Alan E. Richardson

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

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.

Prospects for using soil microorganisms to improve the acquisition of phosphorus by plants

Microorganisms play an important role in the acquisition and transfer of nutrients in soil. For phosphorus (P), soil microorganisms are involved in a range of processes that affect P transformation and thus influence the subsequent availability of P (as phosphate) to plant roots. In particular, microorganisms can solubilize and mineralize P from inorganic and organic pools of total soil P. In addition, microorganisms may effectively increase the surface area of roots. Also, the microbial biomass itself contains a large pool of immobilized P that potentially is available to plants. Given that most soils are deficient in plant-available P and that P fertilizer represents a significant cost for agricultural production throughout the world, there is interest in using soil microorganisms as inoculants to mobilize P from poorly available sources in soil. Although potential clearly exists for developing such inoculants, their widespread application remains limited by a poor understanding of microbial ecology and population dynamics in soil, and by inconsistent performance over a range of environments. Furthermore, promotion of growth of plants in soil, as a consequence of microbial inoculation, may not necessarily be associated with characteristics such as P solubilization, which are manifest under laboratory conditions.

Soil Microorganisms Mediating Phosphorus Availability Update on Microbial Phosphorus

Microorganisms are integral to the soil phosphorus (P) cycle and as such play an important role in mediating the availability of P to plants. Understanding the microbial contribution to plant P nutrition and opportunities for manipulating specific microorganisms to enhance P availability in soil has

Soil microorganisms mediating phosphorus availability

Microorganisms are integral to the soil phosphorus (P) cycle and as such play an important role in mediating the availability of P to plants. Understanding the microbial contribution to plant P nutrition and opportunities for manipulating specific microorganisms to enhance P availability in soil has

Plant mechanisms to optimise access to soil phosphorus.

Phosphorus (P) is an important nutrient required for plant growth and its management in soil is critical to ensure sustainable and profitable agriculture that has minimal impact on the environment. Although soils may contain a large amount of total P, only a small proportion is immediately available to plants. Australian soils often have low availability of P for plant growth and P-based fertilisers are, therefore, commonly used to correct P deficiency and to maintain productivity. For many soils, the sustained use of P fertiliser has resulted in an accumulation of total P, a proportion of which is in forms that are poorly available to most plants. The efficiency with which different P fertilisers are used in agricultural systems depends on their capacity to supply P in a soluble form that is available for plant uptake (i.e. as orthophosphate anions). In addition to fertiliser source, the availability of P in soil is influenced to a large extent by physico-chemical and biological properties of the soil. Plant access to soil P is further affected by root characteristics (e.g. rate of growth, specific root length, and density and length of root hairs) and biochemical processes that occur at the soil–root interface. The ability of roots to effectively explore soil, the release of exudates (e.g. organic anions and phosphatases) from roots that influence soil P availability, and the association of roots with soil microorganisms such as mycorrhizal fungi are particularly important. These processes occur as a natural response of plants to P deficiency and, through better understanding, may provide opportunities for improving plant access to soil and fertiliser P in conventional and organic agricultural systems.

Strategies and agronomic interventions to improve the phosphorus-use efficiency of farming systems

Phosphorus (P)-deficiency is a significant challenge for agricultural productivity on many highly P-sorbing weathered and tropical soils throughout the world. On these soils it can be necessary to apply up to five-fold more P as fertiliser than is exported in products. Given the finite nature of global P resources, it is important that such inefficiencies be addressed. For low P-sorbing soils, P-efficient farming systems will also assist attempts to reduce pollution associated with P losses to the environment. P-balance inefficiency of farms is associated with loss of P in erosion, runoff or leaching, uneven dispersal of animal excreta, and accumulation of P as sparingly-available phosphate and organic P in the soil. In many cases it is possible to minimise P losses in runoff or erosion. Uneven dispersal of P in excreta typically amounts to ~5% of P-fertiliser inputs. However, the rate of P accumulation in moderate to highly P-sorbing soils is a major contributor to inefficient P-fertiliser use. We discuss the causal edaphic, plant and microbial factors in the context of soil P management, P cycling and productivity goals of farms. Management interventions that can alter P-use efficiency are explored, including better targeted P-fertiliser use, organic amendments, removing other constraints to yield, zone management, use of plants with low critical-P requirements, and modified farming systems. Higher productivity in low-P soils, or lower P inputs in fertilised agricultural systems can be achieved by various interventions, but it is also critically important to understand the agroecology of plant P nutrition within farming systems for improvements in P-use efficiency to be realised.

Components of organic phosphorus in soil extracts that are hydrolysed by phytase and acid phosphatase

Abstract Extracts were prepared from soil using water, 50 mM citric acid (pH ∼2.3) or 0.5 M NaHCO3 (pH 8.5), and were incubated with excess phytase from Aspergillus niger to determine the amounts of labile P. Two A. niger phytase preparations were used: (1) a purified form which exhibited a narrow substrate specificity and high specific activity against phytate; and (2) a commercial preparation (Sigma) with activity against a broad range of P compounds. A comparatively large proportion (up to 79%, or 5.7 μg g–1 soil) of the organic P (Po) extracted with citric acid was hydrolysed by the commercial phytase, while between 28% and 40% (up to 3.1 μg g–1 soil) was hydrolysed using purified phytase. By comparison, only small quantities of the Po in water and NaHCO3 soil extracts were enzyme labile. While extractable Po was increased both with increasing concentrations of citric acid (up to 50 mM) and increasing pH (pH 2.3–6.0), enzyme-labile P increased only with citric acid concentration. The labile component of Po in citric acid extracts from soils with contrasting fertiliser histories indicated that enzyme-labile Po is a relatively large soil P pool and is potentially an important source of P for plants.

Utilization of phosphorus by pasture plants supplied with myo-inositol hexaphosphate is enhanced by the presence of soil micro-organisms

A range of pasture grass (Danthonia richardsonii and Phalaris aquatica) and legume (Medicago polymorpha, M. sativa, Trifolium repens and T. subterraneum) species showed limited capacity to obtain phosphorus (P) from inositol hexaphosphate (IHP), when grown in either sterile agar (pH 5.0 or 5.5) or sand-vermiculite media (pH 5.0). The total P content of shoots from IHP-supplied plants grown in agar was between 20% and 34% of that for seedlings supplied with an equivalent amount of P as inorganic phosphate (Pi), while in sand-vermiculite, the total P content of IHP-grown plants was between 5 and 10% of control plants. The poor ability of plants to utilize P from IHP resulted in significantly lower tissue P concentrations and, in general, reduced plant dry weight accumulation. In contrast, the P nutrition of plants supplied with IHP was significantly improved by inoculating media with either a cultured population of total soil micro-organisms or with a specific isolate of Pseudomonas sp., selected for its ability to release phosphate from IHP (strain CCAR59; Richardson and Hadobas, 1997 Can. J. Micro. 43, 509-516). In agar and sand-vermiculite media, respectively, the P content of IHP-grown plants increased with inoculation by up to 3.9- and 6.8-fold, such that the dry weight and P content of the plant material were equivalent to those observed for control plants supplied with Pi. However, the response to inoculation was dependent on the growth medium and the source of micro-organisms used. In sand-vermiculite, the cultured population of soil micro-organisms was effective when IHP was supplied at an equivalent level of Pi required for maximum plant growth. By comparison, inoculation of plants with the Pseudomonas strain was only effective at very high levels of IHP supply (×36), whereas in agar a response to inoculation occurred at all levels of IHP. The ability of pasture plants to acquire P from phytate was, therefore, influenced by the availability of IHP substrate, which was further affected by the presence of soil micro-organisms. Our results show that in addition to having an effect on the sorption characteristics of the growth media, soil micro-organisms also provided a source of phytase for the dephosphorylation of phytate for subsequent utilization of Pi by plants.

Effects of selected root exudate components on soil bacterial communities

Low-molecular-weight organic compounds in root exudates play a key role in plant-microorganism interactions by influencing the structure and function of soil microbial communities. Model exudate solutions, based on organic acids (OAs) (quinic, lactic, maleic acids) and sugars (glucose, sucrose, fructose), previously identified in the rhizosphere of Pinus radiata, were applied to soil microcosms. Root exudate compound solutions stimulated soil dehydrogenase activity and the addition of OAs increased soil pH. The structure of active bacterial communities, based on reverse-transcribed 16S rRNA gene PCR, was assessed by denaturing gradient gel electrophoresis and PhyloChip microarrays. Bacterial taxon richness was greater in all treatments than that in control soil, with a wide range of taxa (88-1043) responding positively to exudate solutions and fewer (<24) responding negatively. OAs caused significantly greater increases than sugars in the detectable richness of the soil bacterial community and larger shifts of dominant taxa. The greater response of bacteria to OAs may be due to the higher amounts of added carbon, solubilization of soil organic matter or shifts in soil pH. Our results indicate that OAs play a significant role in shaping soil bacterial communities and this may therefore have a significant impact on plant growth.

Promoter Analysis of the Barley Pht1;1 Phosphate Transporter Gene Identifies Regions Controlling Root Expression and Responsiveness to Phosphate Deprivation

Previous studies have shown that the promoter from the barley (Hordeum vulgare) phosphate transporter gene, HvPht1;1, activates high levels of expression in rice (Oryza sativa) roots and that the expression level was induced by up to 4-fold in response to phosphorus (P) deprivation. To identify promoter regions controlling gene regulation specificities, successive promoter truncations were made and attached to reporter genes. Promoters of between 856 and 1,400 nucleotides activated gene expression in a number of cell types but with maximal expression in trichoblast (root hair) cells. For shorter promoters the trichoblast specificity was lost, but in other tissues the distribution pattern was unchanged. The low P induction response was unaffected by promoter length. Domain exchange experiments subsequently identified that the region between −856 and −547 nucleotides (relative to the translational start) is required for epidermal cell expression. A second region located between 0 and −195 nucleotides controls root-tip expression. The HvPht1;1 promoter contains one PHO-like motif and three motifs similar to the dicot P1BS element. Analysis of promoters from which the PHO-like element was eliminated (by truncation) showed no change in the gene induction response to P deficiency. In contrast, mutation of the P1BS elements eliminated any induction of gene expression in response to low P. An internal HvPht1;1 promoter fragment, incorporating a single P1BS element, had an increased response to P deprivation in comparison with the unmodified promoter (containing three elements). Together these findings further our understanding of the regulation of the HvPht1;1 gene and provide direct evidence for a functional role of the P1BS element in the expression of P-regulated genes.