{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T15:57:34Z","timestamp":1787932654430,"version":"build-2784847793"},"reference-count":86,"publisher":"MDPI AG","issue":"3","license":[{"start":{"date-parts":[[2026,3,12]],"date-time":"2026-03-12T00:00:00Z","timestamp":1773273600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Deanship of Scientific Research at Imam Mohammad Ibn Saud Islamic University","award":["IMSIU-DDRSP2601"],"award-info":[{"award-number":["IMSIU-DDRSP2601"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Microorganisms"],"abstract":"<jats:p>Environmental pollution caused by persistent chemical compounds, particularly heavy metals, poses a significant global challenge. Current strategies focus on eco-friendly and sustainable approaches, such as the application of microorganisms, to mitigate this issue. In this study, four strains of Bacillus and Pseudomonas were phylogenetically identified and assessed for their resistance to three heavy metals: copper (Cu), chromium (Cr), and cadmium (Cd) up to 500 \u00b5g\/mL. Various tolerance mechanisms related to heavy metal resistance were elucidated, including salinity tolerance, antibiotic resistance, production of exopolysaccharides (EPS), and biosurfactant synthesis. The antifungal activities of these strains were evaluated against the fungal isolates Fusarium oxysporum fs. phaseoli (Fop) and Stemphylium botryosum (St-bt) using dual culture assays. Phylogenetic analysis revealed that three strains belong to the genus Bacillus, while one strain is classified under Pseudomonas. Additionally, these strains exhibited diverse mechanisms for heavy metal tolerance, including salinity tolerance (up to 600 mM), multi-antibiotic resistance (to imipenem, ampicillin, and sodium fusidate), and the production of viscous, slimy colonies indicative of EPS synthesis. Biosurfactant production led to a significant reduction in surface tension, ranging from 10.51 \u00b1 3.87% to 82.89 \u00b1 5.01%. The antifungal assays demonstrated that the strains effectively inhibited the mycelial growth of the fungal isolates, with inhibition percentages varying from 0% to 83.34 \u00b1 2.22%. The strains characterized in this study exhibit considerable potential for application in the bioremediation of metal-contaminated soils and as biocontrol agents.<\/jats:p>","DOI":"10.3390\/microorganisms14030644","type":"journal-article","created":{"date-parts":[[2026,3,12]],"date-time":"2026-03-12T13:12:24Z","timestamp":1773321144000},"page":"644","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Resistance Mechanisms of Rhizospheric Bacillus and Pseudomonas Strains Against Heavy Metal Contamination (Cu, Cr and Cd) and Their Antifungal Properties"],"prefix":"10.3390","volume":"14","author":[{"given":"Slimane","family":"Mokrani","sequence":"first","affiliation":[{"name":"Laboratory of Research on Biological Systems and Geomantic (L.R.S.B.G.), Department of Biology, Mustapha Stambouli University, P.O. Box 305, Mascara 29000, Algeria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-3128-4478","authenticated-orcid":false,"given":"Zahira","family":"Benouguef","sequence":"additional","affiliation":[{"name":"Laboratory of Research on Biological Systems and Geomantic (L.R.S.B.G.), Department of Biology, Mustapha Stambouli University, P.O. Box 305, Mascara 29000, Algeria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Karim","family":"Houali","sequence":"additional","affiliation":[{"name":"Laboratory of Analytical Biochemistry and Biotechnology, Department of Microbiology and Biochemistry, Mouloud Mammeri University, Tizi Ouzou 15000, Algeria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Leila","family":"Bensidhoum","sequence":"additional","affiliation":[{"name":"Laboratoire de Maitrise Des Energies Renouvelables, Universit\u00e9 de Bejaia, Bejaia 06000, Algeria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-1040-1530","authenticated-orcid":false,"given":"Assia","family":"Derguini","sequence":"additional","affiliation":[{"name":"Microbial Ecology Laboratory, Department of Microbiology, Abderrahmane Mira University, Beja\u00efa 06000, Algeria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6382-8807","authenticated-orcid":false,"given":"Nasir A.","family":"Ibrahim","sequence":"additional","affiliation":[{"name":"Department of Biology, College of Science, Imam Mohammad Ibn Saud Islamic University, Riyadh 11623, Saudi Arabia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8814-5957","authenticated-orcid":false,"given":"Nosiba S.","family":"Basher","sequence":"additional","affiliation":[{"name":"Department of Biology, College of Science, Imam Mohammad Ibn Saud Islamic University, Riyadh 11623, Saudi Arabia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8607-0522","authenticated-orcid":false,"given":"El-hafid","family":"Nabti","sequence":"additional","affiliation":[{"name":"Laboratoire de Maitrise Des Energies Renouvelables, Universit\u00e9 de Bejaia, Bejaia 06000, Algeria"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,3,12]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1007\/s11270-024-07702-4","article-title":"Water Quality, Air Pollution, and Climate Change: Investigating the Environmental Impacts of Industrialization and Urbanization","volume":"236","author":"Saxena","year":"2025","journal-title":"Water Air Soil Pollut."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1093\/bmb\/ldg019","article-title":"Environmental pollution and the global burden of disease","volume":"68","author":"Briggs","year":"2003","journal-title":"Br. Med. Bull."},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Rathod, S.V. (2025). Environmental pollution: Threats and challenges for management. Eco-Restoration of Polluted Environment, CRP Press.","DOI":"10.1201\/9781003423393-1"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1016\/bs.aiip.2023.01.005","article-title":"Heavy metal pollutants: The hidden pervasive threat to honey bees and other pollinators","volume":"64","author":"Monchanin","year":"2023","journal-title":"Harrison. Adv. Insect Physiol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1038\/333134a0","article-title":"Quantitative assessment of worldwide contamination of air, water and soils by trace metals","volume":"333","author":"Nriagu","year":"1988","journal-title":"Nature"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"288","DOI":"10.3846\/16486897.2015.1068781","article-title":"Removal of heavy metals using bacterial bio-flocculants of Bacillus sp. and Pseudomonas sp.","volume":"23","author":"Azzam","year":"2015","journal-title":"J. Environ. Eng. Landsc. Manag."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Verma, D.K., Verma, C., and Mahish, P.K. (2023). Physical, chemical, and biological methods of heavy metal management. Heavy Metals in the Environment: Management Strategies for Global Pollution, ACS Publications.","DOI":"10.1021\/bk-2023-1456"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"138861","DOI":"10.1016\/j.chemosphere.2023.138861","article-title":"Biological and green remediation of heavy metal contaminated water and soils: A state-of-the-art review","volume":"332","author":"Sarker","year":"2023","journal-title":"Chemosphere"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1016\/j.jenvman.2016.02.047","article-title":"Beneficial role of bacterial endophytes in heavy metal phytoremediation","volume":"174","author":"Ma","year":"2016","journal-title":"J. Environ. Manag."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1080\/1040841X.2021.1970512","article-title":"Bacterial survival strategies and responses under heavy metal stress: A comprehensive overview","volume":"48","author":"Pal","year":"2022","journal-title":"Crit. Rev. Microbiol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"e2411073","DOI":"10.4014\/jmb.2411.11073","article-title":"Bacterial Heavy Metal Resistance in Contaminated Soil","volume":"35","author":"Iimaa","year":"2025","journal-title":"J. Microbiol. Biotechnol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"855","DOI":"10.1080\/09583157.2012.694413","article-title":"Mechanisms of biocontrol and plant growth-promoting activity in soil bacterial species of Bacillus and Pseudomonas: A review","volume":"22","author":"Santoyo","year":"2012","journal-title":"Biocontrol Sci. Technol."},{"key":"ref_13","unstructured":"Singh, H.B., and Vaishnav, A. (2022). Pseudomonas and Bacillus: A biological tool for crop protection. New and Future Developments in Microbial Biotechnology and Bioengineering, Elsevier."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1868","DOI":"10.1080\/10643389.2020.1863112","article-title":"Heavy metal remediation and resistance mechanism of Aeromonas, Bacillus, and Pseudomonas: A review","volume":"52","author":"Fakhar","year":"2022","journal-title":"Crit. Rev. Environ. Sci. Technol."},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Singh, A.K., Kumar, A., and Singh, P.K. (2019). Tolerance of heavy metal toxicity using PGPR strains of Pseudomonas species. PGPR Amelioration in Sustainable Agriculture, Elsevier.","DOI":"10.1016\/B978-0-12-815879-1.00012-4"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"130243","DOI":"10.1016\/j.chemosphere.2021.130243","article-title":"Optimization of integrated phytoremediation system (IPS) for enhanced lead removal and restoration of soil microbial activities","volume":"277","author":"Manzoor","year":"2021","journal-title":"Chemosphere"},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Nnaji, N.D., Anyanwu, C.U., Miri, T., and Onyeaka, H. (2024). Mechanisms of heavy metal tolerance in bacteria: A review. Sustainability, 16.","DOI":"10.3390\/su162411124"},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Joshi, S., Gangola, S., Bhandari, G., Bhandari, N.S., Nainwal, D., Rani, A., Malik, S., and Slama, P. (2023). Rhizospheric bacteria: The key to sustainable heavy metal detoxification strategies. Front. Microbiol., 14.","DOI":"10.3389\/fmicb.2023.1229828"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"2967","DOI":"10.1007\/s00253-016-7364-4","article-title":"Genetic basis and importance of metal resistant genes in bacteria for bioremediation of contaminated environments with toxic metal pollutants","volume":"100","author":"Das","year":"2016","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_20","first-page":"422","article-title":"Heavy Metal Resistant Bacteria Isolation from Koodankulam Coast: Bacillus sp. and Pseudomonas sp. as Potential Agents for Bioremediation","volume":"44","author":"Punitha","year":"2023","journal-title":"J. Adv. Zool."},{"key":"ref_21","unstructured":"Mokrani, S. (2019). Recherche de Bact\u00e9ries PGPR\/B Isol\u00e9es de l\u2019ouest Alg\u00e9rien et leurs Effets de Biofertilisation et de Lutte Biologique \u00e0 l\u2019\u00e9gard de Quelques Microorganismes Phytopathog\u00e8nes de Phaseolus vulgaris L.. [Ph.D. Thesis, Universit\u00e9 Abderrahmane Mira]."},{"key":"ref_22","first-page":"6876","article-title":"Bacterial genomic DNA isolation using CTAB","volume":"50","author":"William","year":"2012","journal-title":"Sigma"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1007\/BF01731581","article-title":"A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences","volume":"16","author":"Kimura","year":"1980","journal-title":"J. Mol. Evol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1596","DOI":"10.1093\/molbev\/msm092","article-title":"MEGA4: Molecular evolutionary genetics analysis (MEGA) software version 4.0","volume":"24","author":"Tamura","year":"2007","journal-title":"Mol. Biol. Evol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1007\/BF00154233","article-title":"Metal resistance in Acinetobacter and its relation to \u03b2-lactamase production","volume":"6","author":"Deshpande","year":"1993","journal-title":"Biometals"},{"key":"ref_26","first-page":"195946","article-title":"In vitro screening for abiotic stress tolerance in potent biocontrol and plant growth promoting strains of Pseudomonas and Bacillus spp.","volume":"2014","author":"PraveenKumar","year":"2014","journal-title":"Int. J. Bacteriol."},{"key":"ref_27","unstructured":"CA-SFM: Comit\u00e9 de l\u2019Antibiogramme de la Soci\u00e9t\u00e9 Fran\u00e7aise de Microbiologie (2025, October 10). Communiqu\u00e9. Num\u00e9ro sp\u00e9cial SFM et site Internet. Available online: https:\/\/www.sfm-microbiologie.org\/wp-content\/uploads\/2020\/07\/Casfm_2005.pdf."},{"key":"ref_28","unstructured":"Mokrani, S. (2009). La Ruminococcine A et son r\u00f4le Antibact\u00e9rien \u00e0 l\u2019\u00e9gard de Souches Multir\u00e9sitantes Isol\u00e9es des H\u00f4pitaux de la R\u00e9gion de Bejaia. [Master\u2019s Thesis, Universit\u00e9 Abderrahmane Mira]."},{"key":"ref_29","first-page":"65","article-title":"Antibiogramme par diffusion en g\u00e9lose: Essai de standardisation de l\u2019inoculum par la m\u00e9thode \u00abPresto ABG\u00ae+\u00bb","volume":"1998","author":"Cavallo","year":"1998","journal-title":"Rev. Francoph. Lab."},{"key":"ref_30","unstructured":"CA-SFM: Comit\u00e9 de l\u2019Antibiogramme de la Soci\u00e9t\u00e9 Fran\u00e7aise de Microbiologie (2025, October 01). Communiqu\u00e9. Num\u00e9ro sp\u00e9cial SFM et site Internet. Available online: https:\/\/www.sfm-microbiologie.org\/wp-content\/uploads\/2024\/06\/CASFM2024_V1.0.pdf."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"199","DOI":"10.5001\/omj.2010.56","article-title":"In Vitro Susceptibility Test of Different Clinical Isolates against Ceftriaxone","volume":"25","author":"Masood","year":"2010","journal-title":"Oman Med. J."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"1462","DOI":"10.1016\/j.jiph.2019.11.013","article-title":"Evaluation of multidrug-resistant Bacillus strains causing public health risks in powdered infant milk formulas","volume":"13","author":"Alanber","year":"2020","journal-title":"J. Infect. Public Health"},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Gajewska, J., Chaj\u0119cka-Wierzchowska, W., and Zadernowska, A. (2022). Occurrenceand Characteristics of Staphylococcus aureus Strains along the Production Chain of Raw Milk Cheeses in Poland. Molecules, 27.","DOI":"10.3390\/molecules27196569"},{"key":"ref_34","first-page":"655","article-title":"A selective medium for addicting Leuconostoc in mixed-strain starter cultures","volume":"45","author":"Mayeux","year":"1962","journal-title":"J. Dairy. Sci."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1016\/0922-338X(92)90055-Y","article-title":"Isolation of a new surfactin producer Bacillus pumilus A-1, and cloning and nucleotide sequence of the regulator gene, psf-1","volume":"74","author":"Morikawa","year":"1992","journal-title":"J. Ferment. Bioeng."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1016\/j.bej.2008.06.016","article-title":"Isolation and comparison of biosurfactants produced by Bacillus subtilis PT2 and Pseudomonas aeruginosa SP4 for microbial surfactant-enhanced oil recovery","volume":"42","author":"Pornsunthorntawee","year":"2008","journal-title":"Biochem. Eng. J."},{"key":"ref_37","first-page":"7","article-title":"Antifungal activities of Pseudomonas strains against indigenous Phytopathogenic fungi of Phaseolus vulgaris L. isolated from a greenhouse in western Algeria","volume":"2","author":"Mokrani","year":"2019","journal-title":"Res. J. Plant Pathol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"47","DOI":"10.13057\/asianjagric\/g030203","article-title":"Potential of Pseudomonas stutzeri strains isolated from rhizospheric soil endowed with antifungal activities against phytopathogenic fungus Stemphylium botryosum","volume":"3","author":"Mokrani","year":"2019","journal-title":"Asian J. Agric."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1007\/BF02981094","article-title":"Antagonistic activity of bacteria from the Chickpea Rhizosphere against Fusarium oxysporium f. sp. ciceris","volume":"25","author":"Landa","year":"1997","journal-title":"Phytoparasitica"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"420","DOI":"10.5513\/JCEA01\/14.1.1217","article-title":"Inhibition of grey mould in vitro and in vivo with essential oil of fennel (Foeniculum vulgare L.)","volume":"14","author":"Mohammadi","year":"2013","journal-title":"J. Cent. Eur. Agric."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1007\/s40726-015-0024-y","article-title":"Chemical processes affecting the mobility of heavy metals and metalloids in soil environments","volume":"2","author":"Caporale","year":"2016","journal-title":"Curr. Pollut. Rep."},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Timkov\u00e1, I., Sedl\u00e1kov\u00e1-Kadukov\u00e1, J., and Prista\u0161, P. (2018). Biosorption and bioaccumulation abilities of actinomycetes\/streptomycetes isolated from metal contaminated sites. Separations, 5.","DOI":"10.3390\/separations5040054"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"101653","DOI":"10.1016\/j.jksus.2021.101653","article-title":"Health and environmental effects of heavy metals","volume":"34","author":"Zaynab","year":"2022","journal-title":"J. King Saud. Univ. Sci."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1016\/S0960-8524(97)00161-2","article-title":"Resistance of environmental bacteria to heavy metals","volume":"64","author":"Hassen","year":"1998","journal-title":"Bioresour. Technol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1405","DOI":"10.1016\/S0883-2927(01)00042-7","article-title":"Ecotoxicological assessment of heavy metals in sewage sludge amended soils","volume":"16","author":"Obbard","year":"2001","journal-title":"Appl. Geochem."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"3223","DOI":"10.1128\/AEM.69.6.3223-3230.2003","article-title":"Cultivation- dependent approaches for determining bacterial diversity in heavy-metal-contaminated soil","volume":"69","author":"Ellis","year":"2003","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1128\/aem.53.1.185-188.1987","article-title":"High incidence of selenite-resistant bacteria from a site polluted with selenium","volume":"53","author":"Burton","year":"1987","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1080\/03067319508041330","article-title":"Chemical speciation and fractionation in soil and sediment heavy metal analysis: A review","volume":"59","author":"Tack","year":"1995","journal-title":"Int. J. Environ. Anal. Chem."},{"key":"ref_49","first-page":"1","article-title":"Speciation of toxic metals","volume":"28","author":"Hassan","year":"2022","journal-title":"Egy. Sci. J. Pestic."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"170","DOI":"10.3390\/toxics3020170","article-title":"Speciation in metal toxicity and metal-based therapeutics","volume":"3","author":"Templeton","year":"2015","journal-title":"Toxics"},{"key":"ref_51","first-page":"121","article-title":"Physiological and genetic responses of bacteria to osmotic stress","volume":"53","author":"Csonka","year":"1989","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"525","DOI":"10.1007\/s11104-019-04143-5","article-title":"Effect of NaCl on proline and glycinebetaine metabolism in Kosteletzkya pentacarpos exposed to cd and Zn toxicities","volume":"441","author":"Zhou","year":"2019","journal-title":"Plant Soil"},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Li, Y.Q., Chai, Y.H., Wang, X.S., Huang, L.Y., Luo, X.M., Qiu, C., Liu, Q.H., and Guan, X.Y. (2021). Bacterial community in saline farmland soil on the Tibetan plateau: Responding to salinization while resisting extreme environments. BMC Microbiol., 21.","DOI":"10.1186\/s12866-021-02190-6"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"504","DOI":"10.1128\/MMBR.62.2.504-544.1998","article-title":"Biology of moderately halophilic aerobic bacteria","volume":"62","author":"Ventosa","year":"1998","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"2385","DOI":"10.1128\/aem.55.9.2385-2390.1989","article-title":"Survey of metal tolerance in moderately halophilic eubacteria","volume":"55","author":"Nieto","year":"1989","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_56","first-page":"164","article-title":"Isolation and characterization of Pseudomonas resistant to heavy metals contaminants","volume":"3","author":"Singh","year":"2010","journal-title":"Int. J. Pharm. Sci. Rev. Res."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/S0378-1119(96)00323-X","article-title":"Bacterial resistances to toxic metal ions-a review","volume":"179","author":"Silver","year":"1996","journal-title":"Gene"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"418","DOI":"10.1016\/j.psep.2022.11.020","article-title":"Effects of heavy metals and antibiotics on antibiotic resistance genes and microbial communities in soil","volume":"169","author":"Fu","year":"2023","journal-title":"Process Saf. Environ. Prot."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1186\/s42834-021-00088-6","article-title":"Resistance of bacteria isolated from leachate to heavy metals and the removal of Hg by Pseudomonas aeruginosa strain FZ-2 at different salinity levels in a batch biosorption system","volume":"31","author":"Imron","year":"2021","journal-title":"Sustain. Environ. Res."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"242","DOI":"10.1007\/s10924-019-01602-z","article-title":"Characterization of exopolysaccharide produced by Pseudomonas sp. PFAB4 for synthesis of EPS-coated AgNPs with antimicrobial properties","volume":"28","author":"Banerjee","year":"2020","journal-title":"J. Polym. Environ."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"63","DOI":"10.2166\/wst.2005.0182","article-title":"Effect of exopolysaccharides on the adsorption of metal ions by Pseudomonas sp. CU-1","volume":"52","author":"Lau","year":"2005","journal-title":"Water Sci. Technol."},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Bal\u00edkov\u00e1, K., Vojtkov\u00e1, H., Duborsk\u00e1, E., Kim, H., Mat\u00fa\u0161, P., and Ur\u00edk, M. (2022). Role of exopolysaccharides of Pseudomonas in heavy metal removal and other remediation strategies. Polymers, 14.","DOI":"10.3390\/polym14204253"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"236","DOI":"10.1007\/s11515-006-0028-5","article-title":"The production-influencing factors of extracellular polysaccharide (EPS) from a strain of Lactic Acid Bacteria and EPS extraction","volume":"3","author":"Chen","year":"2006","journal-title":"Front. Biol. China"},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Rosenberg, E., DeLong, E.F., Lory, S., Stackebrandt, E., and Thompson, F. (2013). Microbial Exopolysaccharides. The Prokaryotes: Applied Bacteriology and Biotechnology 4e, Springer Reference.","DOI":"10.1007\/978-3-642-31331-8"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"1710","DOI":"10.1101\/gad.246397.114","article-title":"Self-regulation of exopolysaccharide production in Bacillus subtilis by a tyrosine kinase","volume":"28","author":"Elsholz","year":"2014","journal-title":"Genes Dev."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"512","DOI":"10.1134\/S0026261715040153","article-title":"The production of exopolysaccharide by Pseudomonas putida GAP-P45 under various abiotic stress conditions and its role in soil aggregation","volume":"84","author":"Sandhya","year":"2015","journal-title":"Microbiology"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1007\/s12088-011-0148-8","article-title":"Potentials of exopolysaccharides from lactic acid bacteria","volume":"52","author":"Patel","year":"2012","journal-title":"Indian J. Microbiol."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"104024","DOI":"10.1016\/j.resmic.2022.104024","article-title":"Bacterial exopolysaccharides as emerging bioactive macromolecules: From fundamentals to applications","volume":"174","author":"Kaur","year":"2023","journal-title":"Res. Microbiol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"254","DOI":"10.1111\/j.1365-2958.2007.06040.x","article-title":"Bistability and biofilm formation in Bacillus subtilis","volume":"67","author":"Chai","year":"2008","journal-title":"Mol. Microbiol."},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Rane, A.N., Baikar, V.V., Ravi Kumar, V., and Deopurkar, R.L. (2017). Agro-industrial wastes for production of biosurfactant by Bacillus subtilis ANR 88 and its application in synthesis of silver and gold nanoparticles. Front. Microbiol., 8, Erratum in Front. Microbiol. 2017, 8, 878. https:\/\/doi.org\/10.3389\/fmicb.2017.00878. PMID: 28392783; PMCID: PMC5364166.","DOI":"10.3389\/fmicb.2017.00492"},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Sober\u00f3n-Ch\u00e1vez, G. (2011). Biosurfactants: A general overview. Biosurfactants, Springer-Verlag.","DOI":"10.1007\/978-3-642-14490-5"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"919","DOI":"10.1080\/01932691.2011.580183","article-title":"Sorption of chromate and arsenate by surfactant modified erionite (E-SMZ)","volume":"33","author":"Swarnkar","year":"2012","journal-title":"J. Dispers. Sci. Technol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"4832","DOI":"10.1007\/s12010-022-04288-7","article-title":"Biofilm-mediated heavy metal removal from aqueous system by multi-metal-resistant bacterial strain Bacillus sp. GH-s29","volume":"195","author":"Maity","year":"2023","journal-title":"Appl. Biochem. Biotechnol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"15593258221117352","DOI":"10.1177\/15593258221117352","article-title":"Biosorptive potential of Pseudomonas species RY12 toward zinc heavy metal in agriculture soil irrigated with contaminated waste water","volume":"20","author":"Yasmin","year":"2022","journal-title":"Dose-Response"},{"key":"ref_75","first-page":"201","article-title":"Heavy Metal Resistance and Gene Expression Analysis of Metal Resistance Genes in Gram-Positive and Gram-Negative Bacteria Present in Egyptian Soils","volume":"6","author":"Abdelatey","year":"2011","journal-title":"J. Appl. Sci. Environ. Sanit."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"5855","DOI":"10.1038\/s41598-019-41899-3","article-title":"Metal resistant PGPR lowered Cd uptake and expression of metal transporter genes with improved growth and photosynthetic pigments in Lycopersicon esculentum under metal toxicity","volume":"9","author":"Khanna","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"1511","DOI":"10.3390\/microbiolres14040103","article-title":"Evaluation of biocontrol potential of Bacillus spp. and Pseudomonas fluorescens UM270 against postharvest fungal pathogens","volume":"14","author":"Santoyo","year":"2023","journal-title":"Microbiol. Res."},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"He, D.C., He, M.H., Amalin, D.M., Liu, W., Alvindia, D.G., and Zhan, J. (2021). Biological control of plant diseases:An evolutionary and eco-economic consideration. Pathogens, 10.","DOI":"10.3390\/pathogens10101311"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1016\/j.tim.2007.12.009","article-title":"Bacillus lipopeptides: Versatile weapons for plant disease biocontrol","volume":"16","author":"Ongena","year":"2008","journal-title":"Trends Microbiol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1016\/j.jbiotec.2018.07.044","article-title":"Biological control of plant pathogens by Bacillus species","volume":"285","author":"Fira","year":"2018","journal-title":"J. Biotechnol."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1937","DOI":"10.1016\/S0006-3495(95)80370-X","article-title":"Interactions of surfactin with membrane models","volume":"68","author":"Ptak","year":"1995","journal-title":"Biophys. J."},{"key":"ref_82","first-page":"2418","article-title":"Fengycin interaction with lipid monolayers at the air-water interface","volume":"1778","author":"Deleu","year":"2008","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"430","DOI":"10.1094\/MPMI-20-4-0430","article-title":"The iturin and fengycin families of lipopeptides are key factors in antagonism of Bacillus subtilis toward Podosphaera fusca","volume":"20","author":"Romero","year":"2007","journal-title":"Mol. Plant-Microbe Interact."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"1005","DOI":"10.1007\/s00253-012-4641-8","article-title":"Biosurfactants in agriculture","volume":"97","author":"Sachdev","year":"2013","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_85","first-page":"1662","article-title":"Biological control of Rhizoctonia root rot on bean by phenazine- and cyclic lipopeptide-producing Pseudomonas strains","volume":"13","author":"Hua","year":"2011","journal-title":"Environ. Microbiol."},{"key":"ref_86","first-page":"1691","article-title":"Review: Lipopeptide biosurfactants: Classes, production, and applications","volume":"38","author":"Mnif","year":"2015","journal-title":"Bioprocess. Biosyst. Eng."}],"container-title":["Microorganisms"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2607\/14\/3\/644\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,3,12]],"date-time":"2026-03-12T13:56:22Z","timestamp":1773323782000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2607\/14\/3\/644"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,3,12]]},"references-count":86,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2026,3]]}},"alternative-id":["microorganisms14030644"],"URL":"https:\/\/doi.org\/10.3390\/microorganisms14030644","relation":{},"ISSN":["2076-2607"],"issn-type":[{"value":"2076-2607","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,3,12]]}}}