P-deficient Plants Research Articles

A role for redox factors in shaping root architecture under Phosphorus deficiency

The developmental response of the Arabidopsis root system to low phosphorus (P) availability involves the reduction in primary root elongation accompanied by the formation of numerous lateral roots. We studied the roles of selected redox metabolites, namely, radical oxygen species (ROS) and ascorbic acid (ASC) in the regulation of root system architecture by different P availability. Rapidly growing roots of plants grown on P-sufficient medium synthesize ROS in root elongation zone and quiescent centre. We have demonstrated that the arrest of root elongation at low P medium coincides with the disappearance of ROS from the elongation zone. P-starvation resulted in a decrease in ascorbic acid level in roots. This correlated with a decrease in cell division activity. On the other hand, feeding P-deficient plants with ASC, stimulated mitotic activity in the primary root meristem and partly reversed the inhibition of root growth imposed by low P conditions. In this paper, we discuss the idea of the involvement of redox agents in the regulation of root system architecture under low P availability.

MicroRNA expression profiles associated with phosphorus deficiency in white lupin ( Lupinus albus L.)

Phosphate (P) deficiency is one of the major factors limiting crop growth and production. Although physiological and molecular identification of P deficiency in plants has been the focus for many years, little is known about the regulation of P-deficiency responsive genes and signal networks. In this study, we analyzed the global expression of miRNAs in white lupin ( Lupinus albus) under phosphate deficiency. Total RNAs extracted from P-deficient or P-sufficient tissues was hybridized to an array containing 771 conserved plant miRNA sequences. An induction ratio of greater than 2.0 in all tissues was obtained for 167 miRNAs belonging to 35 miRNA families. Analyses of the data revealed that 35 white lupin miRNA families show differential expression under P deficiency, of which, 17, 9 and 10 were found to be up-regulated, while 7, 6 and 12 were down-regulated in roots, stems and leaves, respectively. These miRNAs were confirmed by RT-PCR-based assay. Within the 35 miRNAs, 30 were successfully validated. Our study provided the evidence that a set of miRNAs are involved in the regulation of adaptive response to P deficiency and that miRNAs coordinately regulate the network of downstream genes controlling morphological and metabolic processes in P-deficient plants.

Leaf phosphorus influences the photosynthesis–nitrogen relation: a cross-biome analysis of 314 species

The ecophysiological linkage of leaf phosphorus (P) to photosynthetic capacity (A (max)) and to the A (max)-nitrogen relation remains poorly understood. To address this issue we compiled published and unpublished field data for mass-based A (max), nitrogen (N) and P (n = 517 observations) from 314 species at 42 sites in 14 countries. Data were from four biomes: arctic, cold temperate, subtropical (including Mediterranean), and tropical. We asked whether plants with low P levels have low A (max), a shallower slope of the A (max)-N relationship, and whether these patterns have a geographic signature. On average, leaf P was substantially lower in the two warmer than in the two colder biomes, with the reverse true for N:P ratios. The evidence indicates that the response of A (max) to leaf N is constrained by low leaf P. Using a full factorial model for all data, A (max) was related to leaf N, but not to leaf P on its own, with a significant leaf N x leaf P interaction indicating that the response of A (max) to N increased with increasing leaf P. This was also found in analyses using one value per species per site, or by comparing only angiosperms or only woody plants. Additionally, the slope of the A (max)-N relationship was higher in the colder arctic and temperate than warmer tropical and subtropical biomes. Sorting data into low, medium, and high leaf P groupings also showed that the A (max)-N slope increases with leaf P. These analyses support claims that in P-limited ecosystems the A (max)-N relationship may be constrained by low P, and are consistent with laboratory studies that show P-deficient plants have limited ribulose-1,5-bisphosphate regeneration, a likely mechanism for the P influence upon the A (max)-N relation.

Metabolite Profiling Reveals Distinct Changes in Carbon and Nitrogen Metabolism in Phosphate-Deficient Barley Plants (Hordeum vulgare L.)

Plants modify metabolic processes for adaptation to low phosphate (P) conditions. Whilst transcriptomic analyses show that P deficiency changes hundreds of genes related to various metabolic processes, there is limited information available for global metabolite changes of P-deficient plants, especially for cereals. As changes in metabolites are the ultimate 'readout' of changes in gene expression, we profiled polar metabolites from both shoots and roots of P-deficient barley (Hordeum vulgare) using gas chromatography-mass spectrometry (GC-MS). The results showed that mildly P-deficient plants accumulated di- and trisaccharides (sucrose, maltose, raffinose and 6-kestose), especially in shoots. Severe P deficiency increased the levels of metabolites related to ammonium metabolism in addition to di- and trisaccharides, but reduced the levels of phosphorylated intermediates (glucose-6-P, fructose-6-P, inositol-1-P and glycerol-3-P) and organic acids (alpha-ketoglutarate, succinate, fumarate and malate). The results revealed that P-deficient plants modify carbohydrate metabolism initially to reduce P consumption, and salvage P from small P-containing metabolites when P deficiency is severe, which consequently reduced levels of organic acids in the tricarboxylic acid (TCA) cycle. The extent of the effect of severe P deficiency on ammonium metabolism was also revealed by liquid chromatography-mass spectrometry (LC-MS) quantitative analysis of free amino acids. A sharp increase in the concentrations of glutamine and asparagine was observed in both shoots and roots of severely P-deficient plants. Based on these data, a strategy for improving the ability of cereals to adapt to low P environments is proposed that involves alteration in partitioning of carbohydrates into organic acids and amino acids to enable more efficient utilization of carbon in P-deficient plants.

Phosphorus acquisition characteristics of cotton (Gossypium hirsutum L.), wheat (Triticum aestivum L.) and white lupin (Lupinus albus L.) under P deficient conditions

A rhizobox experiment was conducted to examine the P acquisition characteristics of cotton (Gossypium hirsutum L.), wheat (Triticum aestivum L.) and white lupin (Lupinus albus L.) under P-deficient conditions. We aimed to identify whether cotton is physiologically efficient at acquiring P through release of protons, phosphatases or carboxylates. Plants were pre-grown in the upper compartment of rhizoboxes filled with a sand and soil mixture to create a dense root mat against a 53 μm polyester mesh. For each species, two P treatments (0 and 20 mg P kg −1 ) were applied to the upper compartment in order to create P-deficient and P-sufficient plants. At harvest, the upper compartment with intact plants was used for collection of root exudates while the lower soil compartment was sliced into thin layers (1 mm) parallel to the rhizoplane. Noticeable carboxylates release was only detected for white lupin. All P-deficient plants showed a capacity to acidify their rhizosphere soil to a distance of 3 mm. The activity of acid phosphatase was significantly enhanced in the soil-root interfaces of P-stressed cotton and wheat. Under P-deficient conditions, the P depletion zone of cotton from the lower soil compartment was narrowest (<2 mm) among the species. Phosphorus fractionation of the rhizosphere soil showed that P utilized by cotton mainly come from NaHCO3-Pi and NaOH-Po pools while wheat and white lupin markedly depleted NaHCO3-Pi and HCl-P pools, and the depletion zone extended to 3 mm. Wheat also depleted NaOH-Po to a significant level irrespective of P supply. The study suggests that acquisition of soil P is enhanced through P mobilization by root exudates for white lupin, and possibly proton release and extensive roots for wheat under P deficiency. In contrast, the P acquisition of cotton was associated with increased activity of phosphatases in rhizosphere soil.

Reduction of Arsenic Oxidative Toxicity By Phosphate Is Not Related To Arsenate Reductase Activity In Wheat Plants

ABSTRACT Arsenic toxicity to plants is thought to be induced through the generation of reactive oxygen species. The response of arsenate reductase (AR) activity as well as the oxidative stress to the interaction of arsenate and phosphate in wheat seedlings was studied in this experiment. The results showed that AR activity increased by 108% with arsenate exposure under P-deficient condition and by 130% under P-sufficient condition; but AR activity showed no significant response to P treatment. Hydrogen peroxide (H2O2) and malondialdehyde (MDA) concentrations increased significantly under arsenate exposure, especially in P-deficient plants, increasing by 180% for H2O2 and 51% for MDA. Superoxide dismutase (SOD) and peroxidase (POD) activity decreased significantly; catalase (CAT) activity decreased under P-deficient condition and increased under P-sufficient condition. Glutathione (GSH), Ascorbate (AsA) and Non-protein thiols (NPT) concentrations all increased when exposed to arsenate under P-deficient condition. Arsenate exposure induced oxidative stress in wheat and P-deprivation increased the damage of oxidative stress.

The BnALMT1 Protein that is an Aluminum-Activated Malate Transporter is Localized in the Plasma Membrane

We have previously reported that Al-induces citrate and malate efflux from P- sufficient and P-deficient plants of rape (Brassica napus L.) and that P-deficiency alone could not induce this response. Further investigation showed that the transcript of two genes designated BnALMT1 and BnALMT2 is accumulated in roots by Al-treatment. Transgenic tobacco cells (Nicotiana tabacum) and Xenopus laevis oocytes expressing the BnALMT1 and BnALMT2 proteins released more malate than control cells in the presence of Al, indicating that the BnALMT genes encode an Al-activated malate transporter. The transgenic tobacco cells exposed to toxic level of Al grew better than control cells indicating that the genes can enhance Al-resistance of plant cells. In this study we showed the subcellular localization of BnALMT1 fused to the green fluoresce protein (GFP). The BnALMT1::GFP construct was transiently expressed in protoplasts prepared from Arabidopsis leaves using the polyethylene glycol (PEG) method. The result showed that the BnALMT1 protein is localized in the plasma membrane. This provides further evidence that the BnALMT proteins facilitate the transport of malate across the plasma membrane (PM).

Phosphorus stress in common bean: root transcript and metabolic responses.

Phosphorus (P) is an essential element for plant growth. Crop production of common bean (Phaseolus vulgaris), the most important legume for human consumption, is often limited by low P in the soil. Functional genomics were used to investigate global gene expression and metabolic responses of bean plants grown under P-deficient and P-sufficient conditions. P-deficient plants showed enhanced root to shoot ratio accompanied by reduced leaf area and net photosynthesis rates. Transcript profiling was performed through hybridization of nylon filter arrays spotted with cDNAs of 2,212 unigenes from a P deficiency root cDNA library. A total of 126 genes, representing different functional categories, showed significant differential expression in response to P: 62% of these were induced in P-deficient roots. A set of 372 bean transcription factor (TF) genes, coding for proteins with Inter-Pro domains characteristic or diagnostic for TF, were identified from The Institute of Genomic Research/Dana Farber Cancer Institute Common Bean Gene Index. Using real-time reverse transcription-polymerase chain reaction analysis, 17 TF genes were differentially expressed in P-deficient roots; four TF genes, including MYB TFs, were induced. Nonbiased metabolite profiling was used to assess the degree to which changes in gene expression in P-deficient roots affect overall metabolism. Stress-related metabolites such as polyols accumulated in P-deficient roots as well as sugars, which are known to be essential for P stress gene induction. Candidate genes have been identified that may contribute to root adaptation to P deficiency and be useful for improvement of common bean.

Phosphate Nutrition and Defoliation Effects on Growth and Root Physiology of Alfalfa

ABSTRACT Phosphorus (P) deficiency reduces forage yield and stand persistence of alfalfa (Medicago sativa L.). The objective of this study was to determine the influence of P nutrition and defoliation on alfalfa shoot growth, root carbohydrate and protein metabolism, and steady-state mRNA levels for high-affinity P transporters. In a greenhouse study, P-deprived plants were provided with 0, 0.25, 2, and 6 mM P beginning 7 d before shoot removal. Plants were sampled immediately (day -7) on days -5, -2, 0 (day of shoot removal), and on days 1, 2, 6, and 9 post-shoot removal. Addition of P to P-deficient plants stimulated growth of shoots but not roots. Taproot bark sugar concentrations were reduced significantly in cut plants at any rate of P, whereas only the 6 mM P treatment reduced taproot wood sugar concentrations in uncut plants. There was a significant defoliation-induced decline in both wood and bark sugar and amino-acid concentration that was enhanced at high P rates. Low P reduced utilization of starch and protein reserves in taproots. Transcripts for a high-affinity P transporter were not detected in any root or shoot tissue assayed, irrespective of defoliation or P treatment. The uncertain relationship between P availability and P-transporter transcript abundance in our greenhouse-grown plants requires additional investigation.

Nitrogen Fixation by White Lupin under Phosphorus Deficiency

White lupin is highly adapted to growth in a low-P environment. The objective of the present study was to evaluate whether white lupin grown under P-stress has adaptations in nodulation and N2 fixation that facilitate continued functioning. Nodulated plants were grown in silica sand supplied with N-free nutrient solution containing 0 to 0.5 mm P. At 21 and 37 d after inoculation (DAI) growth, nodulation, P and N concentration, N2 fixation (15N2 uptake and H2 evolution), root/nodule net CO2 evolution and CO2 fixation (14CO2 uptake) were measured. Furthermore, at 21 DAI in-vitro activities and transcript abundance of key enzymes of the C and N metabolism in nodules were determined. Moreover, nodulation in cluster root zones was evaluated. Treatment without P led to a lower P concentration in shoots, roots, and nodules. In both treatments, with or without P, the P concentration in nodules was greater than that in the other organs. At 21 DAI nitrogen fixation rates did not differ between treatments and the plants displayed no symptoms of P or N deficiency on their shoots. Although nodule number at 21 DAI increased in response to P-deficiency, total nodule mass remained constant. Increased nodule number in P-deficient plants was associated with cluster root formation. A higher root/nodule CO2 fixation in the treatment without P led to a lower net CO2 release per unit fixed N, although the total CO2 released per unit fixed N was higher in the treatment without P. The higher CO2 fixation was correlated with increased transcript abundance and enzyme activities of phosphoenolpyruvate carboxylase and malate dehydrogenase in nodules. Between 21 and 37 DAI, shoots of plants grown without P developed symptoms of N- and P-deficiency. By 37 DAI the P concentration had decreased in all organs of the plants treated with no P. At 37 DAI, nitrogen fixation in the treatment without P had almost ceased. Enhanced nodulation in cluster root zones and increased potential for organic acid production in root nodules appear to contribute to white lupin's resilience to P-deficiency.

Aluminum resistance of cowpea as affected by phosphorus-deficiency stress

Plants growing in acid soils suffer both phosphorus (P) deficiency and aluminum (Al) toxicity stresses. Selection of genotypes for adaptation to either P deficiency or Al toxicity has sometimes been unsuccessful because these two soil factors often interact. Two experiments were conducted to evaluate eight cowpea genotypes for Al resistance and to study the combined effect of P deficiency and Al toxicity stress on growth, P uptake, and organic acid anion exudation of two genotypes of contrasting Al resistance selected from the first experiment. Relative root inhibition by 30 microM Al ranged from 14% to 60% and differed significantly among the genotypes. Al significantly induced callose formation, particularly in Al-sensitive genotypes. P accumulation was significantly reduced (28% and 95%) by Al application for both the Al-resistant and the Al-sensitive genotypes. Al supply significantly enhanced malate release of root apices of both genotypes. However, the exudation rate was significantly higher in the Al-resistant genotype. P deprivation induced an enhanced malate exudation in the presence of Al only in the Al-resistant genotype IT89KD-391. Citrate exudation rate of the root apices was lower than malate exudation by a factor of about 10, and was primarily enhanced by P deficiency in both genotypes. Al treatment further enhanced citrate exudation in P-sufficient, but not in P-deficient plants. The level of citrate exudation was consistently higher in the Al-resistant genotype IT89KD-391 particularly in presence of Al. It is concluded that the Al-resistant genotype is better adapted to acid Al-toxic and P-deficient soils than the Al-sensitive genotype since both malate and citrate exudation were more enhanced by combined Al and P-deficiency stresses.

Interactions between the effects of atmospheric CO2 content and P nutrition on photosynthesis in white lupin (Lupinus albus L.)

Phosphorus (P) is a major factor limiting the response of carbon acquisition of plants and ecosystems to increasing atmospheric CO2 content. An important consideration, however, is the effect of P deficiency at the low atmospheric CO2 content common in recent geological history, because plants adapted to these conditions may also be limited in their ability to respond to further increases in CO2 content. To ascertain the effects of low P on various components of photosynthesis, white lupin (Lupinus albus L.) was grown hydroponically at 200, 400 and 750 micromol mol(-1) CO2, under sufficient and deficient P supply (250 and 0.69 microM P, respectively). Increasing growth CO2 content increased photosynthesis only under sufficient growth P. Ribulose 1,5-biphosphate carboxylase/oxygenase (Rubisco) content and activation state were not reduced to the same degree as the net CO2 assimilation rate (A), and the in vivo rate of electron transport was sufficient to support photosynthesis in all cases. The rate of triose phosphate use did not appear limiting either, because all the treatments continued to respond positively to a drop in oxygen levels. We conclude that, at ambient and elevated CO2 content, photosynthesis in low-P plants appears limited by the rate of ribulose biphosphate (RuBP) regeneration, probably through inhibition of the Calvin cycle. This failure of P-deficient plants to respond to rising CO2 content above 200 micromol mol(-1) indicates that P status already imposes a widespread restriction in plant responses to increases in CO2 content from the pre-industrial level to current values.

Transcriptomic analysis indicates putative metabolic changes caused by manipulation of phosphorus availability in rice leaves

Plants have developed several strategies for coping with phosphorus (P) deficiency. However, the details of the regulation of gene expression of adaptations to low P are still unclear. Using a cDNA microarray, transcriptomic analyses were carried out of the rice genes regulated by P deficiency and P re-supply to P-deficient plants. The OsPI1 gene, which was isolated as the most significant up-regulated gene under -P conditions, was also the most significant down-regulated gene following P re-supply. Many starch metabolism-related genes, as well as several genes for P(i)-liberating enzymes, were up-regulated by -P treatment, suggesting a homeostatic contribution to the P(i) concentration in leaf tissues. mRNAs for glucanases were also induced by P re-supply: these are suspected to play a role in loosening the cell wall compounds. Most of the genes up-regulated by -P treatment were down-regulated by P re-supply, suggesting that their responses were specific to -P conditions. Conversely, the number of genes up-regulated by P re-supply was also larger following P re-supply than in the -P condition. It is proposed that the genes up-regulated by P re-supply play an important role in P acquisition by P-deficient plants.

Transport and partitioning of phosphorus in wheat as affected by P withdrawal during flag-leaf expansion

Increasing P-use efficiency within the plant is one of the acclimations to P-limiting conditions. In this work, we studied the effects of P withdrawal during flag-leaf expansion on sink-source relationships and P-use efficiency in two detillered wheat cultivars (Triticum aestivum L., CA9325 and JM2) under controlled conditions. The study period was divided into two phases of one month each. In the first period after withdrawing P from the medium, the rates of dry weight gain were unaffected compared with the control plant. However, the net dry matter deposition in the ear, and P remobilization within the plant were accelerated in both cultivars. In control plants and in the first period, P transported in the xylem came mainly from the roots’ current uptake in both cultivars; in the second period, however, phloem retranslocation of P from the shoot and cycling through the root contributed 86% in CA9325 and 95% in JM2 to the xylem-transported P. In the P-deficient plants of both cultivars, almost all of the P transported in the xylem was remobilized, exported from vegetative organs and recycled through the phloem. Over the entire duration of the experiment, the net dry matter deposition and P allocation to grains were not synchronous, indicating independent regulatory processes. Although withdrawing P from the medium markedly reduced the net dry weight gain of whole plants in both cultivars, the final dry weight of the grains was hardly influenced. The percentage of grain dry weight to whole plant dry weight increased from 42.5% in control plants to 44.7% in P-deficient plants in CA9325, and from 41.0% to 45.0% in JM2, and that of P increased from 24.8% to 87.7% and from 25.5% to 84.3%, respectively. The results showed that withdrawing P from the medium during flag-leaf expansion did not influence grain growth and its final P content. The possible mechanisms to regulate P redistribution and reutilization in plants are discussed.

Changes Induced by Low Oxygen Concentration in Photosynthetic and Respiratory CO&lt;sub&gt;2&lt;/sub&gt; Exchange in Phosphate-Deficient Bean Leaves

Effect of phosphorus deficiency on photosynthetic and respiratory CO2 exchanges were analysed in primary leaves of 2-week-old bean (Phaseolus vulgaris L. cv. Golden Saxa) plants under non-photorespiratory (2 % O2) and photorespiratory (21 % O2) conditions. Low P decreased maximum net photosynthetic rate (PNmax) and increased the time necessary to reach it. In the leaves of P-deficient plants the relative decrease of PNmax at 2 % O2 was larger than at 21 % O2. The results suggested the influence of photorespiration in the cellular turnover of phosphates.

The role of phosphorus in aluminium-induced citrate and malate exudation from rape (Brassica napus).

Exudation of organic anions is believed to be a common tolerance mechanism for both aluminium toxicity and phosphorus deficiency. Nevertheless, which of these stresses that actually elicit the exudation of organic anions from rape (Brassica napus L) remains unknown, and the combined effects of Al toxicity and P deficiency on rape have not been reported before. Therefore, in the current study, Brassica napus var. Natane nourin plants grown with or without 0.25 mM P were exposed to 0 or 50 micro M AlCl(3) and several parameters related to the exudation of organic anions from the roots were investigated. Eight days of P deficiency resulted in a significant growth reduction, but P deficiency alone did not induce exudation of organic anions. In contrast, Al strongly induced organic acid exudation, while simultaneously inhibiting root growth. Increased in-vitro activity of citrate synthase (CS, EC 4.1.3.7), malate dehydrogenase (MDH, EC 1.1.1.37) and phosphoenolpyruvate carboxylase (PEPC, EC 4.1.1.31), together with reduced root respiration, indicated that the Al-induced accumulation and subsequent exudation of citrate and malate were associated with both increased biosynthesis and reduced metabolism of citric and malic acid. Phosphorus-sufficient plants showed more pronounced aluminium-induced accumulation and exudation of organic anions than P-deficient plants. A divided root chamber experiment showed the necessity of direct contact between Al and roots to elicit exudation of organic anions. Prolonged exposure (10 days) to Al resulted in a decrease in the net exudation of citrate and malate, and the rate of decrease was much more rapid in P-deficient plants than in P-sufficient plants. It is concluded that P nutrition affects the level of Al-induced synthesis and exudation of organic anions. However, the mechanism needs further investigation.

Dynamics of ectomycorrhizal mycelial growth and P transfer to the host plant in response to low and high soil P availability

The aim of this work was to investigate the quantitative relationships between the rates of fungal soil exploration and their effects on plant growth and phosphorus (P) nutrition in soil with varying P availability. Ectomycorrhizal associations were established between Pinus pinaster and the basidiomycete Hebeloma cylindrosporum. Plants were grown for 4 and 6 months in mini-rhizoboxes filled with a 0.5 mm soil layer with two contrasting P levels (−P and +P soils), containing 3 or 50 mg of bicarbonate extractable P per kg of dry soil, respectively. Surface areas of the soil layers colonised by the roots and the hyphae were estimated using image analysis. High P availability decreased the rates of fungal soil colonisation, calculated as 0.92 ± 0.19 cm 2 day −1 plant −1 in the −P soil and 0.42 ± 0.1 cm 2 day −1 plant −1 in the +P soil over the 4–6 months period. Four-month old mycorrhizal plants accumulated lower amounts of biomass and total P than non-mycorrhizal plants, regardless the level of P availability. By contrast, 6-month old mycorrhizal plants were larger and contained more P than non-mycorrhizal plants, especially in the +P soil. However, mycorrhizal plants were always different from non-mycorrhizal P-deficient plants, which had an increased root surface and root P allocation. To explain these contradictory results, we propose that P accumulation by mycorrhizal plants derives mainly from fungal P uptake. The net P transfer from the fungus to the plant was estimated as 0.36 and 0.66 μmol of P per cm 2 of mycelium in −P and +P soil, respectively. Our data demonstrated that, despite the inhibitory effect of the high soil P availability on the rates of fungal soil colonisation, the ectomycorrhizal symbiosis was more efficient to improve host plant P nutrition in these conditions.

Differential responses of rice acid phosphatase activities and isoforms to phosphorus deprivation.

Acid phosphatases (APases) play a role in the release of phosphate in organic complexes in soil. We investigated tissue- and isoform-specific responses of APases to phosphorus (P) deficiency in three rice genotypes; Dasan-byeo, Sobi-byeo, and Palawan. The levels of shoot APase activity per protein were similar in the three genotypes. They significantly decreased with P deprivation that was longer than seven days. Root APase activity per protein was two- to three-fold higher in Dasan than in Sobi and Palawan. In all genotypes the APase activity increased in P-deficient plants, but the increase was higher in Sobi and Palawan. After 21 days of P deprivation, secreted APase activity increased more than eight-fold in Dasan and two-fold in Sobi and Palawan. Isoform profiles of shoot and root APases were most diverse in Dasan. The activities of the major isoforms in P-deficient shoots decreased in all three genotypes. Depending on the genotypes, further increases in constitutive isoforms and new induction of one to four isoforms occurred in P-deficient roots. The results indicate that tissue and genotype differences in the response of APase to P deficiency are primarily facilitated by the different responses of the isoforms.

High‐affinity phosphate/arsenate transport in white lupin (Lupinus albus) is relatively insensitive to phosphate status

• The development of proteoid roots under phosphorus deficiency by white lupin (Lupinus albus) may result in increased arsenate uptake, as arsenate is a phosphate analogue. This, together with its high biomass production, rapid growth and ability to survive in soils with low phosphate and nitrogen contents, low pH and high metal contents make them an interesting species to investigate with respect to revegetation, and possibly also for long-term phytoremediation, of arsenic contaminated soils. • Kinetic parameters for arsenate uptake for P-deficient and P-sufficient plants, as well as for proteoid and nonproteoid roots were obtained. Down-regulation of arsenate uptake by phosphate, as well as phosphate/arsenate competition for P-deficient and P-sufficient plants was studied. • Arsenate uptake was reduced by phosphate, but small differences were found between P-deficient and P-sufficient plants. Arsenate uptake by proteoid roots was higher than for nonproteoid roots of P-deficient plants, with higher Vmax and similar Km values. Down-regulation of the high affinity phosphate/arsenate uptake system by phosphate does take place but seems to be slower than in other plants. • This study suggests that the low sensitivity of the phosphate/arsenate uptake system to regulation by phosphate may be related to the adaptations of white lupin to low P available environments. Such adaptation are absent in plants unable to develop proteoid roots.

PHOTOSYNTHATE DISTRIBUTION IN WHEAT VARIETIES DIFFERING IN PHOSPHORUS EFFICIENCY

The distribution of phosphorus (P), photosynthates, and the amount of assimilated 14C secreted as root exudate were measured in wheat genotypes differing in phosphorus efficiency. Two wheat genotypes, one P-efficient (Triticum aestivum L., cv. Xiaoyan 54) and the other inefficient (Triticum aestivum L., cv. Jing 411) were used in this study. The plants were cultivated in solution culture for 4 weeks under controlled conditions. Xiaoyan 54 had higher relative yield, higher P concentration in roots and higher root/shoot ratio than Jing 411 under P-deficient condition. Xiaoyan 54 distributed more P and assimilated carbon to the roots than Jing 411 under P deficiency and also released a higher percentage of assimilated 14C as root exudates. The percentage released was 3.98% for Xiaoyan 54 and 2.47% for Jing 411 under P deficiency, and 2.95% and 3.28%, respectively, under P-sufficiency. However, P deficiency led to more assimilated 14C being partitioned to roots by both wheat genotypes. Regardless of genotypes, the 14C distributed to roots of P-deficient plants was 5 to 6 times that of P-sufficient plants. The P-efficient genotype translocated more assimilated 14C to roots than the P-inefficient genotype under P deficiency and the difference between the two genotypes were smaller under P sufficiency. It is concluded that Xiaoyan 54 has higher relative biomass and better root system under P limited condition, which main reasons are that Xiaoyan 54 can maintain a higher P concentration, release more root exudate, and distribute P to roots.