Phosphorus is the second important key element after nitrogen as a mineral nutrient in terms of quantitative plant requirement. Although abundant in soils, in both organic and inorganic forms, its availability is restricted as it occurs mostly in insoluble forms. The P content in average soil is about 0.05% (w/w) but only 0.1% of the total P is available to plant because of poor solubility and its fixation in soil (Illmer and Schinner, Soil Biol Biochem 27:257-263, 1995). An adequate supply of phosphorus during early phases of plant development is important for laying down the primordia of plant reproductive parts. It plays significant role in increasing root ramification and strength thereby imparting vitality and disease resistance capacity to plant. It also helps in seed formation and in early maturation of crops like cereals and legumes. Poor availability or deficiency of phosphorus (P) markedly reduces plant size and growth. Phosphorus accounts about 0.2 - 0.8% of the plant dry weight. To satisfy crop nutritional requirements, P is usually added to soil as chemical P fertilizer, however synthesis of chemical P fertilizer is highly energy intensive processes, and has long term impacts on the environment in terms of eutrophication, soil fertilility depletion, carbon footprint. Moreover, plants can use only a small amount of this P since 75–90% of added P is precipitated by metal–cation complexes, and rapidly becomes fixed in soils. Such environmental concerns have led to the search for sustainable way of P nutrition of crops. In this regards phosphate-solubilizing microorganisms (PSM) have been seen as best eco-friendly means for P nutrition of crop. Although, several bacterial (pseudomonads and bacilli) and fungal strains (Aspergilli and Penicillium) have been identified as PSM their performance under in situ conditions is not reliable and therefore needs to be improved by using either genetically modified strains or co-inoculation techniques. This review focuses on the diversity of PSM, mechanism of P solubilization, role of various phosphatases, impact of various factors on P solubilization, the present and future scenario of their use and potential for application of this knowledge in managing a sustainable environmental system.

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Phosphate solubilizing microbes: sustainable approach for managing phosphorus deficiency in agricultural soils

In addition to fixing atmospheric nitrogen, some bacterial isolates can also solubilize insoluble phosphates, further contributing to plant growth. The objectives of this study were the following: isolate, select, and identify nodulating bacteria in the cowpea that are efficient not only in biological nitrogen fixation (BNF) but also in the solubilization of insoluble inorganic phosphates; identify and quantify the organic acids produced; and establish the relationship between those acids and the solubilizing capacity. The bacteria were captured from two soils containing high concentrations of insoluble phosphorus from the cities of Lavras and Patos de Minas, using the cowpea [Vigna unguiculata (L.) Walp.] as bait. We obtained 78 strains, which were characterized according to their cultural attributes in culture medium 79 with the strains UFLA 03-84, INPA 03-11B, and BR3267 (approved by the Ministry of Livestock and Supply Agriculture—MAPA, as inoculants for the cowpea) and Burkholderia cepacia (LMG1222T), which was used as a positive control for phosphate solubilization. Strains that were selected for their efficiency in both processes were identified by 16S rDNA sequence analysis. We evaluated the symbiotic efficiency (BNF) in a greenhouse and the solubilization efficiency of CaHPO4, Al(H2PO4)3, and FePO4.2H2O in solid and liquid GELP media. Strains that excelled at the solubilization of these phosphate sources were also evaluated for the production of the following organic acids: oxalic, citric, gluconic, lactic, succinic, and propionic. The presence of Acinetobacter, Bacillus, Firmicutes, Microbacterium, Paenibacillus, and Rhizobium was detected by 16S rDNA sequencing and analysis. Bacterial strains obtained from cowpea nodules varied greatly in the efficiency of their BNF and phosphate solubilization processes, especially in the strains UFLA 03-09, UFLA 03-10, UFLA 03-12, and UFLA 03-13, which were more efficient in both processes. More strains were able to solubilize insoluble inorganic calcium and iron phosphates in liquid medium than in solid medium. The production of organic acids was related to the solubilization of CaHPO4 and FePO4.2H2O for some strains, and the type and concentration of the acid influenced this process. These are the first results obtained with bacterial isolates from tropical soils in which the production of organic acids was detected and quantified to examine the solubilization of insoluble inorganic phosphates.

Biological nitrogen fixation and phosphate solubilization by bacteria isolated from tropical soils

The use of phosphate-solubilizing fungi is a promising biotechnological strategy in the management of phosphorus (P) fertilization, as it enables the utilization of rock phosphates (RP) or the recovery of P fixed in soil particles. The objective of our study was to evaluate fungal isolates for mechanisms of solubilization of P-bearing compounds, such as AlPO4, FePO4, Ca3(PO4)2, Araxa RP, and Catalao RP. Four fungal isolates obtained from Brazilian soils were characterized in liquid media: Aspergillus niger FS1, Penicillium canescens FS23, Eupenicillium ludwigii FS27, and Penicillium islandicum FS30. A. niger FS1 was the only isolate able to solubilize all of the P sources, solubilizing 71, 36, 100, and 14 % of the P from AlPO4, FePO4, Ca3(PO4)2, and RPs, respectively. Medium acidification was an effective solubilization mechanism, particularly for Ca3(PO4)2. The other P sources were mainly solubilized through organic acids produced by the fungi. Oxalic acid, produced exclusively by A. niger FS1, and citric acid were decisive factors in the solubilization of AlPO4 and FePO4. Penicillium isolates produced more gluconic acid than A. niger FS1 in all treatments. However, this higher production did not result in higher solubilization for any of the P sources, showing that gluconic acid contributes little to the solubilization of the P sources evaluated. The higher capacity of medium acidification and the production of organic acids with stronger metal-complexation activity are characteristics that confer to A. niger FS1 a wider action on insoluble P sources. Consequently, this isolate qualifies as a promising candidate for application in the management of P fertilization.

Mechanisms of phosphate solubilization by fungal isolates when exposed to different P sources

Rice husk biochar impacts soil phosphorous availability, phosphatase activities and bacterial community characteristics in three different soil types

Most soils contain large reserves of total phosphorus (P), but its fixation and precipitation with soil constituents cause a major P-deficiency and severely restrict the growth and yield of plants. The use of chemical P-fertilizers is obviously the best means to circumvent P-deficiency, but their use is always limited due to its spiraling cost. In order to increase the availability of P and to reduce the use of chemical fertilizers, solubilization of insoluble P by phosphate-solubilizing microorganisms has provided an alternative to chemical phosphatic fertilizer. Besides P, these organisms promote the growth of plants by N2 fixation, enhancement of other plant nutrients, synthesizing phytohormones, suppressing plant diseases (bio-control) and reducing the toxicity of ethylene through 1-aminocyclopropane-1carboxylate (ACC) deaminase. In this chapter, attention is paid to understanding the fundamental and molecular basis as to how precisely these microbes, notably bacteria and fungi, help plants to grow better in P-deficient soils. Effective use of such microbes is likely to result in an ideal cropping system with a lesser impact on the environment through decreased application of chemical fertilizers.

Recent Advances in Plant Growth Promotion by Phosphate-Solubilizing Microbes

Two of 187 fungal isolates (Aspergillus niger 1B and 6A) displaying superior phosphate (P) solubilization and hydrolytic enzyme secretion were studied using P forms of calcium (Ca-P), iron (Fe-P), and aluminum (Al-P). Phosphate solubilization in a sucrose-basal salt (SB) broth was increased and pH decreased by both isolates. In Ca-P medium, solubilization for 6A was approximately 322 μg P mL−1 and pH decreased by 4.2 units to 2.3 in 72 h. However, when pH value of the SB broth was lowered to 2.5 using HCl, 65.3  ±  0.4 μg mL−1 of P was released from Ca-P, whereas trace amounts of P were released from Fe-P and Al-P. Both isolates displayed enhanced Al-P solubilization using NH4Cl rather than KNO3 as the N source; final pH values were not significantly different. With Ca-P, gluconic acid was predominantly produced by 1B and 6A, whereas oxalic acid predominated with Fe-P and Al-P. Addition of gluconic acid (final concentration of 8.5 μmol mL−1) to Ca-P-supplemented SB lowered pH (2.9) and solubilized phosphate (146.0 ± 1.0 μg mL−1). Similarly, addition of oxalic acid (final concentration 6.6 μmol mL−1) to Ca-P- and Fe-P-amended media solubilized P (60.2 ± 0.9 and 21.6 ± 2.1 μg mL−1, respectively), although these quantities were significantly lower than those detected in unamended SB. The presence of unidentified P solubilized compound(s) in the dialyzed (MW>500) supernatant warrants further study. In pot experiments, significant increases in plant (Brassica chinensis Linn.) dry weight and N and P contents were observed with the addition of isolate 6A, when a small amount of organic fertilizer together with either rock phosphate (South African apatite) or Ca-P served as the main P sources.

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Yu-Lin Kuo

Papers

Solubilization of inorganic phosphates and plant growth promotion by Aspergillus niger