie University, Halifax, NS B3H 4R2, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Balakrishnan","family":"Prithiviraj","sequence":"additional","affiliation":[{"name":"Department of Plant Food and Environmental Sciences, Dalhousie University, Halifax, NS B3H 4R2, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2023,3,7]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1126\/science.218.4571.443","article-title":"Plant productivity and environment","volume":"218","author":"Boyer","year":"1982","journal-title":"Science"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"941","DOI":"10.1104\/pp.124.3.941","article-title":"Genetic analysis of plant salt tolerance using Arabidopsis","volume":"124","author":"Zhu","year":"2000","journal-title":"Plant Physiol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1046\/j.0016-8025.2001.00808.x","article-title":"Comparative physiology of salt and water stress","volume":"25","author":"Munns","year":"2002","journal-title":"Plant Cell Environ."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"756","DOI":"10.1007\/s12033-012-9624-6","article-title":"How the nucleus and mitochondria communicate in energy production during stress: Nuclear MtATP6, an early-stress responsive gene, regulates the mitochondrial F1F0-ATP synthase complex","volume":"54","author":"Moghadam","year":"2013","journal-title":"Mol. Biotechnol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1146\/annurev.arplant.59.032607.092911","article-title":"Mechanisms of salinity tolerance","volume":"59","author":"Munns","year":"2008","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"S3","DOI":"10.1017\/S0007114512000852","article-title":"Review of the health benefits of peas (Pisum sativum L.)","volume":"108","author":"Dahl","year":"2012","journal-title":"Br. J. Nutr."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Acosta-Motos, J.R., Ortu\u00f1o, M.F., Bernal-Vicente, A., Diaz-Vivancos, P., Sanchez-Blanco, M.J., and Hernandez, J.A. (2017). Plant responses to salt stress: Adaptive mechanisms. Agronomy, 7.","DOI":"10.20944\/preprints201702.0083.v2"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"340","DOI":"10.3389\/fpls.2019.00340","article-title":"Submergence and waterlogging stress in plants: A review highlighting research opportunities and understudied aspects","volume":"10","author":"Fukao","year":"2019","journal-title":"Front. Plant Sci."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"277","DOI":"10.17660\/eJHS.2017\/82.6.2","article-title":"Plant biostimulants: Innovative tool for enhancing plant nutrition in organic farming","volume":"82","author":"Rouphael","year":"2018","journal-title":"Eur. J. Hortic. Sci."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"40","DOI":"10.3389\/fpls.2020.00040","article-title":"Editorial: Biostimulants in agriculture","volume":"11","author":"Rouphael","year":"2020","journal-title":"Front. Plant Sci."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.scienta.2015.10.044","article-title":"Biostimulants in horticulture","volume":"196","author":"Colla","year":"2015","journal-title":"Sci. Hortic."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/j.plaphy.2018.02.024","article-title":"Ascophyllum nodosum extract biostimulants and their role in enhancing tolerance to drought stress in tomato plants","volume":"126","author":"Quille","year":"2018","journal-title":"Plant Physiol. Biochem."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1007\/s00344-018-9861-4","article-title":"Physiological and transcriptomics analyses reveal that Ascophyllum nodosum extracts induce salinity tolerance in Arabidopsis by regulating the expression of stress responsive genes","volume":"38","author":"Jithesh","year":"2019","journal-title":"J. Plant Growth Regul."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"655","DOI":"10.3389\/fpls.2019.00655","article-title":"Ascophyllum nodosum-based biostimulants: Sustainable applications in agriculture for the stimulation of plant growth, stress tolerance, and disease management","volume":"10","author":"Shukla","year":"2019","journal-title":"Front. Plant Sci."},{"key":"ref_15","first-page":"573","article-title":"Toward the molecular understanding of the action mechanism of Ascophyllum nodosum extracts on plants","volume":"32","author":"Vereecke","year":"2019","journal-title":"J. Appl. Phycol."},{"key":"ref_16","first-page":"plx051","article-title":"Seaweed extract improve drought tolerance of soybean by regulating stress-response genes","volume":"10","author":"Shukla","year":"2017","journal-title":"AoBP"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/j.scienta.2014.12.004","article-title":"Effects of A. nodosum seaweed extracts on spinach growth, physiology and nutrition value under drought stress","volume":"183","author":"Xu","year":"2015","journal-title":"Sci. Hortic."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1873","DOI":"10.1080\/00103624.2013.790404","article-title":"A commercial extract of brown macroalga (Ascophyllum nodosum) affects yield and the nutritional quality of spinach in vitro","volume":"44","author":"Fan","year":"2013","journal-title":"Commun. Soil Sci. Plant Anal."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/j.scienta.2015.09.012","article-title":"Seaweed extracts as biostimulants in horticulture","volume":"196","author":"Battacharyya","year":"2015","journal-title":"Sci. Hortic."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"292","DOI":"10.14295\/cs.v9i2.2134","article-title":"Is seaweed extract an elicitor compound? Changing proline content in drought-stressed bean plants","volume":"9","author":"Carvalho","year":"2018","journal-title":"Comun. Sci."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1362","DOI":"10.3389\/fpls.2017.01362","article-title":"Ascophyllum nodosum seaweed extract alleviates drought stress in Arabidopsis by affecting photosynthetic performance and related gene expression","volume":"8","author":"Santaniello","year":"2017","journal-title":"Front. Plant Sci."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1007\/s10658-011-9802-6","article-title":"Extracts of the marine brown macroalga, Ascophyllum nodosum, induce jasmonic acid dependent systemic resistance in Arabidopsis thaliana against Pseudomonas syringae pv. tomato DC3000 and Sclerotinia sclerotiorum","volume":"131","author":"Subramanian","year":"2011","journal-title":"Eur. J. Plant Pathol."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Shukla, P.S., Borza, T., Critchley, A.T., Hiltz, D., Norrie, J., and Prithiviraj, B. (2018). Ascophyllum nodosum extract mitigates salinity stress in Arabidopsis thaliana by modulating the expression of miRNA involved in stress tolerance and nutrient acquisition. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0206221"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"2675","DOI":"10.1007\/s10811-018-1439-9","article-title":"Ascophyllum nodosum-based algal extracts act as enhancers of growth, fruit quality, and adaptation to stress in salinized tomato plants","volume":"30","author":"Silletti","year":"2018","journal-title":"J. Appl. Phycol."},{"key":"ref_25","first-page":"357","article-title":"The initial growth of passion fruit plant irrigated with saline water and the application of biostimulants","volume":"10","author":"Gomes","year":"2018","journal-title":"J. Agric. Sci."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Bonomelli, C., Celis, V., Lombardi, G., and M\u00e1rtiz, J. (2018). Salt sress effects on avocado (Persea americana Mill.) plants with and without seaweed extract (Ascophyllum nodosum) application. Agronomy, 8.","DOI":"10.3390\/agronomy8050064"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"198","DOI":"10.2135\/cropsci1994.0011183X003400010035x","article-title":"Alleviation of salinity stress in Kentucky bluegrass by plant growth regulators and iron","volume":"34","author":"Nabati","year":"1994","journal-title":"Crop Sci."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"10537","DOI":"10.1038\/s41598-017-07692-w","article-title":"Sargassum muticum and Jania rubens regulate amino acid metabolism to improve growth and alleviate salinity in chickpea","volume":"7","author":"Srivastava","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"386","DOI":"10.1007\/s00344-009-9103-x","article-title":"Seaweed extracts as biostimulants of plant growth and development","volume":"28","author":"Khan","year":"2009","journal-title":"J. Plant Growth Regul."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1002\/jctb.5000670905","article-title":"The seasonal variation in chemical composition of some of the littoral seaweeds common to Scotland. Part I","volume":"67","author":"Black","year":"1948","journal-title":"Ascophyllum nodosum. J. Soc. Chem. Ind."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1017\/S0025315400056186","article-title":"The seasonal variation in weight and chemical composition of the common British Laminariaceae","volume":"29","author":"Black","year":"1950","journal-title":"J. Mar. Biol. Assoc. UK"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1080\/00071617900650121","article-title":"The polysaccharides of green, red and brown seaweeds: Their basic structure, biosynthesis and function","volume":"14","author":"Percival","year":"1979","journal-title":"Eur. J. Phycol."},{"key":"ref_33","first-page":"126","article-title":"Sulfated and non-sulfated polysaccharides from seaweeds and their uses: An overview","volume":"14","author":"Venugopal","year":"2019","journal-title":"ECronicon Nutr."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"486","DOI":"10.1016\/j.tplants.2016.01.006","article-title":"Mannitol in Plants, Fungi, and Plant-Fungal Interactions","volume":"21","author":"Patel","year":"2016","journal-title":"Trends Plant Sci."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"650","DOI":"10.1007\/s00344-018-9880-1","article-title":"Endophytic bacteria in plant salt stress tolerance: Current and future prospects","volume":"38","author":"Vaishnav","year":"2019","journal-title":"J. Plant Growth Regul."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1016\/j.micres.2018.11.005","article-title":"Interactions between Pseudomonas spp. and their role in improving the red pepper plant growth under salinity stress","volume":"219","author":"Samaddar","year":"2019","journal-title":"Microbiol. Res."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"158","DOI":"10.5958\/j.2229-4473.27.1.025","article-title":"Studies on rhizosphere-bacteria mediated biotic and abiotic stress tolerance in chickpea (Cicer arietinum L.)","volume":"27","author":"Sarkar","year":"2014","journal-title":"Vegetos"},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Chu, T.N., Tran, B.T.H., Van Bui, L., and Hoang, M.T.T. (2019). Plant growth-promoting rhizobacterium Pseudomonas PS01 induces salt tolerance in Arabidopsis thaliana. BMC Res. Notes, 12.","DOI":"10.1186\/s13104-019-4046-1"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"581","DOI":"10.1038\/nprot.2006.83","article-title":"The single-step method of RNA isolation by acid guanidinium thiocyanate\u2013phenol\u2013chloroform extraction: Twenty-something years on","volume":"1","author":"Chomczynski","year":"2006","journal-title":"Nat. Protoc."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1392","DOI":"10.3389\/fmicb.2019.01392","article-title":"The production of ACC deaminase and trehalose by the plant growth promoting bacterium Pseudomonas sp. UW4 synergistically protect tomato plants against salt stress","volume":"10","author":"Duan","year":"2019","journal-title":"Front. Microbiol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1297","DOI":"10.3389\/fmicb.2018.01297","article-title":"Gene expression patterns in roots of Camelina sativa with enhanced salinity tolerance arising from inoculation of soil with plant growth promoting bacteria producing 1-aminocyclopropane-1-carboxylate deaminase or expression the corresponding acdS gene","volume":"9","author":"Heydarian","year":"2018","journal-title":"Front. Microbiol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1506","DOI":"10.3389\/fmicb.2019.01506","article-title":"ACC deaminase producing bacteria with multifarious plant growth promoting traits alleviates salinity stress in French bean (Phaseolus vulgaris) plants","volume":"10","author":"Gupta","year":"2019","journal-title":"Front. Microbiol."},{"key":"ref_43","first-page":"17","article-title":"Pseudomonas chemotaxis","volume":"39","author":"Sampedro","year":"2014","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"14422","DOI":"10.1073\/pnas.0507170102","article-title":"A chemosensory system that regulates biofilm formation through modulation of cyclic diguanylate levels","volume":"102","author":"Hickman","year":"2005","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Mart\u00ednez-Granero, F., Navazo, A., Barahona, E., Redondo-Nieto, M., Gonz\u00e1lez de Heredia, E., Baena, I., Mart\u00edn-Mart\u00edn, I., Rivilla, R., and Mart\u00edn, M. (2014). Identification of flgZ as a flagellar gene encoding a PilZ domain protein that regulates swimming motility and biofilm formation in Pseudomonas. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0087608"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1111\/j.1574-6968.2001.tb10695.x","article-title":"Dual control of hydrogen cyanide biosynthesis by the global activator GacA in Pseudomonas aeruginosa PAO1","volume":"200","author":"Pessi","year":"2001","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"2159","DOI":"10.1099\/mic.0.000391","article-title":"The global regulator ANR is essential for Pseudomonas chlororaphis strain PA23 biocontrol","volume":"162","author":"Nandi","year":"2016","journal-title":"Microbiology"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"3187","DOI":"10.1128\/JB.180.12.3187-3196.1998","article-title":"Characterization of the hcnABC gene cluster encoding hydrogen cyanide synthase and anaerobic regulation by ANR in the strictly aerobic biocontrol agent Pseudomonas fluorescens CHA0","volume":"180","author":"Laville","year":"1998","journal-title":"J. Bacteriol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1630","DOI":"10.1111\/j.1462-2920.2010.02210.x","article-title":"Transcriptional control of the pvdS iron starvation sigma factor gene by the master regulator of sulfur metabolism CysB in Pseudomonas aeruginosa","volume":"12","author":"Imperi","year":"2010","journal-title":"Environ. Microbiol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"424","DOI":"10.15698\/mic2018.10.649","article-title":"The biosynthesis of pyoverdines","volume":"5","author":"Ringel","year":"2018","journal-title":"Microb. Cell"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"478","DOI":"10.1016\/j.tplants.2012.04.001","article-title":"The rhizosphere microbiome and plant health","volume":"17","author":"Berendsen","year":"2012","journal-title":"Trends Plant Sci."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1111\/j.1469-8137.2004.01130.x","article-title":"Plant and mycorrhizal regulation of rhizodeposition","volume":"163","author":"Jones","year":"2004","journal-title":"New Phytol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"666","DOI":"10.1111\/j.1365-3040.2009.01926.x","article-title":"Regulation and function of root exudates","volume":"32","author":"Badri","year":"2009","journal-title":"Plant Cell Environ."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"256","DOI":"10.1016\/j.tplants.2016.01.008","article-title":"Metabolomics in the rhizosphere: Tapping into belowground chemical communication","volume":"21","author":"Bouwmeester","year":"2016","journal-title":"Trends Plant Sci."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Gunupuru, L.R., Patel, J.S., Sumarah, M.W., Renaud, J.B., Mantin, E.G., and Prithiviraj, B. (2019). A plant biostimulant made from the marine brown algae Ascophyllum nodosum and chitosan reduce Fusarium head blight and mycotoxin contamination in wheat. PLoS ONE, 14.","DOI":"10.1371\/journal.pone.0220562"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1007\/BF02186237","article-title":"Seaweed liquid fertilizer from Ascophyllum nodosum contains elicitors of plantd-glycanases","volume":"5","author":"Patier","year":"1993","journal-title":"J. Appl. Phycol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"306","DOI":"10.3389\/fmicb.2017.00306","article-title":"Trichoderma asperellum (T42) and Pseudomonas fluorescens (OKC)-enhances resistance of pea against Erysiphe pisi through enhanced ROS generation and lignifications","volume":"8","author":"Patel","year":"2017","journal-title":"Front. Microbiol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1111\/jph.12358","article-title":"Streptomyces rochei SM3 induces stress tolerance in chickpea against Sclerotinia sclerotiorum and NaCl","volume":"163","author":"Srivastava","year":"2015","journal-title":"J. Phytopathol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1016\/S0065-2296(07)46004-9","article-title":"Iron dynamics in plants","volume":"46","author":"Briat","year":"2007","journal-title":"Adv. Bot. Res."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"2273","DOI":"10.1016\/j.febslet.2007.04.043","article-title":"Mining iron: Iron uptake and transport in plants","volume":"581","author":"Kim","year":"2007","journal-title":"FEBS Lett."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"2167","DOI":"10.1104\/pp.113.233585","article-title":"Scavenging iron: A novel mechanism of plant immunity activation by microbial siderophores","volume":"164","author":"Aznar","year":"2014","journal-title":"Plant Physiol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1104\/pp.15.01537","article-title":"The Pseudomonas fluorescens siderophore pyoverdine weakens Arabidopsis thaliana defense in favor of growth in iron-deficient conditions","volume":"171","author":"Trapet","year":"2016","journal-title":"Plant Physiol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"2681","DOI":"10.1128\/JB.01376-13","article-title":"PvdP is a tyrosinase that drives maturation of the pyoverdine chromophore in Pseudomonas aeruginosa","volume":"196","author":"Koch","year":"2014","journal-title":"J. Bacteriol."},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Bathke, J., Konzer, A., Remes, B., McIntosh, M., and Klug, G. (2019). Comparative analyses of the variation of the transcriptome and proteome of Rhodobacter sphaeroides throughout growth. BMC Genom., 20.","DOI":"10.1186\/s12864-019-5749-3"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"535","DOI":"10.1016\/j.cell.2016.03.014","article-title":"On the dependency of cellular protein levels on mRNA abundance","volume":"165","author":"Liu","year":"2016","journal-title":"Cell"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"891","DOI":"10.1093\/bib\/bbv090","article-title":"Transcriptomic and metabolomic data integration","volume":"17","author":"Cavill","year":"2015","journal-title":"Brief. Bioinform."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"103997","DOI":"10.1016\/j.biocontrol.2019.05.016","article-title":"Phosphate solubilizers as antagonists for bacterial leaf blight with improved rice growth in phosphorus deficit soil","volume":"136","author":"Rasul","year":"2019","journal-title":"Biol. Control"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"4162","DOI":"10.1128\/AEM.00295-09","article-title":"Role of gluconic acid production in the regulation of biocontrol traits of Pseudomonas fluorescens CHA0","volume":"75","author":"Keel","year":"2009","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"1408","DOI":"10.1039\/b817075b","article-title":"Genomics of secondary metabolite production by Pseudomonas spp","volume":"26","author":"Gross","year":"2009","journal-title":"Nat. Prod. Rep."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"34768","DOI":"10.1038\/srep34768","article-title":"Plant growth promoting rhizobacteria Dietzia natronolimnaea modulates the expression of stress responsive genes providing protection of wheat from salinity stress","volume":"6","author":"Bharti","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"2097","DOI":"10.1038\/s41598-019-38702-8","article-title":"Comparative physiological and metabolic analysis reveals a complex mechanism involved in drought tolerance in chickpea (Cicer arietinum L.) induced by PGPR and PGRs","volume":"9","author":"Khan","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1093\/jxb\/erh010","article-title":"Plant growth-promoting bacteria and nitrate availability: Impacts on root development and nitrate uptake","volume":"55","author":"Mantelin","year":"2004","journal-title":"J. Exp. Bot."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"1173","DOI":"10.1094\/MPMI.2002.15.11.1173","article-title":"Flagella-driven chemotaxis towards exudate components is an important trait for tomato root colonization by Pseudomonas fluorescens","volume":"15","author":"Vermeiren","year":"2002","journal-title":"Mol. Plant-Microbe Interact."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1111\/1462-2920.14472","article-title":"Recognition of dominant attractants by key chemoreceptors mediates recruitment of plant growth-promoting rhizobacteria","volume":"21","author":"Feng","year":"2019","journal-title":"Environ. Microbiol."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1016\/j.mib.2018.02.002","article-title":"Stimulus sensing and signal processing in bacterial chemotaxis","volume":"45","author":"Bi","year":"2018","journal-title":"Curr. Opin. Microbiol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/j.1462-2920.2007.01424.x","article-title":"High incidence of plant growth-stimulating bacteria associated with the rhizosphere of wheat grown on salinated soil in Uzbekistan","volume":"10","author":"Egamberdieva","year":"2008","journal-title":"Environ. Microbiol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"406","DOI":"10.5423\/PPJ.OA.03.2019.0066","article-title":"A biostimulant preparation of brown seaweed Ascophyllum nodosum suppresses powdery mildew of strawberry","volume":"35","author":"Bajpai","year":"2019","journal-title":"Plant. Pathol. J."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1007\/s10811-007-9228-x","article-title":"Seaweeds from Vietnam used for functional food, medicine and biofertilizer","volume":"19","author":"Hong","year":"2007","journal-title":"J. Appl. Phycol."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1007\/s10811-010-9583-x","article-title":"Bioassay to detect Ascophyllum nodosum extract-induced cytokinin-like activity in Arabidopsis thaliana","volume":"23","author":"Khan","year":"2011","journal-title":"J. Appl. Phycol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"918","DOI":"10.1007\/s00343-019-8109-x","article-title":"Production of bio-fertilizer from Ascophyllum nodosum and Sargassum muticum (Phaeophyceae)","volume":"37","author":"Silva","year":"2019","journal-title":"J. Oceanol. Limnol."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1007\/s10126-005-0029-4","article-title":"Salt-regulated mannitol metabolism in algae","volume":"7","author":"Iwamoto","year":"2005","journal-title":"Mar. Biotechnol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1093\/aob\/mcq163","article-title":"Genome-wide analysis of thiourea-modulated salinity stress-responsive transcripts in seeds of Brassica juncea: Identification of signalling and effector components of stress tolerance","volume":"106","author":"Srivastava","year":"2010","journal-title":"Ann. Bot."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"234","DOI":"10.1080\/17429145.2012.725480","article-title":"Exogenous application of mannitol and thiourea regulates plant growth and oxidative stress responses in salt-stressed maize (Zea mays L.)","volume":"8","author":"Kaya","year":"2013","journal-title":"J. Plant Interact."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1007\/s00344-008-9068-1","article-title":"An enhancing effect of exogenous mannitol on the antioxidant enzyme activities in roots of wheat under salt stress","volume":"28","author":"Seckin","year":"2009","journal-title":"J. Plant Growth Regul."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/0003-2697(87)90612-9","article-title":"Universal chemical assay for the detection and determination of siderophores","volume":"160","author":"Schwyn","year":"1987","journal-title":"Anal. Biochem."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1111\/j.1574-6968.1999.tb13383.x","article-title":"An efficient microbiological growth medium for screening phosphate solubilizing microorganisms","volume":"170","author":"Nautiyal","year":"1999","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1016\/S0021-9258(18)84756-1","article-title":"The colorimetric determination of phosphorus","volume":"66","author":"Fiske","year":"1925","journal-title":"J. Biol. Chem."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"142","DOI":"10.1111\/j.1399-3054.1948.tb07118.x","article-title":"Production of hydrocyanic acid by bacteria","volume":"1","author":"Lorck","year":"1948","journal-title":"Physiol. Plant."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1016\/j.mimet.2009.08.008","article-title":"Effect of iron and phosphate on bacterial cyanide formation determined by methemoglobin in two-dimensional gradient microcultivations","volume":"79","author":"Furrer","year":"2009","journal-title":"J. Microbiol. Methods."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1007\/s12298-010-0041-7","article-title":"Production of indole acetic acid by Pseudomonas sp.: Effect of coinoculation with Mesorhizobium sp. Cicer on nodulation and plant growth of chickpea (Cicer arietinum)","volume":"17","author":"Malik","year":"2011","journal-title":"Physiol. Mol. Biol. Plants."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1007\/BF02916414","article-title":"Salt-tolerant and thermostable alkaline protease from Bacillus subtilis NCIM No. 64","volume":"38","author":"Kembhavi","year":"1993","journal-title":"Appl. Biochem. Biotechnol."},{"key":"ref_92","doi-asserted-by":"crossref","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(-Delta Delta C(T)) Method","volume":"25","author":"Livak","year":"2001","journal-title":"Methods"}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/12\/6\/1208\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T18:49:33Z","timestamp":1760122173000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/12\/6\/1208"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,3,7]]},"references-count":92,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2023,3]]}},"alternative-id":["plants12061208"],"URL":"https:\/\/doi.org\/10.3390\/plants12061208","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,3,7]]}}}