REVIEW PAPER
Exopolysaccharide-Producing PGPR: Mechanisms
for Alleviating Salinity-Induced Plant Stress
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Department of Biology, College of Science in Zulfi, Majmaah University, Al-Majmaah 11952, Saudi Arabia
Submission date: 2024-09-06
Final revision date: 2024-10-16
Acceptance date: 2024-12-02
Online publication date: 2025-03-27
Publication date: 2026-01-30
Corresponding author
Wardah A. Alhoqail
Department of Biology, College of Science in Zulfi,, Majmaah University, Al-Majmaah Saudi Arabia., 11952, Al-Majmaah, Saudi Arabia
Pol. J. Environ. Stud. 2026;35(1):501-518
KEYWORDS
TOPICS
ABSTRACT
Elevated salinity levels pose a significant challenge to global crop production, affecting approximately
85% of the Earth’s arable land. The stress induces disruptions in vegetative development and yield by
inducing osmotic imbalance, toxicity from ions, and membrane instabilities, ensuing in a substantial
fall in crop production. However, the administration of EPS-producing bacteria promises an optimistic
approach that can minimize the effects of salinity and boost crop resistance in saline conditions. EPSproducing
bacteria have a dynamic role in reducing the harmful effects of salt stress through several
mechanisms, such as restricting the absorption of sodium, improving the intake of beneficial ions such
as K+, Ca2+, Mg2+, and P, elevating levels of antioxidants, and encouraging the establishment of soil
aggregates. These bacteria assist in regulating ion balance in plant cells experiencing salt stress, hence
improving plant development traits and stress tolerance. When the level of salt elevates in the soil, these
bacterial strains form a protective layer (biofilm) around roots and prevent the uptake of sodium ions
into the plant. EPS also forms complexes with sodium ions in the soil, so decreasing their accessibility
for plant uptake and contributing to the maintenance of an ideal ion equilibrium for plant survival.
In addition, bacteria that produce EPS enhance soil structure and fertility by promoting soil aggregation,
enhancing nutrient availability, and improving water retention capacity. EPS-producing bacteria have
a crucial role in enhancing the health and productivity of plants cultivated in saline environments.
In addition, bacteria that produce EPS show positive impacts on the efficiency of photosynthesis,
the amount of water in plants, and markers of oxidative stress in plants that are exposed to salt
stress. These bacteria improve photosynthetic rates by enhancing food availability and water uptake
in the rhizosphere, resulting in higher chlorophyll production and overall plant development.
Furthermore, bacteria that produce EPS increase the water content in plant tissues, resulting in better
hydration and overall plant health. Furthermore, these bacteria reduce levels of oxidative stress markers
such as malondialdehyde (MDA) and hydrogen peroxide (H2O2), therefore increasing antioxidant
defenses and reducing oxidative damage in plants subjected to salt stress.
CONFLICT OF INTEREST
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
REFERENCES (117)
1.
SHARMA A., KUMAR V., SHAHZAD B., RAMAKRISHNAN M., SINGH SIDHU G.P., BALI A.S., HANDA N., KAPOOR D., YADAV P., KHANNA K. Photosynthetic response of plants under different abiotic stresses: a review. Journal of Plant Growth Regulation. 39, 509, 2020.
https://doi.org/10.1007/s00344....
2.
VAROL T., CETIN M., OZEL H.B., SEVIK H., ZEREN CETIN I. The effects of climate change scenarios on Carpinus betulus and Carpinus orientalis in Europe. Water, Air, & Soil Pollution. 233 (2), 45, 2022.
https://doi.org/10.1007/s11270....
3.
BERA K., DUTTA P., SADHUKHAN S. Plant responses under abiotic stress and mitigation options towards agricultural sustainability. In: Plant Stress: Challenges and Management in the New Decade, Chapter 1, Springer, 2022.
https://doi.org/10.1007/978-3-....
4.
CETIN M., ALJAMA A.M.O., ALRABITI O.B.M., ADIGUZEL F., SEVIK H., ZEREN CETIN I. Determination and mapping of regional change of Pb and Cr pollution in Ankara city center. Water, Air, & Soil Pollution. 233 (5), 163, 2022.
https://doi.org/10.1007/s11270....
5.
KHAN A.A., WANG T., HUSSAIN T., AMNA ALI F., SHI F., LATEF A.A.H.A., ALI O.M., HAYAT K., MEHMOOD S. Halotolerant-Koccuria rhizophila (14asp)-induced amendment of salt stress in pea plants by limiting Na+ uptake and elevating production of antioxidants. Agronomy. 11 (10), 1907, 2021.
https://doi.org/10.3390/agrono....
6.
DIKILITAS M., SIMSEK E., KARAKAS S., LATEF A.A.H.A. Abiotic stresses and their interactions with each other on plant growth, development and defense mechanisms. CRC Press. 2021.
https://doi.org/10.1201/978100....
7.
TEKIN O., CETIN M., VAROL T., OZEL H.B., SEVIK H., ZEREN CETIN I. Altitudinal migration of species of Fir (Abies spp.) in adaptation to climate change. Water, Air, & Soil Pollution. 233 (9), 385, 2022.
https://doi.org/10.1007/s11270....
8.
CETIN M., ALJAMA A.M.O., ALRABITI O.B.M., ADIGUZEL F., SEVIK H., ZEREN CETIN I. Using topsoil analysis to determine and map changes in Ni Co pollution. Water, Air, & Soil Pollution. 233 (8), 293, 2022.
https://doi.org/10.1007/s11270....
9.
KHAN I., RAZA M.A., AWAN S.A., SHAH G.A., RIZWAN M., ALI B., TARIQ R., HASSAN M.J., ALYEMENI M.N., BRESTIC M. Amelioration of salt induced toxicity in pearl millet by seed priming with silver nanoparticles (AgNPs): The oxidative damage, antioxidant enzymes and ions uptake are major determinants of salt tolerant capacity. Plant Physiology and Biochemistry. 156, 221, 2020.
https://doi.org/10.1016/j.plap... PMid:32979796.
10.
CETIN M., ADIGUZEL F., ZEREN CETIN I. Determination of the effect of urban forests and other green areas on surface temperature in Antalya. Springer, 2023.
https://doi.org/10.1007/978-98....
11.
HAIDER F.U., LIQUN C., COULTER J.A., CHEEMA S.A., WU J., ZHANG R., WENJUN M., FAROOQ M. Cadmium toxicity in plants: Impacts and remediation strategies. Ecotoxicology and Environmental Safety. 211, 111887, 2021.
https://doi.org/10.1016/j.ecoe... PMid:33450535.
12.
MUKHERJEE P., MITRA A., ROY M. Halomonas Rhizobacteria of Avicennia marina of Indian Sundarbans Promote Rice Growth Under Saline and Heavy Metal Stresses Through Exopolysaccharide Production. Frontiers in Microbiology. 10, 2019.
https://doi.org/10.3389/fmicb.... PMid:31191507 PMCid:PMC6549542.
13.
XIE X., HE Z., CHEN N., TANG Z., WANG Q., CAI Y. The roles of environmental factors in regulation of oxidative stress in plant. BioMed Research International. 2019, 2019.
https://doi.org/10.1155/2019/9... PMid:31205950 PMCid:PMC6530150.
15.
VERMA P., YADAV A.N., KUMAR V., SINGH D.P., SAXENA A.K. Beneficial plant-microbes interactions: biodiversity of microbes from diverse extreme environments and its impact for crop improvement. In: Plant-microbe interactions in agro-ecological perspectives. Chapter 22, Springer Nature Singapore, 2017.
https://doi.org/10.1007/978-98....
16.
SIES H. On the history of oxidative stress: Concept and some aspects of current development. Current Opinion in Toxicology. 7, 122, 2018.
https://doi.org/10.1016/j.coto....
17.
LUSHCHAK V.I. Free radicals, reactive oxygen species, oxidative stress and its classification. Chemico-Biological Interactions. 224, 164, 2014.
https://doi.org/10.1016/j.cbi.... PMid:25452175.
18.
ZAINAB N., AMNA KHAN A.A., AZEEM M.A., ALI B., WANG T., SHI F., ALGHANEM S.M., HUSSAIN MUNIS M.F., HASHEM M. PGPR-mediated plant growth attributes and metal extraction ability of Sesbania sesban L. in industrially contaminated soils. Agronomy. 11 (9), 1820, 2021.
https://doi.org/10.3390/agrono....
19.
ZAINAB N., DIN B.U., JAVED M.T., AFRIDI M.S., MUKHTAR T., KAMRAN M.A., KHAN A.A., ALI J., JATOI W.N., MUNIS M.F.H. Deciphering metal toxicity responses of flax (Linum usitatissimum L.) with exopolysaccharide and ACC-deaminase producing bacteria in industrially contaminated soils. Plant Physiology and Biochemistry. 152, 90, 2020.
https://doi.org/10.1016/j.plap... PMid:32408178.
20.
DUMONT S., RIVOAL J. Consequences of oxidative stress on plant glycolytic and respiratory metabolism. Frontiers in Plant Science. 10, 432113, 2019.
https://doi.org/10.3389/fpls.2... PMid:30833954 PMCid:PMC6387960.
21.
HAYAT K., MENHAS S., BUNDSCHUH J., ZHOU P., NIAZI N.K., AMNA HUSSAIN A., HAYAT S., ALI H., WANG J. Plant growth promotion and enhanced uptake of Cd by combinatorial application of Bacillus pumilus and EDTA on Zea mays L. International Journal of Phytoremediation. 22 (13), 1372, 2020.
https://doi.org/10.1080/152265... PMid:32579378.
22.
SOLIMAN M.H., ALNUSAIRI G.S., KHAN A.A., ALNUSAIRE T.S., FAKHR M.A., ABDULMAJEED A.M., ALDESUQUY H.S., YAHYA M., NAJEEB U. Biochar and selenium nanoparticles induce water transporter genes for sustaining carbon assimilation and grain production in salt-stressed wheat. Journal of Plant Growth Regulation. 42 (3), 1522, 2023.
https://doi.org/10.1007/s00344....
24.
MATHEW L. Understanding The Significance Of Pre-Treatment In The Post-Cryopreservation Survival Of Kiwifruit Shoot Tips Through Biochemical And Ultrastructural Studies. University of Otago, 2015.
25.
HUANG H., ULLAH F., ZHOU D.-X., YI M., ZHAO Y. Mechanisms of ROS regulation of plant development and stress responses. Frontiers in Plant Science. 10, 800, 2019.
https://doi.org/10.3389/fpls.2... PMid:31293607 PMCid:PMC6603150.
26.
AL-MUSHHIN A.A., QARI S.H., FAKHR M.A., ALNUSAIRI G.S., ALNUSAIRE T.S., ALRASHIDI A.A., LATEF A.A.H.A., ALI O.M., KHAN A.A., SOLIMAN M.H. Exogenous myo-inositol alleviates salt stress by enhancing antioxidants and membrane stability via the upregulation of stress responsive genes in Chenopodium quinoa L. Plants. 10 (11), 2416, 2021.
https://doi.org/10.3390/plants... PMid:34834781 PMCid:PMC8623490.
27.
DUMANOVIĆ J., NEPOVIMOVA E., NATIĆ M., KUČA K., JAĆEVIĆ V. The significance of reactive oxygen species and antioxidant defense system in plants: A concise overview. Frontiers in Plant Science. 11, 552969, 2021.
https://doi.org/10.3389/fpls.2... PMid:33488637 PMCid:PMC7815643.
28.
MEHMOOD S., KHAN A.A., SHI F., TAHIR M., SULTAN T., MUNIS M.F.H., KAUSHIK P., ALYEMENI M.N., CHAUDHARY H.J. Alleviation of salt stress in wheat seedlings via multifunctional Bacillus aryabhattai PM34: an in-vitro study. Sustainability. 13 (14), 8030, 2021.
https://doi.org/10.3390/su1314....
29.
ALI S., BHARWANA S.A., RIZWAN M., FARID M., KANWAL S., ALI Q., IBRAHIM M., GILL R.A., KHAN M.D. Fulvic acid mediates chromium (Cr) tolerance in wheat (Triticum aestivum L.) through lowering of Cr uptake and improved antioxidant defense system. Environmental Science and Pollution Research. 22, 10601, 2015.
https://doi.org/10.1007/s11356... PMid:25744818.
30.
KHAN A.A., WANG T., NISA Z.U., ALNUSAIRI G.S., SHI F. Insights into cadmium-induced morphophysiological disorders in Althea rosea cavan and its phytoremediation through the exogeneous citric acid. Agronomy. 12 (11), 2776, 2022.
https://doi.org/10.3390/agrono....
32.
AMNA UD DIN B., SARFRAZ S., XIA Y., KAMRAN M.A., JAVED M.T., SULTAN T., HUSSAIN MUNIS M.F., CHAUDHARY H.J. Mechanistic elucidation of germination potential and growth of wheat inoculated with exopolysaccharide and ACC-deaminase producing Bacillus strains under induced salinity stress. Ecotoxicology and Environmental Safety. 183, 109466, 2019.
https://doi.org/10.1016/j.ecoe... PMid:31408821.
33.
ARAYES M.A., MABROUK M.E.M., SABRY S.A., ABDELLA B. Exopolysaccharide production from Alkalibacillus sp. w3: statistical optimization and biological activity. Biologia. 78 (1), 229, 2023.
https://doi.org/10.1007/s11756....
34.
ASHRAF M., HASNAIN S., HUSSAIN F. Exopolysaccharides (EPS) producing biofilm bacteria in improving physico-chemical characteristics of the salt-affected soils. Proceedings of International Conference Environment and Sustainable Development. COMSATS Institute of Information Technology, Abbottabad, Pakistan. 2005.
35.
NGUYEN P.T., NGUYEN T.T., BUI D.C., HONG P.T., HOANG Q.K., NGUYEN H.T. Exopolysaccharide production by lactic acid bacteria: the manipulation of environmental stresses for industrial applications. AIMS Microbiol. 6 (4), 451, 2020.
36.
SANDHYA V., ALI S.Z. The production of exopolysaccharide by Pseudomonas putida GAP-P45 under various abiotic stress conditions and its role in soil aggregation. Microbiology. 84, 512, 2015.
https://doi.org/10.1134/S00262....
37.
SINGH R., SINGH J., DEVAL R., UPADHYAY S., KUMAR D. The potentiality of selected strain of PGPR: azotobacter, for sustainable agriculture in India. G-Journal of Environmental Science and Technology. 4 (6), 49, 2017.
38.
TEWARI S., SHARMA S. Rhizobial exopolysaccharides as supplement for enhancing nodulation and growth attributes of Cajanus cajan under multi-stress conditions: A study from lab to field. Soil and Tillage Research. 198, 104545, 2020.
https://doi.org/10.1016/j.stil....
39.
WU N., PAN H.-X., QIU D., ZHANG Y.-M. Feasibility of EPS-producing bacterial inoculation to speed up the sand aggregation in the Gurbantunggut Desert, Northwestern China. Journal of Basic Microbiology. 54 (12), 1378, 2014.
https://doi.org/10.1002/jobm.2... PMid:25224518.
40.
RAMADAN E.M., ABDELHAFEZ A.A., HASSAN E.A., SABER F.M. Plant growth promoting rhizobacteria and their potential for biocontrol of phytopathogens. African Journal of Microbiology Research. 10 (15), 486, 2016.
https://doi.org/10.5897/AJMR20....
41.
KUDOYAROVA G., ARKHIPOVA T., MELENT'EV A. Role of bacterial phytohormones in plant growth regulation and their development. Bacterial Metabolites in Sustainable Agroecosystem. 69, 2015.
https://doi.org/10.1007/978-3-....
42.
MAITRA S., BRESTIC M., BHADRA P., SHANKAR T., PRAHARAJ S., PALAI J.B., SHAH M.M.R., BAREK V., ONDRISIK P., SKALICKÝ M. Bioinoculants-natural biological resources for sustainable plant production. Microorganisms. 10 (1), 51, 2021.
https://doi.org/10.3390/microo... PMid:35056500 PMCid:PMC8780112.
43.
HOSSAIN A., DA SILVA J.A.T., LOZOVSKAYA M.V., ZVOLINSKY V.P. The effect of high temperature stress on the phenology, growth and yield of five wheat (Triticum aestivum L.) varieties. Asian and Australasian Journal of Plant Science and Biotechnology. 6 (1), 14, 2012.
44.
ODOH C.K. Plant growth promoting rhizobacteria (PGPR): a bioprotectant bioinoculant for sustainable agrobiology. A review. International Journal of Advanced Research in Biological Sciences. 4 (5), 123, 2017.
https://doi.org/10.22192/ijarb....
45.
PANHWAR Q., RADZIAH O., RAHMAN A.Z., SARIAH M., RAZI I.M., NAHER U. Contribution of phosphate-solubilizing bacteria in phosphorus bioavailability and growth enhancement of aerobic rice. Spanish Journal of Agricultural Research. 9 (3), 810, 2011.
https://doi.org/10.5424/sjar/2....
46.
SWARNALAKSHMI K., YADAV V., TYAGI D., DHAR D.W., KANNEPALLI A., KUMAR S. Significance of Plant Growth Promoting Rhizobacteria in Grain Legumes: Growth Promotion and Crop Production. Plants. 9 (11), 1596, 2020.
https://doi.org/10.3390/plants... PMid:33213067 PMCid:PMC7698556.
47.
PII Y., MIMMO T., TOMASI N., TERZANO R., CESCO S., CRECCHIO C. Microbial interactions in the rhizosphere: beneficial influences of plant growth-promoting rhizobacteria on nutrient acquisition process. A review. Biology and Fertility of Soils. 51 (4), 403, 2015.
https://doi.org/10.1007/s00374....
48.
HASSAN M.K., MCINROY J.A., KLOEPPER J.W. The Interactions of Rhizodeposits with Plant Growth-Promoting Rhizobacteria in the Rhizosphere: A Review. Agriculture. 9 (7), 142, 2019.
https://doi.org/10.3390/agricu....
49.
ANGELIN J., KAVITHA M. Exopolysaccharides from probiotic bacteria and their health potential. International Journal of Biological Macromolecules. 162, 853, 2020.
https://doi.org/10.1016/j.ijbi... PMid:32585269 PMCid:PMC7308007.
50.
POLI A., ANZELMO G., NICOLAUS B. Bacterial exopolysaccharides from extreme marine habitats: production, characterization and biological activities. Marine Drugs. 8 (6), 1779, 2010.
https://doi.org/10.3390/md8061... PMid:20631870 PMCid:PMC2901825.
51.
ATES O. Systems biology of microbial exopolysaccharides production. Frontiers in Bioengineering and Biotechnology. 3, 200, 2015.
https://doi.org/10.3389/fbioe.... PMid:26734603 PMCid:PMC4683990.
52.
FREITAS F., TORRES C.A.V., REIS M.A.M. Engineering aspects of microbial exopolysaccharide production. Bioresource Technology. 245, 1674, 2017.
https://doi.org/10.1016/j.bior... PMid:28554522.
53.
YANG Y., JIANG G., TIAN Y. Biological activities and applications of exopolysaccharides produced by lactic acid bacteria: a mini-review. World Journal of Microbiology and Biotechnology. 39 (6), 155, 2023.
https://doi.org/10.1007/s11274... PMid:37039945.
54.
RACIOPPO A., D'AMELIO A., DE SANTIS A., BEVILACQUA A., CORBO M.R., SINIGAGLIA M. Potential Use of Plant Growth-Promoting Bacteria to Enhance Growth and Soil Fertility in Marginal Areas: Focus on the Apulia Region, Italy. Agronomy. 13 (12), 2983, 2023.
https://doi.org/10.3390/agrono....
55.
CETIN M., ABO AISHA A.E.S. Variation of Al concentrations depending on the growing environment in some indoor plants that used in architectural designs. Environmental Science and Pollution Research. 30 (7), 18748, 2023.
https://doi.org/10.1007/s11356... PMid:36219289.
56.
MASMOUDI F., ALSAFRAN M., JABRI H.A., HOSSEINI H., TRIGUI M., SAYADI S., TOUNSI S., SAADAOUI I. Halobacteria-Based Biofertilizers: A Promising Alternative for Enhancing Soil Fertility and Crop Productivity under Biotic and Abiotic Stresses - A Review. Microorganisms. 11 (5), 1248, 2023.
https://doi.org/10.3390/microo... PMid:37317222 PMCid:PMC10222427.
57.
HASAN A., TABASSUM B., HASHIM M., KHAN N. Role of Plant Growth Promoting Rhizobacteria (PGPR) as a Plant Growth Enhancer for Sustainable Agriculture: A Review. Bacteria. 3 (2), 59, 2024.
https://doi.org/10.3390/bacter....
58.
ALI N., ABBAS S.A.A.A., SHARIF L., SHAFIQ M., KAMRAN Z., MASAH HASEEB M., SHAHID M.A. Microbial extracellular polymeric substance and impacts on soil aggregation, Chapter 13. Elsevier, 2024.
https://doi.org/10.1016/B978-0....
59.
DAR A., ZAHIR Z.A., IQBAL M., MEHMOOD A., JAVED A., HUSSAIN A., BUSHRA AHMAD M. Efficacy of rhizobacterial exopolysaccharides in improving plant growth, physiology, and soil properties. Environmental Monitoring and Assessment. 193 (8), 515, 2021.
https://doi.org/10.1007/s10661... PMid:34304322.
60.
OSEMWEGIE O.O., ADETUNJI C.O., AYENI E.A., ADEJOBI O.I., ARISE R.O., NWONUMA C.O., OGHENEKARO A.O. Exopolysaccharides from bacteria and fungi: current status and perspectives in Africa. Heliyon. 6 (6), e04205, 2020.
https://doi.org/10.1016/j.heli... PMid:32577572 PMCid:PMC7303563.
61.
KEMP B.P., HORNE J., BRYANT A., COOPER R.M. Xanthomonas axonopodis pv. manihotis gumD gene is essential for EPS production and pathogenicity and enhances epiphytic survival on cassava (Manihot esculenta). Physiological and Molecular Plant Pathology. 64 (4), 209, 2004.
https://doi.org/10.1016/j.pmpp....
62.
TEO H.M., AZIZ A., WAHIZATUL A.A., BHUBALAN K., SITI NORDAHLIAWATE M.S., MUHAMAD SYAZLIE C.I., LEE CHUEN Ng. Setting a Plausible Route for Saline Soil-Based Crop Cultivations by Application of Beneficial Halophyte-Associated Bacteria: A Review. Microorganisms. 10 (3), 657, 2022.
https://doi.org/10.3390/microo... PMid:35336232 PMCid:PMC8953261.
63.
LAMICHHANE J.R., SOLTANI E. Sowing and seedbed management methods to improve establishment and yield of maize, rice and wheat across drought-prone regions: A review. Journal of Agriculture and Food Research. 2, 100089, 2020.
https://doi.org/10.1016/j.jafr....
64.
UPADHYAY S.K., SINGH D.P. Effect of salt-tolerant plant growth-promoting rhizobacteria on wheat plants and soil health in a saline environment. Plant Biology. 17 (1), 288, 2015.
https://doi.org/10.1111/plb.12... PMid:24750405.
65.
THAKUR R., SRIVASTAVA S., YADAV S. Multitrait Pseudomonas sp. isolated from the rhizosphere of Bergenia ciliata acts as a growth-promoting bioinoculant for plants. Frontiers in Sustainable Food Systems. 7, 2023.
https://doi.org/10.3389/fsufs.....
66.
NOZARI R.M., ORTOLAN F., ASTARITA L.V., SANTARÉM E.R. Streptomyces spp. enhance vegetative growth of maize plants under saline stress. Brazilian Journal of Microbiology. 52 (3), 1371, 2021.
https://doi.org/10.1007/s42770... PMid:33834385 PMCid:PMC8324692.
67.
THAKUR R., YADAV S. Patent landscaping and citation network analysis to reveal the global research trends in biopriming using microbial inoculants: an insight toward sustainable agriculture. Biologia Futura. 74 (4), 545, 2023.
https://doi.org/10.1007/s42977... PMid:37995047.
68.
THAKUR R., YADAV S. Thermotolerant and halotolerant Streptomyces sp. isolated from Ajuga parviflora having biocontrol activity against Pseudomonas syringae and Xanthomonas campestris acts as a sustainable bioadditive in growth promotion of Cicer arietinum. Physiological and Molecular Plant Pathology. 127, 102059, 2023.
https://doi.org/10.1016/j.pmpp....
69.
CESUR A., ZEREN CETIN I., CETIN M., SEVIK H., OZEL H.B. The use of Cupressus arizonica as a biomonitor of Li, Fe, and Cr pollution in Kastamonu. Water, Air, & Soil Pollution. 233 (6), 193, 2022.
https://doi.org/10.1007/s11270....
70.
KUMARI P., GUPTA A., CHANDRA H., SINGH P., YADAV S. Effects of Salt Stress on the Morphology, Anatomy, and Gene Expression of Crop Plants. In: Physiology of Salt Stress in Plants. pp. 87, Wiley, 2021.
https://doi.org/10.1002/978111... PMid:32918207.
71.
CICEK N., ERDOGAN M., YUCEDAG C., CETIN M. Improving the detrimental aspects of salinity in salinized soils of arid and semi-arid areas for effects of vermicompost leachate on salt stress in seedlings. Water, Air, & Soil Pollution. 233 (6), 197, 2022.
https://doi.org/10.1007/s11270....
72.
JHA S., SINGH A.A., THAKUR N. Role of exopolysaccharide and biofilms in microorganisms for alleviating salt stress. Chapter 10, Academic Press, 2022.
https://doi.org/10.1016/B978-0... PMid:38013818 PMCid:PMC10653561.
73.
MOKRANI S., NABTI E.-H., CRUZ C. Current Advances in Plant Growth Promoting Bacteria Alleviating Salt Stress for Sustainable Agriculture. Applied Sciences. 10 (20), 7025, 2020.
https://doi.org/10.3390/app102....
74.
GIANNELLI G., POTESTIO S., VISIOLI G. The Contribution of PGPR in Salt Stress Tolerance in Crops: Unravelling the Molecular Mechanisms of Cross-Talk between Plant and Bacteria. Plants. 12 (11), 2197, 2023.
https://doi.org/10.3390/plants... PMid:37299176 PMCid:PMC10255858.
75.
SAHA I., DATTA S., BISWAS D. Exploring the Role of Bacterial Extracellular Polymeric Substances for Sustainable Development in Agriculture. Current Microbiology. 77 (11), 3224, 2020.
https://doi.org/10.1007/s00284... PMid:32876713.
76.
CETIN M., ISIK PEKKAN O., BILGE OZTURK G., SENYEL KURKCUOGLU M.A., KUCUKPEHLIVAN T., CABUK A. Examination of the change in the vegetation around the Kirka Boron mine site by using remote sensing techniques. Water, Air, & Soil Pollution. 233 (7), 254, 2022.
https://doi.org/10.1007/s11270....
77.
STEELE D.J., FRANKLIN D.J., UNDERWOOD G.J.C. Protection of cells from salinity stress by extracellular polymeric substances in diatom biofilms. Biofouling. 30 (8), 987, 2014.
https://doi.org/10.1080/089270... PMid:25268215 PMCid:PMC4706044.
78.
TRIVEDI P., MATTUPALLI C., EVERSOLE K., LEACH J.E. Enabling sustainable agriculture through understanding and enhancement of microbiomes. New Phytologist. 230 (6), 2129, 2021.
https://doi.org/10.1111/nph.17... PMid:33657660.
79.
SHRIVASTAVA P., KUMAR R. Soil salinity: A serious environmental issue and plant growth promoting bacteria as one of the tools for its alleviation. Saudi Journal of Biological Sciences. 22 (2), 123, 2015.
https://doi.org/10.1016/j.sjbs... PMid:25737642 PMCid:PMC4336437.
80.
DAS S. Genetic regulation, biosynthesis and applications of extracellular polysaccharides of the biofilm matrix of bacteria. Carbohydrate Polymers. 291, 119536, 2022.
https://doi.org/10.1016/j.carb... PMid:35698329.
81.
SANDHYA V., SK Z.A., GROVER M., REDDY G., VENKATESWARLU B. Alleviation of drought stress effects in sunflower seedlings by the exopolysaccharides producing Pseudomonas putida strain GAP-P45. Biology and Fertility of Soils. 46, 17, 2009.
https://doi.org/10.1007/s00374....
82.
O'CALLAGHAN M., BALLARD R.A., WRIGHT D. Soil microbial inoculants for sustainable agriculture: Limitations and opportunities. Soil Use and Management. 38 (3), 1340, 2022.
https://doi.org/10.1111/sum.12....
83.
HUNGRIA M., NOGUEIRA M.A., ARAUJO R.S. Co-inoculation of soybeans and common beans with rhizobia and azospirilla: strategies to improve sustainability. Biology and Fertility of Soils. 49, 791, 2013.
https://doi.org/10.1007/s00374....
84.
SHAH G., JAN M., AFREEN M., ANEES M., REHMAN S., DAUD M., MALOOK I., JAMIL M. Halophilic bacteria mediated phytoremediation of salt-affected soils cultivated with rice. Journal of Geochemical Exploration. 174, 59, 2017.
https://doi.org/10.1016/j.gexp....
85.
ROJAS-TAPIAS D., MORENO-GALVÁN A., PARDO-DÍAZ S., OBANDO M., RIVERA D., BONILLA R. Effect of inoculation with plant growth-promoting bacteria (PGPB) on amelioration of saline stress in maize (Zea mays). Applied Soil Ecology. 61, 264, 2012.
https://doi.org/10.1016/j.apso....
86.
VIMAL S.R., SINGH J.S., ARORA N.K., SINGS S. Soil-plant-microbe interactions in stressed agriculture management: a review. Pedosphere. 27 (2), 177, 2017.
https://doi.org/10.1016/S1002-....
87.
UPADHYAY S.K., SINGH J.S., SINGH D.P. Exopolysaccharide-Producing Plant Growth-Promoting Rhizobacteria Under Salinity Condition. Pedosphere. 21 (2), 214, 2011.
https://doi.org/10.1016/S1002-....
88.
FATIMA T., MISHRA I., VERMA R., ARORA N.K. Mechanisms of halotolerant plant growth promoting Alcaligenes sp. involved in salt tolerance and enhancement of the growth of rice under salinity stress. 3 Biotech. 10 (8), 361, 2020.
https://doi.org/10.1007/s13205... PMid:32832323 PMCid:PMC7392994.
89.
ABD EL-GHANY M.F., ATTIA M. Effect of Exopolysaccharide-Producing Bacteria and Melatonin on Faba Bean Production in Saline and Non-Saline Soil. Agronomy. 10 (3), 316, 2020.
https://doi.org/10.3390/agrono....
90.
ASHRAF M., HASNAIN S., BERGE O., MAHMOOD T. Inoculating wheat seedlings with exopolysaccharide-producing bacteria restricts sodium uptake and stimulates plant growth under salt stress. Biology and Fertility of Soils. 40, 157, 2004.
https://doi.org/10.1007/s00374....
91.
ANSARI F.A., AHMAD I., PICHTEL J. Growth stimulation and alleviation of salinity stress to wheat by the biofilm forming Bacillus pumilus strain FAB10. Applied Soil Ecology. 143, 45, 2019.
https://doi.org/10.1016/j.apso....
92.
KUMAR A., SINGH S., MUKHERJEE A., RASTOGI R.P., VERMA J.P. Salt-tolerant plant growth-promoting Bacillus pumilus strain JPVS11 to enhance plant growth attributes of rice and improve soil health under salinity stress. Microbiological Research. 242, 126616, 2021.
https://doi.org/10.1016/j.micr... PMid:33115624.
93.
VIVAS A., MARULANDA A., RUIZ-LOZANO J.M., BAREA J.M., AZCÓN R. Influence of a Bacillus sp. on physiological activities of two arbuscular mycorrhizal fungi and on plant responses to PEG-induced drought stress. Mycorrhiza. 13, 249, 2003.
https://doi.org/10.1007/s00572... PMid:14593518.
94.
HAQUE M.M., BISWAS M.S., MOSHARAF M.K., HAQUE M.A., ISLAM M.S., NAHAR K., ISLAM M.M., SHOZIB H.B., ISLAM M.M., FERDOUS E.E. Halotolerant biofilm-producing rhizobacteria mitigate seawater-induced salt stress and promote growth of tomato. Scientific Reports. 12 (1), 5599, 2022.
https://doi.org/10.1038/s41598... PMid:35379908 PMCid:PMC8980105.
95.
TALEBI ATOUEI M., POURBABAEE A.A., SHORAFA M. Alleviation of Salinity Stress on Some Growth Parameters of Wheat by Exopolysaccharide-Producing Bacteria. Iranian Journal of Science and Technology, Transactions A: Science. 43 (5), 2725, 2019.
https://doi.org/10.1007/s40995....
96.
SHULTANA R., KEE ZUAN A.T., YUSOP M.R., SAUD H.M. Characterization of salt-tolerant plant growth-promoting rhizobacteria and the effect on growth and yield of saline-affected rice. PLoS One. 15 (9), e0238537, 2020.
https://doi.org/10.1371/journa... PMid:32886707 PMCid:PMC7473536.
97.
SHULTANA R., TAN KEE ZUAN A., YUSOP M.R., MOHD SAUD H., AYANDA A.F. Effect of salt-tolerant bacterial inoculations on rice seedlings differing in salt-tolerance under saline soil conditions. Agronomy. 10 (7), 1030, 2020.
https://doi.org/10.3390/agrono....
98.
MAQSOOD A., SHAHID M., HUSSAIN S., MAHMOOD F., AZEEM F., TAHIR M., AHMED T., NOMAN M., MANZOOR I., BASIT F. Root colonizing Burkholderia sp. AQ12 enhanced rice growth and upregulated tillering-responsive genes in rice. Applied Soil Ecology. 157, 103769, 2021.
https://doi.org/10.1016/j.apso....
99.
HAN L., ZHANG H., XU Y., LI Y., ZHOU J. Biological characteristics and salt-tolerant plant growth-promoting effects of an ACC deaminase-producing Burkholderia pyrrocinia strain isolated from the tea rhizosphere. Archives of Microbiology. 203, 2279, 2021.
https://doi.org/10.1007/s00203... PMid:33644819.
100.
MAHMOUD O.M.B., HIDRI R., TALBI-ZRIBI O., TAAMALLI W., ABDELLY C., DJÉBALI N. Auxin and proline producing rhizobacteria mitigate salt-induced growth inhibition of barley plants by enhancing water and nutrient status. South African Journal of Botany. 128, 209, 2020.
https://doi.org/10.1016/j.sajb....
101.
MENESES C., GONÇALVES T., ALQUÉRES S., ROUWS L., SERRATO R., VIDAL M., BALDANI J. Gluconacetobacter diazotrophicus exopolysaccharide protects bacterial cells against oxidative stress in vitro and during rice plant colonization. Plant and Soil. 416, 133, 2017.
https://doi.org/10.1007/s11104....
102.
XIONG Y.-W., JU X.-Y., LI X.-W., GONG Y., XU M.-J., ZHANG C.-M., YUAN B., LV Z.-P., QIN S. Fermentation conditions optimization, purification, and antioxidant activity of exopolysaccharides obtained from the plant growth-promoting endophytic actinobacterium Glutamicibacter halophytocola KLBMP 5180. International Journal of Biological Macromolecules. 153, 1176, 2020.
https://doi.org/10.1016/j.ijbi... PMid:31756484.
103.
HIDRI R., MAHMOUD O.M.-B., ZORRIG W., MAHMOUDI H., SMAOUI A., ABDELLY C., AZCON R., DEBEZ A. Plant growth-promoting rhizobacteria alleviate high salinity impact on the halophyte Suaeda fruticosa by modulating antioxidant defense and soil biological activity. Frontiers in Plant Science. 13, 821475, 2022.
https://doi.org/10.3389/fpls.2... PMid:35720566 PMCid:PMC9199488.
104.
QURASHI A.W., SABRI A.N. Bacterial exopolysaccharide and biofilm formation stimulate chickpea growth and soil aggregation under salt stress. Brazilian Journal of Microbiology. 43 (3), 1183, 2012.
https://doi.org/10.1590/S1517-... PMid:24031943 PMCid:PMC3768896.
105.
ZHANG H., KIM M.-S., SUN Y., DOWD S.E., SHI H., PARÉ P.W. Soil bacteria confer plant salt tolerance by tissue-specific regulation of the sodium transporter HKT1. Molecular Plant-Microbe Interactions. 21 (6), 737, 2008.
https://doi.org/10.1094/MPMI-2... PMid:18624638.
106.
MASMOUDI F., ABDELMALEK N., TOUNSI S., DUNLAP C.A., TRIGUI M. Abiotic stress resistance, plant growth promotion and antifungal potential of halotolerant bacteria from a Tunisian solar saltern. Microbiological Research. 229, 126331, 2019.
https://doi.org/10.1016/j.micr... PMid:31521945.
107.
MISRA S., DIXIT V.K., MISHRA S.K., CHAUHAN P.S. Demonstrating the potential of abiotic stress-tolerant Jeotgalicoccus huakuii NBRI 13E for plant growth promotion and salt stress amelioration. Annals of Microbiology. 69 (4), 419, 2019.
https://doi.org/10.1007/s13213....
108.
KUSALE S.P., ATTAR Y.C., SAYYED R., EL ENSHASY H., HANAPI S.Z., ILYAS N., ELGORBAN A.M., BAHKALI A.H., MARRAIKI N. Inoculation of Klebsiella variicola alleviated salt stress and improved growth and nutrients in wheat and maize. Agronomy. 11 (5), 927, 2021.
https://doi.org/10.3390/agrono....
109.
SUN L., LEI P., WANG Q., MA J., ZHAN Y., JIANG K., XU Z., XU H. The Endophyte Pantoea alhagi NX-11 Alleviates Salt Stress Damage to Rice Seedlings by Secreting Exopolysaccharides. Frontiers in Microbiology. 10, 2020.
https://doi.org/10.3389/fmicb.... PMid:32038554 PMCid:PMC6987256.
110.
ALHOQAIL W.A. ACC-Deaminase producing Pseudomonas putida RT12 inoculation: A promising strategy for improving Brassica juncea tolerance to salinity stress. Notulae Botanicae Horti Agrobotanici Cluj-Napoca. 52 (1), 13550, 2024.
https://doi.org/10.15835/nbha5....
111.
VAISHNAV A., KUMARI S., JAIN S., VARMA A., TUTEJA N., CHOUDHARY D.K. PGPR‐mediated expression of salt tolerance gene in soybean through volatiles under sodium nitroprusside. Journal of Basic Microbiology. 56 (11), 1274, 2016.
https://doi.org/10.1002/jobm.2... PMid:27439917.
112.
LIU X., CHAI J., ZHANG Y., ZHANG C., LEI Y., LI Q., YAO T. Halotolerant rhizobacteria mitigate the effects of salinity stress on maize growth by secreting exopolysaccharides. Environmental and Experimental Botany. 204, 105098, 2022.
https://doi.org/10.1016/j.enve....
113.
DEKA P., GOSWAMI G., DAS P., GAUTOM T., CHOWDHURY N., BORO R.C., BAROOAH M. Bacterial exopolysaccharide promotes acid tolerance in Bacillus amyloliquefaciens and improves soil aggregation. Molecular Biology Reports. 46, 1079, 2019.
https://doi.org/10.1007/s11033... PMid:30554311.
114.
CHENG C., SHANG-GUAN W., HE L., SHENG X. Effect of Exopolysaccharide-Producing Bacteria on Water-Stable Macro-Aggregate Formation in Soil. Geomicrobiology Journal. 37 (8), 738, 2020.
https://doi.org/10.1080/014904....
115.
ISFAHANI F.M., TAHMOURESPOUR A., HOODAJI M., ATAABADI M., MOHAMMADI A. Characterizing the new bacterial isolates of high yielding exopolysaccharides under hypersaline conditions. Journal of Cleaner Production. 185, 922, 2018.
https://doi.org/10.1016/j.jcle....
116.
CHANRATANA M., HAN G.H., ROY CHOUDHURY A., SUNDARAM S., HALIM M.A., KRISHNAMOORTHY R., KANG Y., SA T. Assessment of Methylobacterium oryzae CBMB20 aggregates for salt tolerance and plant growth promoting characteristics for bio-inoculant development. AMB Express. 7, 1, 2017.
https://doi.org/10.1186/s13568... PMid:29164352 PMCid:PMC5698239.
117.
LIU X., EUSTERHUES K., THIEME J.R., CIOBOTA V., HÖSCHEN C., MUELLER C.W., KÜSEL K., KÖGEL-KNABNER I., RÖSCH P., POPP J.R. STXM and NanoSIMS investigations on EPS fractions before and after adsorption to goethite. Environmental Science & Technology. 4 (7), 3158, 2013.
https://doi.org/10.1021/es3039... PMid:23451805.