{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,17]],"date-time":"2026-10-17T02:59:21Z","timestamp":1792205961337,"version":"4.4.0"},"reference-count":117,"publisher":"MDPI AG","issue":"12","license":[{"start":{"date-parts":[[2022,12,16]],"date-time":"2022-12-16T00:00:00Z","timestamp":1671148800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001871","name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["UIDB\/04551\/2020"],"award-info":[{"award-number":["UIDB\/04551\/2020"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["UIDP\/04551\/2020"],"award-info":[{"award-number":["UIDP\/04551\/2020"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["050\/BII\/2022"],"award-info":[{"award-number":["050\/BII\/2022"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["UIDB\/04551\/2020"],"award-info":[{"award-number":["UIDB\/04551\/2020"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["UIDP\/04551\/2020"],"award-info":[{"award-number":["UIDP\/04551\/2020"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["050\/BII\/2022"],"award-info":[{"award-number":["050\/BII\/2022"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biology"],"abstract":"<jats:p>Agricultural production is being affected by increasingly harsh conditions caused by climate change. The vast majority of crops suffer growth and yield declines due to a lack of water or intense heat. Hence, commercial legume crops suffer intense losses of production (20\u201380%). This situation is even more noticeable in plants used as fodder for animals, such as alfalfa and pitch trefoil, since their productivity is linked not only to the number of seeds produced, but also to the vegetative growth of the plant itself. Thus, we decided to study the microbiota associated with their seeds in different locations on the Iberian Peninsula, with the aim of identifying culturable bacteria strains that have adapted to harsh environments and that can be used as biotreatments to improve plant growth and resistance to stress. As potentially inherited microbiota, they may also represent a treatment with medium- and long-term adaptative effects. Hence, isolated strains showed no clear relationship with their geographical sampling location, but had about 50% internal similarity with their model plants. Moreover, out of the 51 strains isolated, about 80% were capable of producing biofilms; around 50% produced mid\/high concentrations of auxins and grew notably in ACC medium; only 15% were characterized as xerotolerant, while more than 75% were able to sporulate; and finally, 65% produced siderophores and more than 40% produced compounds to solubilize phosphates. Thus, Paenibacillus amylolyticus BB B2-A, Paenibacillus xylanexedens MS M1-C, Paenibacillus pabuli BB Oeiras A, Stenotrophomonas maltophilia MS M1-B and Enterobacter hormaechei BB B2-C strains were tested as plant bioinoculants in lentil plants (Lens culinaris Medik.), showing promising results as future treatments to improve plant growth under stressful conditions.<\/jats:p>","DOI":"10.3390\/biology11121838","type":"journal-article","created":{"date-parts":[[2022,12,19]],"date-time":"2022-12-19T05:55:43Z","timestamp":1671429343000},"page":"1838","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":16,"title":["Geographically Disperse, Culturable Seed-Associated Microbiota in Forage Plants of Alfalfa (Medicago sativa L.) and Pitch Clover (Bituminaria bituminosa L.): Characterization of Beneficial Inherited Strains as Plant Stress-Tolerance Enhancers"],"prefix":"10.3390","volume":"11","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5095-7578","authenticated-orcid":false,"given":"Marla","family":"Niza-Costa","sequence":"first","affiliation":[{"name":"iPlantMicro Lab, Instituto de Tecnologia Qu\u00edmica e Biol\u00f3gica (ITQB)-NOVA, Oeiras, 2784-501 Lisboa, Portugal"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ana Sof\u00eda","family":"Rodr\u00edguez-dos Santos","sequence":"additional","affiliation":[{"name":"iPlantMicro Lab, Instituto de Tecnologia Qu\u00edmica e Biol\u00f3gica (ITQB)-NOVA, Oeiras, 2784-501 Lisboa, Portugal"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"In\u00eas","family":"Rebelo-Rom\u00e3o","sequence":"additional","affiliation":[{"name":"iPlantMicro Lab, Instituto de Tecnologia Qu\u00edmica e Biol\u00f3gica (ITQB)-NOVA, Oeiras, 2784-501 Lisboa, Portugal"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mar\u00eda Victoria","family":"Ferrer","sequence":"additional","affiliation":[{"name":"iPlantMicro Lab, Instituto de Tecnologia Qu\u00edmica e Biol\u00f3gica (ITQB)-NOVA, Oeiras, 2784-501 Lisboa, Portugal"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cristina","family":"Sequero L\u00f3pez","sequence":"additional","affiliation":[{"name":"GeoBioTec, Department of Earth Sciences, NOVA School of Sciences and Technology, Universidade NOVA de Lisboa (Campus de Caparica), 1070-312 Caparica, Portugal"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4524-7384","authenticated-orcid":false,"given":"Juan Ignacio","family":"V\u00edlchez","sequence":"additional","affiliation":[{"name":"iPlantMicro Lab, Instituto de Tecnologia Qu\u00edmica e Biol\u00f3gica (ITQB)-NOVA, Oeiras, 2784-501 Lisboa, Portugal"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2022,12,16]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"4991","DOI":"10.1038\/s41467-020-18725-w","article-title":"Crop switching reduces agricultural losses from climate change in the United States by half under RCP 8.5","volume":"11","author":"Rising","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Sulser, T., Wiebe, K.D., Dunston, S., Cenacchi, N., Nin-Pratt, A., Mason-D\u2019Croz, D., Robertson, R.D., Willenbockel, D., and Rosegrant, M.W. (2021). Climate Change and Hunger: Estimating Costs of Adaptation in the Agrifood System, International Food Policy Research Institute.","DOI":"10.2499\/9780896294165"},{"key":"ref_3","unstructured":"Nelson, G.C., Rosegrant, M.W., Koo, J., Robertson, R.D., Sulser, T., Zhu, T., Ringler, C., Msangi, S., Palazzo, A., and Batka, M. (2009). Climate Change: Impact on Agriculture and Costs of Adaptation, International Food Policy Research Institute."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"972","DOI":"10.1038\/s41558-018-0299-2","article-title":"Climate change and interconnected risks to sustainable development in the Mediterranean","volume":"8","author":"Cramer","year":"2018","journal-title":"Nat. Clim. Chang."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Lange, M.A. (2019). Impacts of climate change on the Eastern Mediterranean and the Middle East and North Africa Region and the water\u2013energy nexus. Atmosphere, 10.","DOI":"10.3390\/atmos10080455"},{"key":"ref_6","first-page":"100032","article-title":"A review on drought stress in plants: Implications, mitigation and the role of plant growth promoting rhizobacteria","volume":"5","author":"Ahluwalia","year":"2021","journal-title":"Resour. Environ. Sustain."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"618092","DOI":"10.3389\/fsufs.2021.618092","article-title":"Climate change and salinity effects on crops and chemical communication between plants and plant growth-promoting microorganisms under stress","volume":"5","author":"Ullah","year":"2021","journal-title":"Front. Sustain. Food Syst."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"694121","DOI":"10.3389\/fsufs.2021.694121","article-title":"Legumes as a cornerstone of the transition toward more sustainable agri-food systems and diets in Europe","volume":"5","author":"Ferreira","year":"2021","journal-title":"Front. Sustain. Food Syst."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"100520","DOI":"10.1016\/j.gfs.2021.100520","article-title":"Legumes as a sustainable source of protein in human diets","volume":"28","author":"Semba","year":"2021","journal-title":"Glob. Food Secur."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1016\/j.livsci.2006.09.029","article-title":"Nutritive value of forage species grown in the warm temperate climate of Australia for dairy cows: Grasses and legumes","volume":"107","author":"Fulkerson","year":"2007","journal-title":"Livest. Sci."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Daryanto, S., Wang, L., and Jacinthe, P.A. (2015). Global synthesis of drought effects on food legume production. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0127401"},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Khatun, M., Sarkar, S., Era, F.M., Islam, A.K.M.M., Anwar, M.P., Fahad, S., Datta, R., and Islam, A.K.M.A. (2021). Drought stress in grain legumes: Effects, tolerance mechanisms and management. Agronomy, 11.","DOI":"10.3390\/agronomy11122374"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1015","DOI":"10.3389\/fpls.2016.01015","article-title":"Drought stress responses in soybean roots and nodules","volume":"7","author":"Kunert","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Nadeem, M., Li, J., Yahya, M., Sher, A., Ma, C., Wang, X., and Qiu, L. (2019). Research progress and perspective on drought stress in legumes: A review. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20102541"},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Singh, S., Kudapa, H., Garg, V., and Varshney, R.K. (2021). Comprehensive analysis and identification of drought-responsive candidate NAC genes in three semi-arid tropics (SAT) legume crops. BMC Genom., 22.","DOI":"10.1186\/s12864-021-07602-5"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1534","DOI":"10.1111\/1365-2745.12940","article-title":"Drought-adapted plants dramatically downregulate dinitrogen fixation: Evidences from Mediterranean legume shrubs","volume":"106","author":"Dovrat","year":"2018","journal-title":"J. Ecol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"20160034","DOI":"10.1098\/rstb.2016.0034","article-title":"Domestication impacts on plant-herbivore interactions: A meta-analysis","volume":"372","author":"Whitehead","year":"2017","journal-title":"Philos. Trans. R. Soc. Lond. B Biol. Sci."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1586","DOI":"10.1111\/nph.17452","article-title":"Intentional and unintentional selection during plant domestication: Herbivore damage, plant defensive traits and nutritional quality of fruit and seed crops","volume":"231","author":"Fernandez","year":"2021","journal-title":"New Phytol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1573","DOI":"10.1093\/pcp\/pcac083","article-title":"Unintended consequences of plant domestication","volume":"63","author":"Singh","year":"2022","journal-title":"Plant Cell Physiol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"936","DOI":"10.1038\/s42003-021-02467-6","article-title":"The effect of plant domestication on host control of the microbiota","volume":"4","author":"Soldan","year":"2021","journal-title":"Commun. Biol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1654","DOI":"10.1093\/pcp\/pcac108","article-title":"Effects of domestication on plant-microbiome interactions","volume":"63","author":"Gutierrez","year":"2022","journal-title":"Plant Cell Physiol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"806915","DOI":"10.3389\/fpls.2021.806915","article-title":"Domestication impacts the wheat-associated microbiota and the rhizosphere colonization by seed- and soil-originated microbiomes, across different fields","volume":"12","author":"Abdullaeva","year":"2022","journal-title":"Front. Plant Sci."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"635","DOI":"10.1007\/s11103-015-0337-7","article-title":"Impact of plant domestication on rhizosphere microbiome assembly and functions","volume":"90","author":"Mendes","year":"2016","journal-title":"Plant Mol. Biol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"688533","DOI":"10.3389\/fpls.2021.688533","article-title":"Tomato cultivars with variable tolerances to water deficit differentially modulate the composition and interaction patterns of their rhizosphere microbial communities","volume":"12","author":"Gaete","year":"2021","journal-title":"Front. Plant Sci."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1753","DOI":"10.1111\/1365-2745.13157","article-title":"Domesticated tomatoes are more vulnerable to negative plant\u2013soil feedbacks than their wild relatives","volume":"107","author":"Carrillo","year":"2019","journal-title":"J. Ecol."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Smulders, L., Ben\u00edtez, E., Moreno, B., L\u00f3pez-Garc\u00eda, \u00c1., Pozo, M.J., Ferrero, V., de la Pe\u00f1a, E., and Alcal\u00e1 Herrera, R. (2021). Tomato domestication affects potential functional molecular pathways of root-associated soil bacteria. Plants, 10.","DOI":"10.3390\/plants10091942"},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Riley, A.B., Grillo, M.A., Epstein, B., Tiffin, P., and Heath, K.D. (2022). Discordant population structure among rhizobium divided genomes and their legume hosts. Mol. Ecol.","DOI":"10.22541\/au.166004242.26292927\/v1"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"2124","DOI":"10.3389\/fpls.2017.02124","article-title":"Evolution of the crop rhizosphere: Impact of domestication on root exudates in tetraploid wheat (Triticum turgidum L.)","volume":"8","author":"Iannucci","year":"2017","journal-title":"Front. Plant Sci."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"126106","DOI":"10.1016\/j.syapm.2020.126106","article-title":"Plant microbiota modified by plant domestication","volume":"43","year":"2020","journal-title":"Syst. Appl. Microbiol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"234","DOI":"10.1186\/s40168-021-01177-9","article-title":"Leave no stone unturned: Individually adapted xerotolerant Thaumarchaeota sheltered below the boulders of the Atacama Desert hyperarid core","volume":"9","author":"Hwang","year":"2021","journal-title":"Microbiome"},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"L\u00f3pez, M.C., Gal\u00e1n, B., Carmona, M., Navarro Llorens, J.M., Peret\u00f3, J., Porcar, M., Getino, L., Olivera, E.R., Luengo, J.M., and Castro, L. (2021). Xerotolerance: A new property in Exiguobacterium genus. Microorganisms, 9.","DOI":"10.3390\/microorganisms9122455"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"743512","DOI":"10.3389\/fmicb.2021.743512","article-title":"The role of plant-associated bacteria, fungi, and viruses in drought stress mitigation","volume":"12","author":"Poudel","year":"2021","journal-title":"Front. Microbiol."},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Gamalero, E., and Glick, B.R. (2022). Recent advances in bacterial amelioration of plant drought and salt stress. Biology, 11.","DOI":"10.3390\/biology11030437"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"822","DOI":"10.1007\/s00248-020-01543-4","article-title":"Unveiling the microbiota diversity of the xerophyte Argania spinosa L. skeels root system and residuesphere","volume":"80","author":"Mapelli","year":"2020","journal-title":"Microb. Ecol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1038\/nrmicro.2017.16","article-title":"Xerotolerant bacteria: Surviving through a dry spell","volume":"15","author":"Lebre","year":"2017","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"553223","DOI":"10.3389\/fmicb.2020.553223","article-title":"Beneficial plant-associated microorganisms from semiarid regions and seasonally dry environments: A review","volume":"11","author":"Bonatelli","year":"2021","journal-title":"Front. Microbiol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"100173","DOI":"10.1016\/j.cpb.2020.100173","article-title":"Reclamation of arid and semi-arid soils: The role of plant growth-promoting archaea and bacteria","volume":"25","author":"Ayangbenro","year":"2021","journal-title":"Curr. Plant Biol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1007\/s11104-017-3289-7","article-title":"The seed microbiome: Origins, interactions, and impacts","volume":"422","author":"Nelson","year":"2018","journal-title":"Plant Soil"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"559728","DOI":"10.3389\/fmicb.2020.559728","article-title":"Dynamic changes in the microbiome of rice during shoot and root growth derived from seeds","volume":"11","author":"Wang","year":"2020","journal-title":"Front. Microbiol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"2152","DOI":"10.1111\/nph.16284","article-title":"Analysis of leaf microbiome composition of near-isogenic maize lines differing in broad-spectrum disease resistance","volume":"225","author":"Wagner","year":"2020","journal-title":"New Phytol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"2199","DOI":"10.1111\/1462-2920.15392","article-title":"Experimental evidence of microbial inheritance in plants and transmission routes from seed to phyllosphere and root","volume":"23","author":"Abdelfattah","year":"2021","journal-title":"Environ. Microbiol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"40","DOI":"10.1094\/PBIOMES-04-19-0023-R","article-title":"Heritability and functional importance of the Setaria viridis bacterial seed microbiome","volume":"4","author":"Antonielli","year":"2020","journal-title":"Phytobiomes J."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"11","DOI":"10.3389\/fmicb.2017.00011","article-title":"A new approach to modify plant microbiomes and traits by introducing beneficial bacteria at flowering into progeny seeds","volume":"8","author":"Mitter","year":"2017","journal-title":"Front. Microbiol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"11916","DOI":"10.1038\/s41598-021-91351-8","article-title":"Profiling, isolation and characterisation of beneficial microbes from the seed microbiomes of drought tolerant wheat","volume":"11","author":"Hone","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"fiw112","DOI":"10.1093\/femsec\/fiw112","article-title":"Seed biopriming with plant growth promoting rhizobacteria: A review","volume":"92","author":"Mahmood","year":"2016","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"24","DOI":"10.3389\/fpls.2018.00024","article-title":"What is there in seeds? Vertically transmitted endophytic resources for sustainable improvement in plant growth","volume":"9","author":"Shahzad","year":"2018","journal-title":"Front. Plant Sci."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1812","DOI":"10.1111\/1462-2920.15240","article-title":"The plant endosphere world\u2014Bacterial life within plants","volume":"23","author":"Compant","year":"2021","journal-title":"Environ. Microbiol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1448","DOI":"10.1111\/nph.18037","article-title":"Seed microbiota revealed by a large-scale meta-analysis including 50 plant species","volume":"234","author":"Simonin","year":"2022","journal-title":"New Phytol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"e00600","DOI":"10.1016\/j.btre.2021.e00600","article-title":"Whole genome sequence of Pantoea agglomerans ANP8, a salinity and drought stress-resistant bacterium isolated from alfalfa (Medicago sativa L.) root nodules","volume":"29","author":"Noori","year":"2021","journal-title":"Biotechnol. Rep."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1016\/j.ecoenv.2018.06.092","article-title":"Mining alfalfa (Medicago sativa L.) nodules for salinity tolerant non-rhizobial bacteria to improve growth of alfalfa under salinity stress","volume":"162","author":"Noori","year":"2018","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Issah, G., Schoenau, J.J., Lardner, H.A., and Knight, J.D. (2020). Nitrogen fixation and resource partitioning in alfalfa (Medicago sativa L.), cicer milkvetch (Astragalus cicer L.) and sainfoin (Onobrychis viciifolia Scop.) using 15N enrichment under controlled environment conditions. Agronomy, 10.","DOI":"10.3390\/agronomy10091438"},{"key":"ref_52","first-page":"2206","article-title":"Physiological responses of Bituminaria bituminosa to drought and heavy metals","volume":"168","year":"2012","journal-title":"J. Plant Physiol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1034","DOI":"10.1071\/CP12216","article-title":"Drought resistance at the seedling stage in the promising fodder plant tedera (Bituminaria bituminosa var. albomarginata)","volume":"63","author":"Foster","year":"2012","journal-title":"Crop Pasture Sci."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"357","DOI":"10.4067\/S0718-58392011000300003","article-title":"Evaluation of perennial forage legumes and herbs in six Mediterranean environments","volume":"71","author":"Real","year":"2011","journal-title":"Chil. J. Agric. Res."},{"key":"ref_55","unstructured":"do Ros\u00e1rio, L. (2004). Indicadores de Desertifica\u00e7ao para Portugal Continental."},{"key":"ref_56","unstructured":"Enne, C., Zanolla, C., and Peter, D. (2000). Managing desertification in a national context. Desertification in Europe: Mitigation Strategies, Land Use Planning, European Communities."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Page, A.L. (1983). Soil pH and lime requirement. Methods of Soil Analysis, Part 2: Chemical and Microbiological Properties, American Society of Agronomy, Inc.","DOI":"10.2134\/agronmonogr9.2.2ed"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"116","DOI":"10.1016\/j.chnaes.2013.06.004","article-title":"Determination and analysis of phreatic water evaporation in extra-arid dune region","volume":"34","author":"Li","year":"2014","journal-title":"Acta Ecol. Sin."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1817","DOI":"10.1016\/j.rse.2010.03.009","article-title":"Assessment and monitoring of land condition in the Iberian Peninsula, 1989\u20132000","volume":"114","author":"Puigdefabregas","year":"2010","journal-title":"Remote Sens. Environ."},{"key":"ref_60","first-page":"e3976","article-title":"Measurements of root colonized bacteria species","volume":"11","author":"Yang","year":"2021","journal-title":"Bio-Protocol"},{"key":"ref_61","unstructured":"Breakwell, D., Woolverton, C., MacDonald, B., Smith, K., and Robison, R. (2007, January 29). Colony morphology protocol. Proceedings of the 14th American Society for Microbiology Conference for Undergraduate Educators (ASMCUE), Buffalo, NY, USA."},{"key":"ref_62","first-page":"117","article-title":"Heat treatment of bacteria: A simple method of DNA extraction for molecular techniques","volume":"41","author":"Dashti","year":"2009","journal-title":"Kuwait Med. J."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"W293","DOI":"10.1093\/nar\/gkab301","article-title":"Interactive Tree Of Life (iTOL) v5: An online tool for phylogenetic tree display and annotation","volume":"49","author":"Letunic","year":"2021","journal-title":"Nucleic Acids Res."},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Heberle, H., Meirelles, G.V., da Silva, F.R., Telles, G.P., and Minghim, R. (2015). InteractiVenn: A web-based tool for the analysis of sets through Venn diagrams. BMC Bioinform., 16.","DOI":"10.1186\/s12859-015-0611-3"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1111\/j.1574-6976.2010.00249.x","article-title":"Comparative genomics and functional analysis of niche-specific adaptation in Pseudomonas putida","volume":"35","author":"Wu","year":"2011","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"5254","DOI":"10.1128\/AEM.00855-10","article-title":"Rapid method for isolation of desiccation-tolerant strains and xeroprotectants","volume":"76","author":"Barba","year":"2010","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"631","DOI":"10.1007\/978-1-4939-0473-0_48","article-title":"Biofilm formation in the 96-well microtiter plate","volume":"1149","author":"Coffey","year":"2014","journal-title":"Methods Mol. Biol."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1128\/jmbe.v12i1.249","article-title":"Use of blue agar CAS assay for siderophore detection","volume":"12","author":"Louden","year":"2011","journal-title":"J. Microbiol. Biol. Educ."},{"key":"ref_69","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_70","doi-asserted-by":"crossref","first-page":"1368","DOI":"10.1104\/pp.18.00217","article-title":"EZ-Root-VIS: A Software pipeline for the rapid analysis and visual reconstruction of root system architecture","volume":"177","author":"Shahzad","year":"2018","journal-title":"Plant Physiol."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Johnston-Monje, D., and Raizada, M.N. (2011). Conservation and diversity of seed associated endophytes in Zea across boundaries of evolution, ethnography and ecology. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0020396"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1007\/s10658-020-01998-8","article-title":"Kosakonia cowanii as the new bacterial pathogen affecting soybean (Glycine max Willd.)","volume":"157","author":"Krawczyk","year":"2020","journal-title":"Eur. J. Plant Pathol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1007\/0-387-30746-X_15","article-title":"Erwinia and related genera","volume":"Volume 6","author":"Kado","year":"2006","journal-title":"The Prokaryotes"},{"key":"ref_74","first-page":"37","article-title":"Wheat seeds harbour bacterial endophytes with potential as plant growth promoters and biocontrol agents of Fusarium graminearum","volume":"186\u2013187","author":"Grossi","year":"2016","journal-title":"Microbiol. Res."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1186\/s40793-021-00389-8","article-title":"The treasure inside barley seeds: Microbial diversity and plant beneficial bacteria","volume":"16","author":"Bziuk","year":"2021","journal-title":"Environ. Microbiome"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"5659","DOI":"10.1111\/1462-2920.15450","article-title":"Paenibacillus polymyxa, a Jack of all trades","volume":"23","author":"Langendries","year":"2021","journal-title":"Environ. Microbiol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1186\/s12934-016-0603-7","article-title":"Current knowledge and perspectives of Paenibacillus: A review","volume":"15","author":"Grady","year":"2016","journal-title":"Microb. Cell Factories"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"438","DOI":"10.1016\/j.bcab.2018.04.012","article-title":"Stress tolerance and plant growth promotion potential of Enterobacter ludwigii PS1 isolated from seabuckthorn rhizosphere","volume":"14","author":"Dolkar","year":"2018","journal-title":"Biocatal. Agric. Biotechnol."},{"key":"ref_79","doi-asserted-by":"crossref","unstructured":"Tsotetsi, T., Nephali, L., Malebe, M., and Tugizimana, F. (2022). Bacillus for plant growth promotion and stress resilience: What have we learned?. Plants, 11.","DOI":"10.3390\/plants11192482"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"568289","DOI":"10.3389\/fmicb.2020.568289","article-title":"Halotolerant PGPR Stenotrophomonas maltophilia BJ01 induces salt tolerance by modulating physiology and biochemical activities of Arachis hypogaea","volume":"11","author":"Alexander","year":"2020","journal-title":"Front. Microbiol."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1945","DOI":"10.3389\/fmicb.2017.01945","article-title":"The PGPR Stenotrophomonas maltophilia SBP-9 Augments resistance against biotic and abiotic stress in wheat plants","volume":"8","author":"Singh","year":"2017","journal-title":"Front. Microbiol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"7292","DOI":"10.1128\/AEM.71.11.7292-7300.2005","article-title":"Paenibacillus polymyxa invades plant roots and forms biofilms","volume":"71","author":"Timmusk","year":"2005","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"951","DOI":"10.1094\/MPMI.1999.12.11.951","article-title":"The plant-growth-promoting rhizobacterium Paenibacillus polymyxa induces changes in Arabidopsis thaliana gene expression: A possible connection between biotic and abiotic stress responses","volume":"12","author":"Timmusk","year":"1999","journal-title":"Mol. Plant-Microbe Interact."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1016\/j.plaphy.2018.06.001","article-title":"Paenibacillus pabuli strain P7S promotes plant growth and induces anthocyanin accumulation in Arabidopsis thaliana","volume":"129","author":"Trinh","year":"2018","journal-title":"Plant Physiol. Biochem."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"880125","DOI":"10.3389\/fpls.2022.880125","article-title":"Role of rhizospheric Bacillus megaterium HGS7 in maintaining mulberry growth under extremely abiotic stress in hydro-fluctuation belt of three gorges reservoir","volume":"13","author":"Ou","year":"2022","journal-title":"Front. Plant Sci."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"539359","DOI":"10.3389\/fmicb.2020.539359","article-title":"Metabolites secreted by a plant-growth-promoting Pantoea agglomerans strain improved rooting of Pyrus communis L. cv dar gazi cuttings","volume":"11","author":"Luziatelli","year":"2020","journal-title":"Front. Microbiol."},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Khan, M.A., Asaf, S., Khan, A.L., Jan, R., Kang, S.-M., Kim, K.-M., and Lee, I.-J. (2020). Thermotolerance effect of plant growth-promoting Bacillus cereus SA1 on soybean during heat stress. BMC Microbiol., 20.","DOI":"10.1186\/s12866-020-01822-7"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1016\/j.micres.2019.03.001","article-title":"Bacterial seed endophyte community of annual plants modulated by plant photosynthetic pathways","volume":"223\u2013225","author":"Girsowicz","year":"2019","journal-title":"Microbiol. Res."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"46","DOI":"10.7845\/kjm.2013.004","article-title":"Growth promotion of tomato plant under drought conditions by treatment of rhizobacteria producing ACC deaminase and phytohormones","volume":"49","author":"Seo","year":"2013","journal-title":"Korean J. Microbiol."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"1208","DOI":"10.1007\/s00284-021-02368-1","article-title":"Enterobacter hormaechei as plant growth-promoting bacteria for improvement in Lycopersicum esculentum","volume":"78","author":"Ranawat","year":"2021","journal-title":"Curr. Microbiol."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"730","DOI":"10.1007\/s40333-020-0019-4","article-title":"Endophytic bacteria associated with halophyte Seidlitzia rosmarinus Ehrenb. ex Boiss. from saline soil of Uzbekistan and their plant beneficial traits","volume":"12","author":"Shurigin","year":"2020","journal-title":"J. Arid Land"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"3732","DOI":"10.1128\/AEM.00049-12","article-title":"Psychrotolerant Paenibacillus tundrae isolates from barley grains produce new cereulide-like depsipeptides (Paenilide and Homopaenilide) that are highly toxic to mammalian cells","volume":"78","author":"Rasimus","year":"2012","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_93","doi-asserted-by":"crossref","unstructured":"Vargas, P., Bosmans, L., Van Kerckhove, S., Van Calenberge, B., Raaijmakers, J.M., Lievens, B., and Rediers, H. (2021). Optimizing biocontrol activity of Paenibacillus xylanexedens for management of hairy root disease in tomato grown in hydroponic greenhouses. Agronomy, 11.","DOI":"10.3390\/agronomy11050817"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"1708","DOI":"10.1099\/ijs.0.004572-0","article-title":"Paenibacillus tundrae sp. nov. and Paenibacillus xylanexedens sp. nov., psychrotolerant, xylan-degrading bacteria from Alaskan tundra","volume":"59","author":"Nelson","year":"2009","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"241","DOI":"10.3389\/fmicb.2015.00241","article-title":"Friends or foes: Can we make a distinction between beneficial and harmful strains of the Stenotrophomonas maltophilia complex?","volume":"6","author":"Berg","year":"2015","journal-title":"Front. Microbiol."},{"key":"ref_96","doi-asserted-by":"crossref","unstructured":"Cherif-Silini, H., Thissera, B., Bouket, A.C., Saadaoui, N., Silini, A., Eshelli, M., Alenezi, F.N., Vallat, A., Luptakova, L., and Yahiaoui, B. (2019). Durum wheat stress tolerance induced by endophyte Pantoea agglomerans with genes contributing to plant functions and secondary metabolite arsenal. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20163989"},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"1004833","DOI":"10.3389\/fpls.2022.1004833","article-title":"Potential of mineral-solubilizing bacteria for physiology and growth promotion of Chenopodium quinoa Willd","volume":"13","author":"Rafique","year":"2022","journal-title":"Front Plant Sci"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"622","DOI":"10.1111\/mpp.12918","article-title":"Analogous wheat root rhizosphere microbial successions in field and greenhouse trials in the presence of biocontrol agents Paenibacillus peoriae SP9 and Streptomyces fulvissimus FU14","volume":"21","author":"Araujo","year":"2020","journal-title":"Mol. Plant Pathol."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"15545","DOI":"10.1038\/s41598-021-94820-2","article-title":"Transcriptomics differentiate two novel bioactive strains of Paenibacillus sp. isolated from the perennial ryegrass seed microbiome","volume":"11","author":"Li","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"e00715-20","DOI":"10.1128\/MRA.00715-20","article-title":"Draft genome sequences of seven strains of Paenibacillus spp. (Phylum Firmicutes) inhabiting the seeds of Cucumis melo L. (Cantaloupe) and exhibiting plant probiotic traits","volume":"9","author":"Khalaf","year":"2020","journal-title":"Microbiol. Resour. Announc."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"542","DOI":"10.1111\/nph.12693","article-title":"Simultaneous profiling of seed-associated bacteria and fungi reveals antagonistic interactions between microorganisms within a shared epiphytic microbiome on Triticum and Brassica seeds","volume":"202","author":"Links","year":"2014","journal-title":"New Phytol."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"E1621","DOI":"10.1073\/pnas.1218984110","article-title":"Bacillus subtilis biofilm induction by plant polysaccharides","volume":"110","author":"Beauregard","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1111\/jam.14609","article-title":"Microbial biofilms in nature: Unlocking their potential for agricultural applications","volume":"129","author":"Pandit","year":"2020","journal-title":"J. Appl. Microbiol."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1186\/s13568-022-01432-8","article-title":"Biofilm forming rhizobacteria affect the physiological and biochemical responses of wheat to drought","volume":"12","author":"Karimi","year":"2022","journal-title":"AMB Express"},{"key":"ref_105","doi-asserted-by":"crossref","unstructured":"Yin, W., Wang, Y., Liu, L., and He, J. (2019). Biofilms: The microbial \u201cprotective clothing\u201d in extreme environments. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20143423"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"e1071004","DOI":"10.1080\/15592324.2015.1071004","article-title":"Synergistic effect of Pseudomonas putida and Bacillus amyloliquefaciens ameliorates drought stress in chickpea (Cicer arietinum L.)","volume":"11","author":"Kumar","year":"2016","journal-title":"Plant Signal. Behav."},{"key":"ref_107","doi-asserted-by":"crossref","unstructured":"Wang, D.C., Jiang, C.H., Zhang, L.N., Chen, L., Zhang, X.Y., and Guo, J.H. (2019). Biofilms positively contribute to Bacillus amyloliquefaciens 54-induced drought tolerance in tomato plants. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20246271"},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"055082","DOI":"10.1088\/1742-6596\/1402\/5\/055082","article-title":"Screening of the drought tolerant phosphate solubilizing bacteria in dissolving P-inorganic","volume":"1402","author":"Susilowati","year":"2019","journal-title":"J. Phys. Conf. Ser."},{"key":"ref_109","doi-asserted-by":"crossref","unstructured":"Kang, S.-M., Khan, M.-A., Hamayun, M., Kim, L.-R., Kwon, E.-H., Kang, Y.-S., Kim, K.-Y., Park, J.-J., and Lee, I.-J. (2021). Phosphate-solubilizing Enterobacter ludwigii AFFR02 and Bacillus megaterium Mj1212 rescues alfalfa\u2019s growth under post-drought stress. Agriculture, 11.","DOI":"10.3390\/agriculture11060485"},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"898979","DOI":"10.3389\/fmicb.2022.898979","article-title":"Mechanistic insights and potential use of siderophores producing microbes in rhizosphere for mitigation of stress in plants grown in degraded land","volume":"13","author":"Singh","year":"2022","journal-title":"Front. Microbiol."},{"key":"ref_111","doi-asserted-by":"crossref","unstructured":"Breitkreuz, C., Reitz, T., Schulz, E., and Tarkka, M.T. (2021). Drought and plant community composition affect the metabolic and genotypic diversity of pseudomonas strains in grassland soils. Microorganisms, 9.","DOI":"10.3390\/microorganisms9081677"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"188","DOI":"10.1007\/s00248-011-9883-y","article-title":"Endophytes of grapevine flowers, berries, and seeds: Identification of cultivable bacteria, comparison with other plant parts, and visualization of niches of colonization","volume":"62","author":"Compant","year":"2011","journal-title":"Microb. Ecol."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"3264","DOI":"10.1111\/1462-2920.15926","article-title":"Back to our roots: Exploring the role of root morphology as a mediator of beneficial plant\u2013microbe interactions","volume":"24","author":"Herms","year":"2022","journal-title":"Environ. Microbiol."},{"key":"ref_114","doi-asserted-by":"crossref","unstructured":"Saberi Riseh, R., Ebrahimi-Zarandi, M., Gholizadeh Vazvani, M., and Skorik, Y.A. (2021). Reducing drought stress in plants by encapsulating plant growth-promoting bacteria with polysaccharides. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms222312979"},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1038\/nature03184","article-title":"The PIN auxin efflux facilitator network controls growth and patterning in Arabidopsis roots","volume":"433","author":"Blilou","year":"2005","journal-title":"Nature"},{"key":"ref_116","doi-asserted-by":"crossref","unstructured":"Camaille, M., Fabre, N., Cl\u00e9ment, C., and Ait Barka, E. (2021). Advances in wheat physiology in response to drought and the role of plant growth promoting rhizobacteria to trigger drought tolerance. Microorganisms, 9.","DOI":"10.3390\/microorganisms9040687"},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1186\/s42269-021-00546-6","article-title":"Enhancement of drought tolerance in Triticum aestivum L. seedlings using Azospirillum brasilense NO40 and Stenotrophomonas maltophilia B11","volume":"45","author":"Kasim","year":"2021","journal-title":"Bull. Natl. Res. Cent."}],"container-title":["Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2079-7737\/11\/12\/1838\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T01:42:56Z","timestamp":1760146976000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2079-7737\/11\/12\/1838"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,12,16]]},"references-count":117,"journal-issue":{"issue":"12","published-online":{"date-parts":[[2022,12]]}},"alternative-id":["biology11121838"],"URL":"https:\/\/doi.org\/10.3390\/biology11121838","relation":{},"ISSN":["2079-7737"],"issn-type":[{"value":"2079-7737","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,12,16]]}}}