Mechanisms and applications of plant growth promoting rhizobacteria: Current perspective

During the last 35 years of studies of Azospirillum–plant interaction, over 20 proposals were suggested for the mechanism of action by which Azospirillum spp., the most intensively studied plant growth-promoting bacteria, enhances plant growth. The proposals include a single phytohormone activity, multiple phytohormones, nitrogen fixation, assortments of small-sized molecules and enzymes, enhanced membrane activity, proliferation of the root system, enhanced water and mineral uptake, mobilization of minerals, mitigation of environmental stressors of plants, and direct and indirect biological control of numerous phytopathogens. By volume, the largest number of published information involves hormonal activities, nitrogen fixation, and root proliferation. After analyzing the accumulated knowledge, it was concluded that this versatile genus possesses a large array of potential mechanisms by which it can effect plant growth. Consequently, this review proposes the “Multiple Mechanisms Theory,” based on the assumption that there is no single mechanism involved in promotion of plant growth by Azospirillum, but a combination of a few or many mechanisms in each case of inoculation. These may vary according to the plant species, the Azospirillum strain, and environmental conditions when the interaction occurred. The effect can be cumulative, an “additive hypothesis” (proposed before), where the effects of small mechanisms operating at the same time or consecutively create a larger final effect on plant. Additionally, the observed effect on plant growth can be the result of a tandem or a cascade of mechanisms in which one mechanism stimulates another, yielding enhanced plant growth, such as the plausible relations among phytohormones, nitric oxide, membrane activities, and proliferation of roots. Finally, the growth promotion can also be a combination of unrelated mechanisms that operate under environmental or agricultural conditions needed by the crop at particular locations, such as mitigating stress (salt, drought, toxic compounds, adverse environment), and the need for biological control of or reducing pathogenic microflora.

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How the Plant Growth-Promoting Bacterium Azospirillum Promotes Plant Growth—A Critical Assessment

Nitrogen is generally considered one of the major limiting nutrients in plant growth. The biological process responsible for reduction of molecular nitrogen into ammonia is referred to as nitrogen fixation. A wide diversity of nitrogen-fixing bacterial species belonging to most phyla of the Bacteria domain have the capacity to colonize the rhizosphere and to interact with plants. Leguminous and actinorhizal plants can obtain their nitrogen by association with rhizobia or Frankia via differentiation on their respective host plants of a specialized organ, the root nodule. Other symbiotic associations involve heterocystous cyanobacteria, while increasing numbers of nitrogen-fixing species have been identified as colonizing the root surface and, in some cases, the root interior of a variety of cereal crops and pasture grasses. Basic and advanced aspects of these associations are covered in this review.

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Nitrogen-fixing bacteria associated with leguminous and non-leguminous plants

The rhizosphere is the soil-plant root interphase and in practice consists of the soil adhering to the root besides the loose soil surrounding it. Plant growth-promoting rhizobacteria (PGPR) are potential agents for the biological control of plant pathogens. A biocontrol strain should be able to protect the host plant from pathogens and fulfill the requirement for strong colonization. Numerous compounds that are toxic to pathogens, such as HCN, phenazines, pyrrolnitrin, and pyoluteorin as well as, other enzymes, antibiotics, metabolites and phytohormones are the means by which PGPR act, just as quorum sensing and chemotaxis which are vital for rhizosphere competence and colonization. The presence of root exudates has a pronounced effect on the rhizosphere where they serve as an energy source, promoting growth and influencing the root system for the rhizobacteria. In certain instances they have products that inhibit the growth of soil-borne pathogens to the advantage of the plant root. A major source of concern is reproducibility in the field due to the complex interaction between the plant (plant species), microbe and the environment (soil fertility and moisture, day length, light intensity, length of growing season, and temperature). This review listed most of the documented PGPR genera and discussed their exploitation.

Beneficial bacteria of agricultural importance

Plant growth-promoting rhizobacteria act as biostimulants in horticulture

Bacteria of the genus Azospirillum are considered to be plant-growth promoting bacteria (PGPR) and stimulate plant growth directly either by synthesising phyto-hormones or by promoting nutrition by the process of biological nitrogen fixation (BNF). Although this genus extensively studied, the effects of inoculation and the possible BNF contribution of the Azospirillum amazonense specie are very scarce. The aim of this study was to isolate, characterise and evaluate auxin production and nitrogenase activity of this species and to select, by inoculation of rice plants, promising isolates based on their ability to improve plant growth, yield and the BNF contribution. One hundred and ten isolates obtained from rice were characterised and grouped according to colony features. Forty-two isolates, confirmed as A. amazonense by the fluorescent in situ hybridization (FISH) technique, were tested for auxin production and nitrogenase activity in vitro. Subsequently plant growth promotion related to plant nutrition effect was evaluated, in vitro and in greenhouse experiments. The BNF contribution to plant growth was evaluated using the 15N isotope dilution technique. All A. amazonense strains tested produced indoles, but only 10% of them showed high production, above 1.33 μM mg protein−1. The nitrogenase activity also was variable and only 9% of isolates showed high nitrogenase activity and the majority (54%) exhibited a low potential. The inoculation of selected strains in rice under gnotobiotic conditions reduced the growth of root and aerial part when compared to the control, showing the negative effects of excess of phytohormone accumulation in the medium. However, in the greenhouse experiment, inoculation of strains of A. amazonense increased grain dry matter accumulation (7 to 11.6%), the number of panicles (3 to 18.6%) and nitrogen accumulation at grain maturation (3.5 to 18.5%). BNF contributions up to 27% were observed for A. amazonense Y2 (wild type strain). The data presented here is the first report that the PGPR effect of A. amazonense for rice plants grown under greenhouse conditions is mainly due the BNF contribution as measured by 15N isotope dilution technique, in contrast to the hormonal effect observed with other Azospirillum species studied.

https://link.springer.com/content/pdf/10.1007%2Fs11104-008-9840-9.pdf

Azospirillum amazonense inoculation: effects on growth, yield and N2 fixation of rice (Oryza sativa L.)

VIDEIRA, S. S.; OLIVEIRA, A. L. M.; BALDANI, J. I. and BALDANI, V. L. D. Identity and genetic variability of isolates of nitrogen-fi xing bacteria associated with rice (Oryza sativa). Revista Universidade Rural: Serie Ciencias da Vida, Seropedica, RJ: EDUR, v. 24, n.2, p. 35-40, jul.-dez., 2004. The study had for objective to evaluateis 14 bacterial isolates, obtained from roots and leaves of the rice cultivars IR42 and IAC 4440. These isolates presented the following properties: presence of an aerotactic pellicle when grown in JNFb semi-solid N-free medium, curved rods, Gram negative, microaerophilic, motile with 5 polar fl agella in one pole, and elevated nitrogenase activity (determined by acetylene reduction). The hybridization with the oligonucleotide probes (FISH) indicate that the isolates belong to the a subdivision of the class Proteobacteria. The results obtained with restriction analysis of rDNA amplifi ed (ARDRA) showed low diversity within the isolates and the partial sequencing of 16S rDNA region showed that they belong to the genus Sphingomonas.

André Luiz Martinez de Oliveira

Papers

Azospirillum amazonense inoculation: effects on growth, yield and N2 fixation of rice (Oryza sativa L.)

IDENTIFICAÇÃO E AVALIAÇÃO DA DIVERSIDADE DE NOVOS ISOLADOS DE BACTÉRIAS DIAZOTRÓFICAS EM PLANTAS DE ARROZ (Oryza sativa)