Contents
I. Introduction  424
II. The phosphorus conundrum 424
III. Adaptations to low P 424
IV. Uptake of P  424
V. P deficiency alters root development and function 426
VI. P deficiency modifies carbon metabolism 431
VII. Acid phosphatase 436
VIII. Genetic regulation of P responsive genes 437
IX. Improving P acquisition 439
X. Synopsis  440


Summary
Phosphorus (P) is limiting for crop yield on > 30% of the world's arable land and, by some estimates, world resources of inexpensive P may be depleted by 2050. Improvement of P acquisition and use by plants is critical for economic, humanitarian and environmental reasons. Plants have evolved a diverse array of strategies to obtain adequate P under limiting conditions, including modifications to root architecture, carbon metabolism and membrane structure, exudation of low molecular weight organic acids, protons and enzymes, and enhanced expression of the numerous genes involved in low-P adaptation. These adaptations may be less pronounced in mycorrhizal-associated plants. The formation of cluster roots under P-stress by the nonmycorrhizal species white lupin (Lupinus albus), and the accompanying biochemical changes exemplify many of the plant adaptations that enhance P acquisition and use. Physiological, biochemical, and molecular studies of white lupin and other species response to P-deficiency have identified targets that may be useful for plant improvement. Genomic approaches involving identification of expressed sequence tags (ESTs) found under low-P stress may also yield target sites for plant improvement. Interdisciplinary studies uniting plant breeding, biochemistry, soil science, and genetics under the large umbrella of genomics are prerequisite for rapid progress in improving nutrient acquisition and use in plants.

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Phosphorus acquisition and use: critical adaptations by plants for securing a nonrenewable resource

P is an important plant macronutrient, making up about 0.2% of a plant's dry weight. It is a component of key molecules such as nucleic acids, phospholipids, and ATP, and, consequently, plants cannot grow without a reliable supply of this nutrient. Pi is also involved in controlling key enzyme

Phosphorus Uptake by Plants: From Soil to Cell

Phosphate (Pi) is considered to be one of the least available plant nutrients in the soil. High-affinity Pi transporters are generally accepted as entry points for Pi in the roots. The physiological, genetic, molecular and biochemical analysis of phosphate starvation response mechanisms highlight the ability of plants to adapt and thrive under phosphate limiting conditions. These responses help them enhance the availability of Pi, increase its uptake and improve the use-efficiency of Pi within a plant. Enhanced ability to acquire Pi appears to be regulated at the level of transcription of high-affinity phosphate transporters. These transporters are encoded by a family of small number of genes having characteristic tissue and organ associated expression patterns. Many of them are strongly induced during phosphate deficiency thus providing plants with enhanced ability to acquire and transfer phosphate. In addition, plants also activate biochemical mechanisms that could lead to increased acquisition of phosphate from both inorganic and organic phosphorus sources in the soil. Furthermore, altered root morphology and mycorrhizal symbiosis further enhance the ability of plants to acquire Pi. Interestingly most of these responses appear to be coordinated by changes in cellular phosphate levels. It is becoming apparent that phosphate acquisition and utilization are associated with activation or inactivation of a host of genes in plants. In this article we describe molecular, biochemical and physiological factors associated with phosphate acquisition by plants.

Phosphate Acquisition

Legumes, broadly defined by their unusual flower structure, podded fruit, and the ability of 88% of the species examined to date to form nodules with rhizobia ([de Faria et al., 1989][1]), are second only to the Graminiae in their importance to humans. The 670 to 750 genera and 18,000 to 19,000

Legumes: Importance and Constraints to Greater Use

Plants have evolved a number of adaptive responses to cope with growth in conditions of limited phosphate (Pi) supply involving biochemical, metabolic, and developmental changes. We prepared an EMS-mutagenized M(2) population of an Arabidopsis thaliana transgenic line harboring a reporter gene specifically responsive to Pi starvation (AtIPS1::GUS), and screened for mutants altered in Pi starvation regulation. One of the mutants, phr1 (phosphate starvation response 1), displayed reduced response of AtIPS1::GUS to Pi starvation, and also had a broad range of Pi starvation responses impaired, including the responsiveness of various other Pi starvation-induced genes and metabolic responses, such as the increase in anthocyanin accumulation. PHR1 was positionally cloned and shown be related to the PHOSPHORUS STARVATION RESPONSE 1 (PSR1) gene from Chlamydomonas reinhardtii. A GFP::PHR1 protein fusion was localized in the nucleus independently of Pi status, as is the case for PSR1. PHR1 is expressed in Pi sufficient conditions and, in contrast to PSR1, is only weakly responsive to Pi starvation. PHR1, PSR1, and other members of the protein family share a MYB domain and a predicted coiled-coil (CC) domain, defining a subtype within the MYB superfamily, the MYB-CC family. Therefore, PHR1 was found to bind as a dimer to an imperfect palindromic sequence. PHR1-binding sequences are present in the promoter of Pi starvation-responsive structural genes, indicating that this protein acts downstream in the Pi starvation signaling pathway.

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A conserved MYB transcription factor involved in phosphate starvation signaling both in vascular plants and in unicellular algae

Phosphorus (P) is the most important nutrient element (after nitrogen) limiting agricultural production in most regions of the world. It is extremely chemically reactive, and more than 170 phosphate minerals have been identified. In all its natural forms, including organic forms, P is very stable or insoluble, and only a very small proportion exists in the soil solution at any one time. Plant-available P may be considered in either its quantitative or intensive dimension. The quantity of available P is time-specific and crop-specific, because it is the amount of P that will come into the soil solution and be taken up by the crop during its life cycle. The intensity of available P (availability) is most easily identified with its concentration in the soil solution. The soil property controlling the relationship between the solid phase P and its concentration in solution is known as the buffering capacity. The solid phase P involved in this relationship is only a small proportion of the total P, and is known as labile P. It is usually measured by isotopic exchange, but this exchangeable P component does not include the sparingly soluble compounds that also replenish the soil solution as its concentration is depleted by plant uptake. The buffering capacity is the ability of the soil solution to resist a change in its P concentration as P is removed by plant uptake or added in fertilisers or organic materials. Buffering capacity is synonymous with sorptivity, which is a preferable term in the context of the reactivity of P fertiliser with soil. It is usually measured from an adsorption isotherm. By fitting a suitable equation, such as the Langmuir, the total sorption capacity as well as the sorption strength can be determined. Both parameters are important in understanding P availability in soils. Buffering capacity has a major effect on the uptake of labile P because it is inversely related to the ease of desorption of solid phase P and its diffusion. Available P therefore is a direct function of the quantity of labile P and an inverse function of buffering capacity. This has been demonstrated in plant uptake studies. Similarly, the most effective methods of measuring available P (soil tests) are those which remove a proportion of labile P that is inversely related to buffer capacity. Soil tests which measure the concentration of P in solution actually measure availability rather than available P, and their efficacy on a range of soils will depend on the uniformity of the soils" buffer capacities. The most effective soil test usually consists of an anionic extractant. Acidic lactate or fluoride have been found most effective in New South Wales, on a wide range of soils, except calcareous soils which neutralise the acidic component (usually hydrochloric or acetic acid) of the extractant. Sodium bicarbonate (pH 8 · 5) has been found effective on calcareous soils and is widely used throughout the world. It has proved unreliable on NSW soils, and may need more thorough evaluation on non-calcareous soils in other parts of Australia.

I. C. R. Holford

Papers

Soil phosphorus: its measurement, and its uptake by plants