Plant growth-promoting rhizobacteria (PGPR) are the rhizosphere bacteria that can enhance plant growth by a wide variety of mechanisms like phosphate solubilization, siderophore production, biological nitrogen fixation, rhizosphere engineering, production of 1-Aminocyclopropane-1-carboxylate deaminase (ACC), quorum sensing (QS) signal interference and inhibition of biofilm formation, phytohormone production, exhibiting antifungal activity, production of volatile organic compounds (VOCs), induction of systemic resistance, promoting beneficial plant-microbe symbioses, interference with pathogen toxin production etc. The potentiality of PGPR in agriculture is steadily increased as it offers an attractive way to replace the use of chemical fertilizers, pesticides and other supplements. Growth promoting substances are likely to be produced in large quantities by these rhizosphere microorganisms that influence indirectly on the overall morphology of the plants. Recent progress in our understanding on the diversity of PGPR in the rhizosphere along with their colonization ability and mechanism of action should facilitate their application as a reliable component in the management of sustainable agricultural system. The progress to date in using the rhizosphere bacteria in a variety of applications related to agricultural improvement along with their mechanism of action with special reference to plant growth-promoting traits are summarized and discussed in this review.

Plant growth-promoting rhizobacteria (PGPR): emergence in agriculture

Mechanisms and applications of plant growth promoting rhizobacteria: Current perspective

The rhizosphere is a complex environment where roots interact with physical, chemical and biological properties of soil. Structural and functional characteristics of roots contribute to rhizosphere processes and both have significant influence on the capacity of roots to acquire nutrients. Roots also interact extensively with soil microorganisms which further impact on plant nutrition either directly, by influencing nutrient availability and uptake, or indirectly through plant (root) growth promotion. In this paper, features of the rhizosphere that are important for nutrient acquisition from soil are reviewed, with specific emphasis on the characteristics of roots that influence the availability and uptake of phosphorus and nitrogen. The interaction of roots with soil microorganisms, in particular with mycorrhizal fungi and non-symbiotic plant growth promoting rhizobacteria, is also considered in relation to nutrient availability and through the mechanisms that are associated with plant growth promotion.

Acquisition of phosphorus and nitrogen in the rhizosphere and plant growth promotion by microorganisms

Soil bacteria are very important in biogeochemical cycles and have been used for crop production for decades. Plant–bacterial interactions in the rhizosphere are the determinants of plant health and soil fertility. Free-living soil bacteria beneficial to plant growth, usually referred to as plant growth promoting rhizobacteria (PGPR), are capable of promoting plant growth by colonizing the plant root. PGPR are also termed plant health promoting rhizobacteria (PHPR) or nodule promoting rhizobacteria (NPR). These are associated with the rhizosphere, which is an important soil ecological environment for plant–microbe interactions. Symbiotic nitrogen-fixing bacteria include the cyanobacteria of the genera Rhizobium, Bradyrhizobium, Azorhizobium, Allorhizobium, Sinorhizobium and Mesorhizobium. Free-living nitrogen-fixing bacteria or associative nitrogen fixers, for example bacteria belonging to the species Azospirillum, Enterobacter, Klebsiella and Pseudomonas, have been shown to attach to the root and efficiently colonize root surfaces. PGPR have the potential to contribute to sustainable plant growth promotion. Generally, PGPR function in three different ways: synthesizing particular compounds for the plants, facilitating the uptake of certain nutrients from the soil, and lessening or preventing the plants from diseases. Plant growth promotion and development can be facilitated both directly and indirectly. Indirect plant growth promotion includes the prevention of the deleterious effects of phytopathogenic organisms. This can be achieved by the production of siderophores, i.e. small metal-binding molecules. Biological control of soil-borne plant pathogens and the synthesis of antibiotics have also been reported in several bacterial species. Another mechanism by which PGPR can inhibit phytopathogens is the production of hydrogen cyanide (HCN) and/or fungal cell wall degrading enzymes, e.g., chitinase and s-1,3-glucanase. Direct plant growth promotion includes symbiotic and non-symbiotic PGPR which function through production of plant hormones such as auxins, cytokinins, gibberellins, ethylene and abscisic acid. Production of indole-3-ethanol or indole-3-acetic acid (IAA), the compounds belonging to auxins, have been reported for several bacterial genera. Some PGPR function as a sink for 1-aminocyclopropane-1-carboxylate (ACC), the immediate precursor of ethylene in higher plants, by hydrolyzing it into α-ketobutyrate and ammonia, and in this way promote root growth by lowering indigenous ethylene levels in the micro-rhizo environment. PGPR also help in solubilization of mineral phosphates and other nutrients, enhance resistance to stress, stabilize soil aggregates, and improve soil structure and organic matter content. PGPR retain more soil organic N, and other nutrients in the plant–soil system, thus reducing the need for fertilizer N and P and enhancing release of the nutrients.

Soil beneficial bacteria and their role in plant growth promotion: a review

Plant growth-promoting rhizobacteria (PGPR) are naturally occurring soil bacteria that aggressively colonize plant roots and benefit plants by providing growth promotion. Inoculation of crop plants with certain strains of PGPR at an early stage of development improves biomass production through direct effects on root and shoots growth. Inoculation of ornamentals, forest trees, vegetables, and agricultural crops with PGPR may result in multiple effects on early-season plant growth, as seen in the enhancement of seedling germination, stand health, plant vigor, plant height, shoot weight, nutrient content of shoot tissues, early bloom, chlorophyll content, and increased nodulation in legumes. PGPR are reported to influence the growth, yield, and nutrient uptake by an array of mechanisms. They help in increasing nitrogen fixation in legumes, help in promoting free-living nitrogen-fixing bacteria, increase supply of other nutrients, such as phosphorus, sulphur, iron and copper, produce plant hormones, enhance other beneficial bacteria or fungi, control fungal and bacterial diseases and help in controlling insect pests. There has been much research interest in PGPR and there is now an increasing number of PGPR being commercialized for various crops. Several reviews have discussed specific aspects of growth promotion by PGPR. In this review, we have discussed various bacteria which act as PGPR, mechanisms and the desirable properties exhibited by them.

Plant Growth Promoting Rhizobacteria: A Critical Review

Screening of free-living rhizospheric bacteria for their multiple plant growth promoting activities

A total of 21 bacterial isolates (Azotobacter sp., 10 and fluorescent Pseudomonas sp., 11) were isolated from different rhizospheric soils in the vicinity of Aligarh city and characterized as per standard methods. These isolates were further tested for the production of indole acetic acid (IAA) in a medium with 0, 1, 2 and 5 mg/ml of tryptophan. A low amount (2.68-10.80 mg/ml) of IAA production was recorded by Azotobacter strains without tryptophan addition. Seven Azotobacter isolates showed high level (7.3 to 32.8 mg/ml) production of IAA at 5 mg/ml of tryptophan while at 1 and 2 mg/ml the production was in the range of 1.47 to 11.88 and 5.99 to 24.8 mg/ml, respectively. Production of IAA in fluorescent Pseudomonas isolates increased with an increase in tryptophan concentration from 1 to 5 mg/ml in the majority of isolates. In the presence of 5mg/ ml of tryptophan, 5 isolates of Pseudomonas produced high levels (41.0 to 53.2 mg/ml) of IAA while 6 other isolates produced IAA in the range of 23.4 to 36.2 mg/ml. Production of IAA was further confirmed by extraction of crude IAA from 3 isolates of Azotobacter (Azs1, Azs6 and Azs9 ) and three isolates of Pseudomonas (Ps1, Ps4 and Ps7) and subsequent TLC analysis. A specific spot from the extracted IAA preparation was found corresponding with the standard spot of IAA with same Rf value. Pseudomonas isolates (Ps1, Ps4 and Ps7) showed inhibitory effects on the growth of root elongation of Sesbania aculeata and Vigna radiata at all concentrations of tryptophan compared to the control. However, the isolates of Azotobacter (Azs1, Azs6 and Azs9) demonstrated stimulatory effects on both plants. Increasing the concentration of tryptophan from 1 mg/ml to 5 mg/ml resulted in decreased growth in both S. aculeata and V. radiata. On a comparative basis isolate Azs9 was most promising in promoting plant growth. On the other hand, high concentration of exogenous tryptophan could exhibit toxic effects on plant growth.

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Indole Acetic Acid Production by the Indigenous Isolates of Azotobacter and Fluorescent Pseudomonas in the Presence and Absence of Tryptophan

Microbes and Microbial Technology

Two isolates belonging to the predominant genera Aspergillus and Rhizopus isolated from agricultural field treated with sewage/ industrial effluents were selected for the biosorption potential evaluation of Cr and Cd. Pretreated, dead biomass of above fungi was used for bioadsorption experiment at pH value 4.5 with the biomass, 1-5 mg in a 100 ml metal solution of different concentration (2, 4, 6 and 8 mM) with a contact time of 18 hrs and agitation, 120 rpm. Bioadsorption of Cr ranged from 6.20-9.5 mg/g of dry mass at one or other initial metal concentrations by Aspergillus and Rhizopus sp. The bioadsorption of Cd was ranged from 2.3-8.21mg/g. On the comparative basis Rhizopus sp. could bioadsorbed higher concentration of both metals as compared to Aspergillus sp. Bioadsorption of Cd and Cr was influenced by initial metal concentration and nature of organism. The findings revealed that fungi of metal polluted sites showed higher metal tolerance and bioadsorption capacity of chromium and cadmium. Journal of Applied Sciences and Environmental Management Vol. 9(1) 2005: 123-126

Heavy Metal Biosorption potential of Aspergillus and Rhizopus sp. isolated from Wastewater treated soil

Roots serve a multitude of functions in plants including anchorage, acquisition of nutrients and water, and production of exudates with growth regulatory properties. The root–soil interface, or rhizosphere, is the site of greatest biological and chemical activity within the soil matrix. Plant growth-promoting rhizobacteria (PGPR) are known to influence plant health by controlling plant pathogens or via direct enhancement of plant development in the laboratory and in greenhouse experiments. Unfortunately, however, results in the field have been less consistent. The colonization of roots by inoculated bacteria is an important step in the interaction between beneficial bacteria and the host plant. However, colonization is a complex phenomenon influenced by many biotic and abiotic parameters, some of which are only now apparent. Monitoring fate and metabolic activity of microbial inoculants as well as their impact on rhizosphere and soil microbial communities are needed to guarantee safe and reliable application, independent of whether they are genetically modified or not. The first and most crucial prerequisite for effective use of PGPRs is that strain identity and activity are continuously confirmed. A combination of both classical and molecular techniques must be perfected for more effective monitoring of inoculants strain (both genetically modified and unmodified) after release into the soil. Recent developments in techniques for studying rhizobacterial communities and detection and tracking systems of inoculated bacteria are important in future application and assessment of effectiveness and consistent performance of microbial inoculants in crop production and protection.

Farah Ahmad

Papers

Screening of free-living rhizospheric bacteria for their multiple plant growth promoting activities

Indole Acetic Acid Production by the Indigenous Isolates of Azotobacter and Fluorescent Pseudomonas in the Presence and Absence of Tryptophan

Microbes and Microbial Technology

Heavy Metal Biosorption potential of Aspergillus and Rhizopus sp. isolated from Wastewater treated soil

Rhizosphere and Root Colonization by Bacterial Inoculants and Their Monitoring Methods: A Critical Area in PGPR Research