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Journal of Integrative Agriculture  2026, Vol. 25 Issue (11): 4621-4636    DOI: 10.1016/j.jia.2026.01.039
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Engineering of plant growth-promoting bacteria for sustainable nutrient management and agricultural sustainability

Jiajia Chen1*, Lanxiang Wang2, 3*, Xianglong Wei1, Muyu Xiong4, Sudarshanee Geekiyanage5, Meisam Zargar6, Lei Dai3, Gefei Hao1#, Moxian Chen1#

1 State Key Laboratory of Green Pesticide, Center for R&D of Fine Chemicals, Guizhou University, Guiyang 550025, China

2 State Key Laboratory for Development and Utilization of Forest Food Resources, State Key Laboratory of Tree Genetics and Breeding, the Southern Modern Forestry Collaborative Innovation Center, College of Life Sciences, Nanjing Forestry University, Nanjing 210037, China

3 State Key Laboratory of Quantitative Synthetic Biology, Shenzhen Institute of Synthetic Biology, Shenzhen Institutes of Advanced Technology, Chinese Academy of Sciences, Shenzhen 518055, China

4 Key Laboratory of Plant Resource Conservation and Germplasm Innovation in Mountainous Region (Ministry of Education), College of Life Sciences/Institute of Agro-bioengineering, Guizhou University, Guiyang 550025, China

5 Department of Agricultural Biology, Faculty of Agriculture University of Ruhuna, Mapalana Kamburupitiya 81100, Sri Lanka

6 Department of Agrobiotechnology, Institute of Agriculture, RUDN University, Moscow 117198, Russia

 Highlights 
● Engineering oxygen tolerance, nitrogen feedback control, and electron transfer pathways enhances BNF.
● Genetic editing of genes related to organic acid, phosphatase, and phytase synthesis strengthens the phosphorus and potassium solubilizing capabilities of PSB and KSB.
● Rationally engineered SynComs provide an effective strategy for improving nutrient use efficiency and biofertilizer performance.
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摘要  

氮(N)、磷(P)和钾(K)是植物生长发育过程中的关键营养元素。然而,土壤中的有效氮、磷、钾的供应常难以满足植物需求,需要通过施用化肥进行补充。长期过量施用化肥会使得土壤、空气和地下水受到污染,严重威胁农业的可持续发展。植物促生细菌(plant growth-promoting bacteria,PGPB)可通过固氮、溶磷、解钾等机制改善土壤肥力,促进植物生长,具有部分甚至完全替代化肥的潜力,从而推动可持续农业的发展。尽管许多 PGPB 菌株在实验条件下表现出良好的应用前景,但在田间环境中,其固氮、溶磷、解钾活性往往会受到限制。近年来,基因编辑技术的发展为精准增强PGPB功能、实现更高效且稳定的生物肥料生产提供了新方法。本文综述了非共生固氮菌(asymbiotic nitrogen fixing bacteria,ANFB)、溶磷菌(phosphorus solubilizing bacteria,PSB)和解钾菌(potassium solubilizing bacteria,KSB)提升土壤中可用性氮、磷、钾含量的机制,并总结了近年来通过基因编辑手段提升其养分转化效率的研究进展。同时,我们提出构建由基因改造菌株组成的合成微生物群落(SynCom),以进一步增强工程菌的促生效果并提高养分利用效率。



Abstract  

Nitrogen (N), phosphorus (P), and potassium (K) are plant growth and development nutrients with key functions in the life cycle of plants.  However, the levels of these nutrients in soil are often insufficient for plant uptake, so they must be supplemented using chemical fertilizers.  Excessive use of these fertilizers has polluted the soil, air, and groundwater, which is a serious threat to the sustainability of agriculture.  Plant growth-promoting bacteria (PGPB) facilitate plant growth by improving nitrogen fixation, as well as phosphorus and potassium solubilization.  These bacteria have the potential to partially or even entirely replace chemical fertilizers, an achievement that could improve soil fertility, boost crop production, and ensure sustainable agriculture.  Even though many PGPB strains can be used instead of chemical fertilizers, they have been shown to exhibit limited activity in N fixation and P or K solubilization when plants are grown in the field.  Recent advances in gene editing technologies now provide new avenues for enhancing these abilities in PGPB toward more efficient and consistent biofertilizer production.  This review discusses the mechanisms whereby asymbiotic nitrogen-fixing bacteria (ANFB) as well as phosphorus- and potassium-solubilizing bacteria (PSB and KSB) enhance plant growth, and it outlines the recent advances in gene editing efforts for improving microbial function.  We suggest creating a synthetic community (SynCom) comprised of genetically modified bacteria to enhance the PGP effects of engineered bacteria and promote efficient nutrient utilization.

Keywords:  PGPB       asymbiotic nitrogen fixation       phosphorus and potassium solubilization       gene editing       SynCom  
Received: 23 September 2025   Accepted: 23 December 2025 Online: 29 January 2026  
Fund: 

This work was financially supported by the National Key Research and Development Program of China (2024YFE0214300), the Guizhou Provincial Basic Research Program (Natural Science), China (ZK[2023]-099), the Post-Subsidy Fund Project of Guizhou International Science and Technology Cooperation Base, China (Qiankehe Platform GHJD (2025) 008), the National Natural Science Foundation of China (NSFC32572900), the Guizhou Provincial Postgraduate Research Fund, China (2024YJSKYJJ078), the RUDN University Strategic Academic Leadership Program, Russia, and the Central Government Guides Local Science and Technology Development Fund Projects, China (Qiankehezhongyindi (2024) 007).

About author:  Jiajia Chen, E-mail: chenjj220@163.com; Lanxiang Wang, E-mail: amelia0610@163.com; #Correspondence Gefei Hao, E-mail: gefei_hao@foxmail.com; Moxian Chen, E-mail: cmx2009920734@gmail.com * These authors contributed equally to this study.

Cite this article: 

Jiajia Chen, Lanxiang Wang, Xianglong Wei, Muyu Xiong, Sudarshanee Geekiyanage, Meisam Zargar, Lei Dai, Gefei Hao, Moxian Chen. 2026. Engineering of plant growth-promoting bacteria for sustainable nutrient management and agricultural sustainability. Journal of Integrative Agriculture, 25(11): 4621-4636.

Abdel L, Abu A, Kordrostami M, Abo B, Zakir A. 2020. Inoculation with Azospirillum lipoferum or Azotobacter chroococcum reinforces maize growth by improving physiological activities under saline conditions. Journal of Plant Growth Regulation, 39, 1293–1306.

Adediji A O, Ojo J A, Olowoake A A, Alabi K O, Atiri G I. 2024. Complete genome of Achromobacter xylosoxidans, a nitrogen-fixing bacteria from the rhizosphere of cowpea (Vigna unguiculata [L.] Walp) tolerant to cucumber mosaic virus infection. Current Microbiology, 81, 356.

Adhikary H, Sanghavi P B, Macwan S R, Archana G, Naresh Kumar G. 2014. Artificial citrate operon confers mineral phosphate solubilization ability to diverse fuorescent pseudomonads. PLoS ONE, 9, e107554.

An R, Moe L A. 2016. Regulation of pyrroloquinoline quinone-dependent glucose dehydrogenase activity in the model rhizosphere-dwelling bacterium Pseudomonas putida KT2440. Applied and Environmental Microbiology, 82, 4955–4964.

Bageshwar U K, Srivastava M, Pardha S P, Paul S, Gothandapani S, Jaat R S, Shankar P, Yadav R, Biswas D R, Kumar P A, Padaria J C, Mandal P K, Annapurna K, Das H K. 2017. An environmentally friendly engineered Azotobacter strain that replaces a substantial amount of urea fertilizer while sustaining the same wheat yield. Applied and Environmental Microbiology, 83, 00590–00617.

Barney B M, Eberhart L J, Ohlert J M, Knutson C M, Plunkett M H. 2015. Gene deletions resulting in increased nitrogen release by Azotobacter vinelandii: Application of a novel nitrogen biosensor. Applied and Environmental Microbiology, 81, 4316–4328.

Bautista C A, Antonio R B, Martínez V C M, Báez P A. 2019. Phosphate-solubilizing bacteria improve Agave angustifolia Haw. growth under field conditions. Journal of the Science of Food and Agriculture, 99, 6601–6607.

Bharwad K, Rajkumar S. 2019. Modulation of PQQ-dependent glucose dehydrogenase (mGDH and sGDH) activity by succinate in phosphate solubilizing plant growth promoting Acinetobacter sp. SK2. 3 Biotech, 10, 5.

Bloch S E, Clark R, Gottlieb S S, Wood L K, Shah N, Mak S M, Lorigan J G, Johnson J, Davis-Richardson A G, Williams L, Mckellar M, Soriano D, Petersen M, Horton A, Smith O, Wu L, Tung E, Broglie R, Tamsir A, Temme K. 2020. Biological nitrogen fixation in maize: optimizing nitrogenase expression in a root-associated diazotroph. Journal of Experimental Botany, 71, 4591–4603.

Bouizgarne B, Bakki M, Boutasknit A, Banane B, Ouarrat H, Maalem S, Amenzou A, Ghousmi A, Meddich A. 2023. Phosphate and potash solubilizing bacteria from moroccan phosphate mine showing antagonism to bacterial canker agent and inducing effective tomato growth promotion. Frontiers in Plant Science, 14, 970382.

Buch A, Archana G, Naresh Kumar G. 2010. Heterologous expression of phosphoenolpyruvate carboxylase enhances the phosphate solubilizing ability of fluorescent pseudomonads by altering the glucose catabolism to improve biomass yield. Bioresource Technology, 101, 679–687.

Buch A D, Archana G, Kumar G N. 2009. Enhanced citric acid biosynthesis in Pseudomonas fluorescens ATCC 13525 by overexpression of the Escherichia coli citrate synthase gene. Microbiology, 155, 2620–2629.

Bueno B M, Dixon R. 2019. Manipulating nitrogen regulation in diazotrophic bacteria for agronomic benefit. Biochemical Society Transactions, 47, 603–614.

Chemla Y, Sweeney C J, Wozniak C A, Voigt C A. 2025. Design and regulation of engineered bacteria for environmental release. Nature Microbiology, 10, 281–300.

Chen Q, Long C, Bao Y, Men X, Zhang Y, Cheng X. 2024. The dominant genera of nitrogenase (nifH) affects soil biological nitrogen fixation along an elevational gradient in the Hengduan mountains. Chemosphere, 347, 140722.

Chen Q, Zhao Q, Xie B, Lu X, Guo Q, Liu G, Zhou M, Tian J, Lu W, Chen K, Tian J, Liang C. 2024. Soybean (Glycine max) rhizosphere organic phosphorus recycling relies on acid phosphatase activity and specific phosphorus-mineralizing-related bacteria in phosphate deficient acidic soils. Journal of Integrative Agriculture, 23, 1685–1702.

Chen Y, Yang H, Shen Z, Ye J. 2022. Whole-genome sequencing and potassium-solubilizing mechanism of Bacillus aryabhattai SK1-7. Frontiers in Microbiology, 12, 722379.

Cheng Y, Narayanan M, Shi X, Chen X, Li Z, Ma Y. 2023. Phosphate-solubilizing bacteria: Their agroecological function and optimistic application for enhancing agro-productivity. Science of The Total Environment, 901, 166468.

Compant S, Cassan F, Kostić T, Johnson L, Brader G, Trognitz F, Sessitsch A. 2025. Harnessing the plant microbiome for sustainable crop production. Nature Reviews Microbiology, 23, 9–23.

Dai Z, Liu G, Chen H, Chen C, Wang J, Ai S, Wei D, Li D, Ma B, Tang C, Brookes P C, Xu J. 2019. Long-term nutrient inputs shift soil microbial functional profiles of phosphorus cycling in diverse agroecosystems. The ISME Journal, 14, 757–770.

Davies B T, Friedlingstein P. 2020. The global distribution of biological nitrogen fixation in terrestrial natural ecosystems. Global Biogeochemical Cycles, 34, e2019GB006387.

Matos G F, Rouws L F M, Simões A J L, Baldani J I. 2021. Evolution and function of nitrogen fixation gene clusters in sugarcane associated Bradyrhizobium strains. Environmental Microbiology, 23, 6148–6162.

Vries D W. 2021. Impacts of nitrogen emissions on ecosystems and human health: a mini review. Current Opinion in Environmental Science and Health, 21, 100249.

Delgado B M, Singh B K, Liu Y R, Sáez S T, Coleine C, Muñoz R M, Bastida F, Trivedi P. 2025. Integrating ecological and evolutionary frameworks for SynCom success. New Phytologist, 246, 1922–1933.

Elhaissoufi W, Ghoulam C, Barakat A, Zeroual Y, Bargaz A. 2022. Phosphate bacterial solubilization: a key rhizosphere driving force enabling higher P use efficiency and crop productivity. Journal of Advanced Research, 38, 13–28.

Emami S, Alikhani H A, Pourbabaei A A, Etesami H, Sarmadian F, Motessharezadeh B. 2019. Effect of rhizospheric and endophytic bacteria with multiple plant growth promoting traits on wheat growth. Environmental Science and Pollution Research, 26, 19804–19813.

Etesami H, Adl SM. 2020. Plant growth-promoting rhizobacteria (PGPR) and their action mechanisms in availability of nutrients to plants. In: Kumar M, Kumar V, Prasad R, eds., Phyto-Microbiome in Stress Regulation. Environmental and Microbial Biotechnology. Springer, Singapore. pp. 147–203.

Etesami H, Emami S, Alikhani H A. 2017. Potassium solubilizing bacteria (KSB): mechanisms, promotion of plant growth, and future prospects: a review. Journal of Soil Science and Plant Nutrition, 17, 897–911.

Etesami H, Jeong B R, Glick B R. 2021. Contribution of arbuscular mycorrhizal fungi, phosphate-solubilizing bacteria, and silicon to P uptake by plant. Frontiers in Plant Science, 12, 699618.

Florentino A P, Weijma J, Stams A J M, Sánchez-Andrea I. 2016. Ecophysiology and application of acidophilic sulfur-reducing microorganisms. Biotechnology of Extremophiles (Advances and Challenges), 1, 141–175.

Fraga R, Rodríguez H, González T. 2001. Transfer of the gene encoding the NapA acid phosphatase of Morganella morganii to a Burkholderia cepacia strain. Acta Biotechnologica, 21, 359–369.

Gandhi R, Prittesh P, Jinal H N, Chavan S M, Paul D, Amaresan N. 2023. Evaluation of the effect of potassium solubilizing bacterial strains on the growth of wheat (Triticum aestivum L.). Journal of Plant Nutrition, 46, 1479–1490.

Gao J, Feng P, Zhang J, Dong C, Wang Z, Chen M, Yu Z, Zhao B, Hou X, Wang H, Wu Z, Jemim R S, Yu H, Sun D, Jing P, Chen J, Song W, Zhang X, Zhou Z, Wu J. 2023. Enhancing maize’s nitrogen-fixing potential through ZmSBT3, a gene suppressing mucilage secretion. Journal of Integrative Plant Biology, 65, 2645–2659.

Gao J, Xu M, Uwiringiyimana E. 2025. Isolation of highly efficient potassium solubilizing bacteria and their effects on nutrient acquisition and growth promotion in tobacco seedlings. BMC Plant Biology, 25, 745.

Gao W, Zhao J, Guo X, Wang F, Chen X, Zhu Z, Ge T, Wang L, Kuzyakov Y, Wu J, Jia Z. 2024. Intensive N2 fixation accelerates microbial turnover in cropland soils. Science of The Total Environment, 916, 170081.

Ghani A, Rajan S, Lee A. 1994. Enhancement of phosphate rock solubility through biological processes. Soil Biology and Biochemistry, 26, 127–136.

Hara S, Morikawa T, Wasai S, Kasahara Y, Koshiba T, Yamazaki K, Fujiwara T, Tokunaga T, Minamisawa K. 2019. Identification of nitrogen-fixing Bradyrhizobium associated with roots of field-grown sorghum by metagenome and proteome analyses. Frontiers in Microbiology, 10, 407.

Harris D F, Yang Z, Dean D R, Seefeldt L C, Hoffman B M. 2018. Kinetic understanding of N2 reduction versus H2 evolution at the E4(4H) janus state in the three nitrogenases. Biochemistry, 57, 5706–5714.

Haskett T L, Tkacz A, Poole P S. 2020. Engineering rhizobacteria for sustainable agriculture. The ISME Journal, 15, 949–964.

He H, Zheng W, Xiao S, Gong L, Li H, Zhou K, Zhang L, Tu Q, Zhu Y Z, Zhang Y. 2024. Deciphering the nitrogen fixation gene cluster in Vibrio natriegens: a study on optimized expression and application of nitrogenase. Journal of Agricultural and Food Chemistry, 72, 12618–12629.

Huang L, Jia X, Zhang G, Shao M. 2017. Soil organic phosphorus transformation during ecosystem development: a review. Plant and Soil, 417, 17–42.

Illmer P, Schinner F. 1995. Solubilization of inorganic calcium phosphates-solubilization mechanisms. Soil Biology and Biochemistry, 27, 257–263.

Jansson J K, Mcclure R, Egbert R G. 2023. Soil microbiome engineering for sustainability in a changing environment. Nature Biotechnology, 41, 1716–1728.

Ji C, Liu Z, Hao L, Song X, Wang C, Liu Y, Li H, Li C, Gao Q, Liu X. 2020. Effects of Enterobacter cloacae HG-1 on the nitrogen-fixing community structure of wheat rhizosphere soil and on salt tolerance. Frontiers in Plant Science, 11, 01094.

Ji C, Tian H, Wang X, Song X, Ju R, Li H, Gao Q, Li C, Zhang P, Li J, Hao L, Wang C, Zhou Y, Xu R, Liu Y, Du J, Liu X. 2022. Bacillus subtilis HG-15, a halotolerant rhizoplane bacterium, promotes growth and salinity tolerance in wheat (Triticum aestivum). Biomed Research International, 2022, 9506227.

Jiao H, Wang R, Qin W, Yang J. 2024. Screening of rhizosphere nitrogen fixing, phosphorus and potassium solubilizing bacteria of Malus sieversii (Ldb.) Roem. and the effect on apple growth. Journal of Plant Physiology, 292, 154142.

Jini D, Ganga V S, Greeshma M B, Sivashankar R, Thirunavukkarasu A. 2024. Sustainable agricultural practices using potassium-solubilizing microorganisms (KSMs) in coastal regions: a critical review on the challenges and opportunities. Environment, Development and Sustainability, 26, 13641–13664.

Johnson R, Vishwakarma K, Hossen M S, Kumar V, Shackira A M, Puthur J T, Abdi G, Sarraf M, Hasanuzzaman M. 2022. Potassium in plants: growth regulation, signaling, and environmental stress tolerance. Plant Physiology and Biochemistry, 172, 56–69.

Jousset A, Becker J, Chatterjee S, Karlovsky P, Scheu S, Eisenhauer N. 2014. Biodiversity and species identity shape the antifungal activity of bacterial communities. Ecology, 95, 1184–1190.

Kaur T, Devi R, Kumar S, Sheikh I, Kour D, Yadav A N. 2022. Microbial consortium with nitrogen fixing and mineral solubilizing attributes for growth of barley (Hordeum vulgare L.). Heliyon, 8, e09326.

Kou C, Song F, Li D, Xu H, Zhang S, Yang W, Shi W, Gao Z. 2024. A necessary considering factor for crop resistance: precise regulation and effective utilization of beneficial microorganisms. New Crops, 1, 1000023.

Lee C C, Stiebritz M T, Hu Y, Ribbe M W. 2021. Assembly and function of nitrogenase. In: Moura J J G, Moura I, Maia L B, eds., Enzymes for Solving Humankind’s Problems. Springer, Cham. pp. 155–184.

Li G, Shi M, Wan W, Wang Z, Ji S, Yang F, Jin S, Zhang J. 2024. Maize endophytic plant growth-promoting bacteria Peribacillus simplex can alleviate plant saline and alkaline stress. International Journal of Molecular Sciences, 25, 10870.

Li J, Yang C, Jousset A, Yang K, Wang X, Xu Z, Yang T, Mei X, Zhong Z, Xu Y, Shen Q, Friman V P, Wei Z. 2023. Engineering multifunctional rhizosphere probiotics using consortia of Bacillus amyloliquefaciens transposon insertion mutants. eLife, 12, e90726.

Li P, Tian Y, Yang K, Tian M, Zhu Y, Chen X, Hu R, Qin T, Liu Y, Peng S, Yi Z, Liu Z, Ao H, Li J. 2024. Mechanism of microbial action of the inoculated nitrogen-fixing bacterium for growth promotion and yield enhancement in rice (Oryza sativa L.). Advanced Biotechnology, 2, 32.

Li X, Liu Q, Liu X, Shi H, Chen S. 2016. Using synthetic biology to increase nitrogenase activity. Microbial Cell Factories, 15, 43.

Li X, Zhao X, Dong X, Ma J, Shen R. 2021. Secretion of gluconic acid from Nguyenibacter sp. L1 is responsible for solubilization of aluminum phosphate. Frontiers in Microbiology, 12, 784025.

Li Y, Li Y, Zhang H, Wang M, Chen S. 2019. Diazotrophic Paenibacillus beijingensis BJ-18 provides nitrogen for plant and promotes plant growth, nitrogen uptake and metabolism. Frontiers in Microbiology, 10, 01119.

Li Z, Fang F, Wu L, Gao F, Li M, Li B, Wu K, Hu X, Wang S, Wei Z, Chen Q, Zhang M, Liu Z. 2024. The microbial community, nutrient supply and crop yields differ along a potassium fertilizer gradient under wheat-maize double-cropping systems. Journal of Integrative Agriculture, 23, 3592–3609.

Liu D, Liberton M, Yu J, Pakrasi H B, Bhattacharyya-Pakrasi M. 2018. Engineering nitrogen fixation activity in an oxygenic phototroph. mBio, 9, 01018–01029.

Liu Y, Evans S E, Friesen M L, Tiemann L K. 2022. Root exudates shift how N mineralization and N fixation contribute to the plant-available N supply in low fertility soils. Soil Biology and Biochemistry, 165, 108541.

Liu Y, Wang Y, Kong W, Liu W, Xie X, Wu X. 2020. Identification, cloning and expression patterns of the genes related to phosphate solubilization in Burkholderia multivorans WS-FJ9 under different soluble phosphate levels. AMB Express, 10, 108–119.

Mahala D M, Maheshwari H S, Yadav R K, Prabina B J, Bharti A, Reddy K K, Kumawat C, Ramesh A. 2020. Microbial transformation of nutrients in soil: an overview. Rhizosphere Microbes (Soil and Plant Functions), 23, 175–211.

Martin J S, Rigsbee J, Bueno B M, Mus F, Rubio L M, Einsle O, Peters J W, Dixon R, Dean D R, Santos P C. 2022. Overview of physiological, biochemical, and regulatory aspects of nitrogen fixation in Azotobacter vinelandii. Critical Reviews in Biochemistry and Molecular Biology, 57, 492–538.

Mrnjavac N, Degli E M, Mizrahi I, Martin W F, Allen J F. 2024. Three enzymes governed the rise of O2 on earth. Biochimica et Biophysica Acta (BBA)-Bioenergetics, 1865, 149495.

Mukherjee A, Singh S, Gaurav A K, Chouhan G K, Jaiswal D K, De Araujo Pereira A P, Passari A K, Abdel-Azeem A M, Verma J P. 2022. Harnessing of phytomicrobiome for developing potential biostimulant consortium for enhancing the productivity of chickpea and soil health under sustainable agriculture. Science of the Total Environment, 836, 155550.

Mullaney E, Ullah A. 2007. Phytases: attributes, catalytic mechanisms and applications. In: Turner B L, Richardson A E, Mullaney E J, eds., Inositol Phosphates: Linking Agriculture and the Environment. CABI, United Kingdom. pp. 97–110.

Narehood S M, Cook B D, Srisantitham S, Eng V H, Shiau A A, Mcguire K L, Britt R D, Herzik M A, Tezcan F A. 2025. Structural basis for the conformational protection of nitrogenase from O2. Nature, 637, 991–997.

Nawaz A, Qamar Z U, Marghoob M U, Imtiaz M, Imran A, Mubeen F. 2023. Contribution of potassium solubilizing bacteria in improved potassium assimilation and cytosolic K+/Na+ ratio in rice (Oryza sativa L.) under saline-sodic conditions. Frontiers in Microbiology, 14, 1196024.

Neal A L, Blackwell M, Akkari E, Guyomar C, Clark I, Hirsch P R. 2018. Phylogenetic distribution, biogeography and the effects of land management upon bacterial non-specific acid phosphatase gene diversity and abundance. Plant and Soil, 427, 175–189.

Pan L, Cai B. 2023. Phosphate-solubilizing bacteria: advances in their physiology, molecular mechanisms and microbial community effects. Microorganisms, 11, 2904.

Pang Z, Mao X, Zhou S, Yu S, Liu G, Lu C, Wan J, Hu L, Xu P. 2023. Microbiota-mediated nitrogen fixation and microhabitat homeostasis in aerial root-mucilage. Microbiome, 11, 85.

Parveen F A M, Hari O, Bholanath S, Pushpendra K. 2019. Free Living Nitrogen Fixation and their Response to Agricultural Crops. Apple Academic Press, United States. pp.173–200.

Pérez G A, Jimenez V E, Gies E J, Salinero L A, Yang Z, Einsle O, Seefeldt L C, Dean D R. 2021. Specificity of NifEN and VnfEN for the assembly of nitrogenase active site cofactors in Azotobacter vinelandii. mBio, 12, 1110–1128.

Petrović M, Janakiev T, Grbić M L, Unković N, Stević T, Vukićević S, Dimkić I. 2023. Insights into endophytic and rhizospheric bacteria of five sugar beet hybrids in terms of their diversity, plant-growth promoting, and biocontrol properties. Microbial Ecology, 87, 19.

Poole R K, Hill S. 1997. Respiratory protection of nitrogenase activity in Azotobacter vinelandii - roles of the terminal oxidases. Bioscience Reports, 17, 303–317.

Qiu L, Fan M, Li Y, Yang Q, Gou X, Kong Y, Zhou S, Chen G, Xie J, Chen Y, Liu L, Tang Y, Tu L. 2023. Soil phosphorus drives variation in diazotrophic communities in a subtropical nitrogen-rich forest. Forest Ecology and Management, 544, 121164.

Rajawat M V S, Ansari W A, Singh D, Singh R. 2019. Potassium solubilizing bacteria (KSB). Microbial Interventions in Agriculture and Environment, 3, 189–209.

Rao V R. 1978. Effect of carbon sources on asymbiotic nitrogen fixation in a paddy soil. Soil Biology and Biochemistry, 10, 319–321.

Rawat P, Das S, Shankhdhar D, Shankhdhar S C. 2021. Phosphate-solubilizing microorganisms: mechanism and their role in phosphate solubilization and uptake. Journal of Soil Science and Plant Nutrition, 21, 49–68.

Raza W, Jiang G, Eisenhauer N, Huang Y, Wei Z, Shen Q, Kowalchuk G A, Jousset A. 2024. Microbe-induced phenotypic variation leads to overyielding in clonal plant populations. Nature Ecology and Evolution, 8, 392–399.

Raza W, Wang J, Jousset A, Friman V P, Mei X, Wang S, Wei Z, Shen Q. 2020. Bacterial community richness shifts the balance between volatile organic compound-mediated microbe–pathogen and microbe–plant interactions. Proceedings of the Royal Society (B: Biological Sciences), 287, 20200403.

Robson R L. 1979. Characterization of an oxygen-stable nitrogenase complex isolated from Azotobacter chroococcum. Biochemical Journal, 181, 569–575.

Romero M A, Aroca R. 2023. A non-K+-solubilizing PGPB (Bacillus megaterium) increased K+ deprivation tolerance in Oryza sativa seedlings by up-regulating root K+ transporters. Plant Physiology and Biochemistry, 196, 774–782.

Rossi F, Carles L, Donnadieu F, Batisson I, Artigas J. 2021. Glyphosate-degrading behavior of five bacterial strains isolated from stream biofilms. Journal of Hazardous Materials, 420, 126651.

Ryu M H, Zhang J, Toth T, Khokhani D, Geddes B A, Mus F, Garcia C A, Peters J W, Poole P S, Ané J M, Voigt C A. 2020. Control of nitrogen fixation in bacteria that associate with cereals. Nature Microbiology, 5, 314–330.

Schlesier J, Rohde M, Gerhardt S, Einsle O. 2016. A conformational switch triggers nitrogenase protection from oxygen aamage by shethna protein II (FeSII). Journal of the American Chemical Society, 138, 239–247.

Sebilo M, Mayer B, Nicolardot B, Pinay G, Mariotti A. 2013. Long-term fate of nitrate fertilizer in agricultural soils. Proceedings of the National Academy of Sciences of the United States of America, 110, 18185–18189.

Setten L, Soto G, Mozzicafreddo M, Fox A R, Lisi C, Cuccioloni M, Angeletti M, Pagano E, Díaz-Paleo A, Ayub N D. 2013. Engineering Pseudomonas protegens Pf-5 for nitrogen fixation and its application to improve plant growth under nitrogen-deficient conditions. PLoS ONE, 8, e63666.

Shao J, Liu Y, Xie J, Štefanič P, Lv Y, Fan B, Mandic M I, Zhang R, Shen Q, Xu Z. 2022. Annulment of bacterial antagonism improves plant beneficial activity of a Bacillus velezensis consortium. Applied and Environmental Microbiology, 88, 00240-22.

Sharma R, Sindhu S S, Glick B R. 2024. Potassium solubilizing microorganisms as potential biofertilizer: a sustainable climate-resilient approach to improve soil fertility and crop production in agriculture. Journal of Plant Growth Regulation, 43, 2503–2535.

Shen L, Wu X, Zeng Q, Liu H. 2016. Regulation of soluble phosphate on the ability of phytate mineralization and β-propeller phytase gene expression of Pseudomonas fluorescens JZ-DZ1, a phytate-mineralizing rhizobacterium. Current Microbiology, 73, 915–923.

Shi S, Richardson A E, O’callaghan M, Deangelis K M, Jones E E, Stewart A, Firestone M K, Condron L M. 2011. Effects of selected root exudate components on soil bacterial communities. FEMS Microbiology Ecology, 77, 600–610.

Shi W, Zhao H, Chen Y, Wang J, Han B, Li C, Lu J, Zhang L. 2021. Organic manure rather than phosphorus fertilization primarily determined asymbiotic nitrogen fixation rate and the stability of diazotrophic community in an upland red soil. Agriculture, Ecosystems and Environment, 319, 107535.

Shi Z Y, Guo Y X, Wang Y Y, Yan X, Guo X, Lei Z H, Niu J P, Liang J P, Li Z Y. 2024. Nitrogen-fixing bacteria promote growth and bioactive components accumulation of Astragalus mongholicus by regulating plant metabolism and rhizosphere microbiota. BMC Microbiology, 24, 261.

Shrivastava M, Srivastava P C, D’souza S F. 2018. Phosphate-solubilizing microbes: diversity and phosphates solubilization mechanism. In: Meena V, ed., Role of Rhizospheric Microbes in Soil. Springer, Singapore. pp. 137–165.

Shulse C N, Chovatia M, Agosto C, Wang G, Hamilton M, Deutsch S, Yoshikuni Y, Blow M J. 2019. Engineered root bacteria release plant-available phosphate from phytate. Applied and Environmental Microbiology, 85, e01210-19.

Si J, Su X, Jin Z, Duan S. 2025. Uncovering essential lncRNAs through transcriptome-scale CRISPR-Cas13 screening. Advanced Biotechnology, 3, 27.

Singh B, Boukhris I, Pragya, Kumar V, Yadav A N, Farhat-Khemakhem A, Kumar A, Singh D, Blibech M, Chouayekh H, Alghamdi O A. 2020. Contribution of microbial phytases to the improvement of plant growth and nutrition: a review. Pedosphere, 30, 295–313.

Singh B K, Hu H, Macdonald C A, Xiong C. 2025. Microbiome-facilitated plant nutrient acquisition. Cell Host and Microbe, 33, 869–881.

Singh P, Singh R K, Li H, Guo D, Sharma A, Lakshmanan P, Malviya M K, Song X, Solanki M K, Verma K K, Yang L, Li Y. 2021. Diazotrophic bacteria Pantoea dispersa and Enterobacter asburiae promote sugarcane growth by inducing nitrogen uptake and defense-related gene expression. Frontiers in Microbiology, 11, 600417.

Smercina D N, Bowsher A W, Evans S E, Friesen M L, Eder E K, Hoyt D W, Tiemann L K. 2021. Switchgrass rhizosphere metabolite chemistry driven by nitrogen availability. Phytobiomes Journal, 5, 88–96.

Solomon J B, Lee C C, Liu Y A, Duffin C, Ribbe M W, Hu Y. 2024. Ammonia synthesis via an engineered nitrogenase assembly pathway in Escherichia coli. Nature Catalysis, 7, 1130–1141.

Su C, Jiang S, Dai J. 2025. Development of a versatile synthetic microbe for the concurrent degradation of organic pollutants. Advanced Biotechnology, 3, 16.

Sun F, Ou Q, Wang N, Guo Z X, Ou Y, Li N, Peng C. 2020. Isolation and identification of potassium-solubilizing bacteria from Mikania micrantha rhizospheric soil and their effect on M. micrantha plants. Global Ecology and Conservation, 23, e01141.

Takimoto R, Tatemichi Y, Aoki W, Kosaka Y, Minakuchi H, Ueda M, Kuroda K. 2022. A critical role of an oxygen-responsive gene for aerobic nitrogenase activity in Azotobacter vinelandii and its application to Escherichia coli. Scientific Reports, 12, 4182.

Tatemichi Y, Nakahara T, Ueda M, Kuroda K. 2021. Construction of recombinant Escherichia coli producing nitrogenase-related proteins from Azotobacter vinelandii. Bioscience, Biotechnology, and Biochemistry, 85, 2209–2216.

Tian S, Xu Y, Zhong Y, Qiao Y, Wang D, Wu L, Yang X, Yang M, Wu Z. 2024. Exploring the organic acid secretion pathway and potassium solubilization ability of Pantoea vagans ZHS-1 for enhanced rice growth. Plants, 13, 1945.

Tovilla C, Eiteman M, Mark A. 2020. Engineered citrate synthase alters acetate accumulation in Escherichia coli. Metabolic Engineering, 61, 171–180.

Upadhyay S K, Rajput V D, Kumari A, Espinosa S D, Menendez E, Minkina T, Dwivedi P, Mandzhieva S. 2023. Plant growth-promoting rhizobacteria: a potential bio-asset for restoration of degraded soil and crop productivity with sustainable emerging techniques. Environmental Geochemistry and Health, 45, 9321–9344.

Van D A, Zamora P, Delaux P M, Heitmann C, Jayaraman D, Rajasekar S, Graham D, Maeda J, Gibson D, Schwartz K D, Berry A M, Bhatnagar S, Jospin G, Darling A, Jeannotte R, Lopez J, Weimer B C, Eisen J A, Shapiro H Y, Ané J M, et al. 2018. Nitrogen fixation in a landrace of maize is supported by a mucilage-associated diazotrophic microbiota. PLoS Biology, 16, e2006352.

Vitousek P M, Porder S, Houlton B Z, Chadwick O A. 2010. Terrestrial phosphorus limitation: mechanisms, implications, and nitrogen-phosphorus interactions. Ecological Applications, 20, 5–15.

Wagh J, Chanchal K, Sonal S, Praveena B, Archana G, Kumar G N. 2016. Inoculation of genetically modified endophytic Herbaspirillum seropedicae Z67 endowed with gluconic and 2-ketogluconic acid secretion, confers beneficial effects on rice (Oriza sativa) plants. Plant and Soil, 409, 51–64.

Wagh J, Shah S, Bhandari P, Archana G, Kumar G N. 2014. Heterologous expression of pyrroloquinoline quinone (pqq) gene cluster confers mineral phosphate solubilization ability to Herbaspirillum seropedicae Z67. Applied Microbiology and Biotechnology, 98, 5117–5129.

Wan W, Qin Y, Wu H, Zuo W, He H, Tan J, Wang Y, He D. 2020. Isolation and characterization of phosphorus solubilizing bacteria with multiple phosphorus sources utilizing capability and their potential for lead immobilization in soil. Frontiers in Microbiology, 11, 00752.

Wang J, Li R, Zhang H, Wei G, Li Z. 2020. Beneficial bacteria activate nutrients and promote wheat growth under conditions of reduced fertilizer application. BMC Microbiology, 20, 38.

Wang L, Zhang L, Liu Z, Zhao D, Liu X, Zhang B, Xie J, Hong Y, Li P, Chen S, Dixon R, Li J. 2013. A minimal nitrogen fixation gene cluster from Paenibacillus sp. WLY78 enables expression of active nitrogenase in Escherichia coli. PLoS Genetics, 9, e1003865.

Wang Z, Zhang H, Liu L, Li S, Xie J, Xue X, Jiang Y. 2022. Screening of phosphate-solubilizing bacteria and their abilities of phosphorus solubilization and wheat growth promotion. BMC Microbiology, 22, 296.

Wen A, Havens K L, Bloch S E, Shah N, Higgins D A, Davis-Richardson A G, Sharon J, Rezaei F, Mohiti-Asli M, Johnson A, Abud G, Ane J M, Maeda J, Infante V, Gottlieb S S, Lorigan J G, Williams L, Horton A, Mckellar M, Soriano D, et al. 2021. Enabling biological nitrogen fixation for cereal crops in fertilized fields. ACS Synthetic Biology, 10, 3264–3277.

Wu J, Liu S, Zhang H, Chen S, Si J, Liu L, Wang Y, Tan S, Du Y, Jin Z, Xie J, Zhang D. 2025. Flavones enrich rhizosphere Pseudomonas to enhance nitrogen utilization and secondary root growth in Populus. Nature Communications, 16, 1461.

Xiao D, Xiao L, Che R, Tan Y, Liu X, Yang R, Zhang W, He X, Wang K. 2020. Phosphorus but not nitrogen addition significantly changes diazotroph diversity and community composition in typical karst grassland soil. Agriculture, Ecosystems and Environment, 301, 106987.

Xu M, Fan L, Li A, Liu Q, Yu G, Wang S, Zhang B, Ye Q, Mo J, Zheng M. 2024. Plant and microbial carbon are important drivers of free-living nitrogen fixation in tropical forest soils: a new discovery of carbon-driven nitrogen input. Geophysical Research Letters, 51, e2024GL111238.

Xu X, Dinesen C, Pioppi A, Kovács Á T, Lozano-Andrade C N. 2025. Composing a microbial symphony: synthetic communities for promoting plant growth. Trends in Microbiology, 33, 738–751.

Yang J, Xie X, Wang X, Dixon R, Wang Y P. 2014. Reconstruction and minimal gene requirements for the alternative iron-only nitrogenase in Escherichia coli. Proceedings of the National Academy of Sciences of the United States of America, 111, e3718-e3725.

You M, Fang S, Macdonald J, Xu J, Yuan Z. 2020. Isolation and characterization of Burkholderia cenocepacia CR318, a phosphate solubilizing bacterium promoting corn growth. Microbiological Research, 233, 126395.

Yu H, Wu X, Zhang G, Zhou F, Harvey P R, Wang L, Fan S, Xie X, Li F, Zhou H, Zhao X, Zhang X. 2022. Identification of the phosphorus-solubilizing bacteria strain JP233 and its effects on soil phosphorus leaching loss and crop growth. Frontiers in Microbiology, 13, 892533.

Zahra S T, Tariq M, Abdullah M, Zafar M, Yasmeen T, Shahid M S, Zaki H E M, Ali A. 2023. Probing the potential of salinity-tolerant endophytic bacteria to improve the growth of mungbean [Vigna radiata (L.) Wilczek]. Frontiers in Microbiology, 14, 1149004.

Zeffa D M, Perini L J, Silva M B, De Sousa N V, Scapim C A, Oliveira A L M D, Amaral Júnior A T D, Azeredo Gonçalves L S. 2019. Azospirillum brasilense promotes increases in growth and nitrogen use efficiency of maize genotypes. PLoS ONE, 14, e0215332.

Zehr J P. 2011. Nitrogen fixation by marine cyanobacteria. Trends in Microbiology, 19, 162–173.

Zeng Q, Tang L, Zhang Y, Shao Y, Wu W, Wang J, Ding X, Han X, Muhammad B. 2023. Isolation and characterization of phosphate-solubilizing bacteria from rhizosphere of poplar on road verge and their antagonistic potential against various phytopathogens. BMC Microbiology, 23, 221.

Zhang B, Jiang H, Zheng G, Zhang Z, Wang T, Chen Q, Chen X, Lu X, Liang C, Tian J. 2025. Genistein exudation drives spatial root allocation and heterogeneous microbial communities to enhance phosphorus acquisition in soybean-maize intercropping. Plant, Cell and Environment, 48, 7297–7312.

Zhang H, Han L, Jiang B, Long C. 2021. Identification of a phosphorus-solubilizing Tsukamurella tyrosinosolvens strain and its effect on the bacterial diversity of the rhizosphere soil of peanuts growth-promoting. World Journal of Microbiology and Biotechnology, 37, 109.

Zhang L, Zhang M, Huang S, Li L, Gao Q, Wang Y, Zhang S, Huang S, Yuan L, Wen Y, Liu K, Yu X, Li D, Zhang L, Xu X, Wei H, He P, Zhou W, Philippot L, Ai C. 2022. A highly conserved core bacterial microbiota with nitrogen-fixation capacity inhabits the xylem sap in maize plants. Nature Communications, 13, 3361.

Zhang T, Yan Y, He S, Ping S, Alam K M, Han Y, Liu X, Lu W, Zhang W, Chen M, Xiang W, Wang X, Lin M. 2012. Involvement of the ammonium transporter AmtB in nitrogenase regulation and ammonium excretion in Pseudomonas stutzeri A1501. Research in Microbiology, 163, 332–339.

Zhang Y, Hu T, Wang H, Jin H, Liu Q, Chen Z, Xie Z. 2022a. Nitrogen content and C/N ratio in straw are the key to affect biological nitrogen fixation in a paddy field. Plant and Soil, 481, 535–546.

Zhang Y, Ye C, Su Y, Peng W, Lu R, Liu Y, Huang H, He X, Yang M, Zhu S. 2022b. Soil acidification caused by excessive application of nitrogen fertilizer aggravates soil-borne diseases: evidence from literature review and field trials. Agriculture, Ecosystems and Environment, 340, 108176.

Zhao Y, Liang H, Zhang J, Chen Y, Dhital Y P, Zhao T, Wang Z. 2024. Isolation and characterization of potassium-solubilizing rhizobacteria (KSR) promoting cotton growth in saline–sodic regions. Microorganisms, 12, 1474.

Zheng B, Zhang D, Wang Y, Hao X, Wadaan M, Hozzein W N, Peñuelas J, Zhu Y, Yang X. 2019. Responses to soil pH gradients of inorganic phosphate solubilizing bacteria community. Scientific Reports, 9, 25.

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