Phosphate transporters of Medicago truncatula and arbuscular mycorrhizal fungi

Most vascular plants acquire phosphate from their environment either directly, via the roots, or indirectly, via a symbiotic interaction with arbuscular mycorrhizal (AM) fungi. The symbiosis develops in the plant roots where the fungi colonize the cortex of the root to obtain carbon from the plant host, while assisting the plant with acquisition of phosphate and other mineral nutrients from the soil solution. As a first step toward understanding the molecular basis of the symbiosis and phosphate utilization, we have cloned and characterized phosphate transporter genes from the AM fungi Glomus versiforme and Glomus intraradices, and from the roots of a host plant, Medicago truncatula. Expression analyses and localization studies indicate that each of these transporters has a role in phosphate uptake from the soil solution.     

植物磷稳态的调控机制

磷是植物必需的重要营养元素之一,是生物大分子的重要组成部分,在植物生命过程中发挥着不可或缺的作用.维持体内磷稳态对于植物的生长发育和环境应答至关重要.多种信号分子参与调控植物对磷的吸收和转运.植物维持磷稳态主要包括土壤磷的活化、磷的吸收、转运、存储和再利用等过程,涉及磷胁迫响应、转录因子调节、miRNA调节、菌根共生、...     

Phosphate transport in plants

Transport of inorganic phosphate (P_i) through plant membranes is mediated by a number of families of transporter proteins. Studies on the topology, function, regulation and sites of expression of the genes that encode the members of these transporter families are enabling roles to be ascribed to each of them. The Pht1 family, of which there are nine members in the Arabidopsis genome, includes proteins involved in the uptake of P_i from the soil solution and the redistribution of P_i within the plant. Members of this family are H₂PO₄ ^–/H⁺ symporters. Most of the genes of the Pht1 family that are expressed in roots are up-regulated in P-stressed plants. Two members of the Pht1 family have been isolated from the cluster roots of white lupin. These same genes are expressed in non-cluster roots. The evidence available to date suggests that there are no major differences between the types of transport systems that cluster roots and non-cluster roots use to acquire P_i. Differences in uptake rates between cluster and non-cluster roots can be ascribed to more high-affinity P_i transporters in the plasma membranes of cluster roots, rather than any difference in the characteristics of the transporters. The efficient acquisition of P_i by cluster roots arises primarily from their capacity to increase the availability of soil P_i immediately adjacent to the rootlets by excretion of carboxylates, protons and phosphatases within the cluster. This paper reviews P_i transport processes, concentrating on those mediated by the Pht1 family of transporters, and attempts to relate those processes involved in P_i acquisition to likely P_i transport processes in cluster roots.     

Phosphate stress response in hydroponically grown maize

The long-term response of hydroponically grown maize plants to variations in the phosphate concentration in the growth medium was studied. There was a 5-week lag period before any differences between experimental and control groups could be seen. After this period, the plants grown without phosphate devoted a higher percentage of their total mass to roots than did the controls. The roots of the phosphate-free plants were longer and less bushy than those of the control plants. Plants grown without phosphate showed an increase in the amount of acid phosphatase extractable from the external surfaces of the roots by a high salt solution. These phosphate stress responses were induced by 5 M phosphate but not by 25 M phosphate.     

The rhizosphere and plant nutrition: a quantitative approach

The role of the rhizosphere in relation to mineral nutrition is discussed within a quantitative framework using the Barber-Cushman model as a starting point. The uptake or release of nutrients by roots growing in soil leads to concentration gradients forming in the soil: the zone so affected is termed the rhizosphere. The nature of these gradients depends on three factors: the rate of uptake/release; the mobility of the nutrient in soil; and the rate of conversion between available and unavailable forms. The interplay between these factors determines the amount of mineral nutrients acquired by the plant and it is the complexity of the interplay which demands the use of mathematical models in order to understand which factors most limit uptake. Despite extensive experimental evidence of root-mediated changes to the physical, chemical and biological status of rhizosphere soil, the quantitative significance of these changes for mineral nutrition has not been assessed. The problems of making this quantitative transition are reviewed.     

Phosphate uptake, efflux and deficiency in the water fern, Azolla

High phosphorus status (High-P) Azolla mexicana plants (P content 15.5 μmoles g fr wt^?1, doubling time ca. 2.2 d) and Low-P plants with early signs of P-deficiency (P content 6.2 μmoles g fr wt^?1, doubling time ca. 3.2 d) were used to study Pi uptake, efflux and deficiency. When High-P plants were transferred to medium lacking Pi, uptake capacity increased 1.5-fold within 12 h and before any detectable change in growth rate (24–48 h). When High-P and Low-P plants were compared, uptake rates from 0.3–10000 mmoles m^?3 Pi were 2.6–1.7 times higher in Low-P than High-P plants (18–1150 vs 7–665 μmoles g fr wt^?1 h^?1). The relationship of uptake rate to concentration was interpreted as arising from a combined operation of a high- and a low-affinity uptake system. Higher uptake in Low-P plants involved a 3.4-fold increase in V_max (high affinity), no change in K_m (high affinity), and a 1.5 to two-fold increase in both V_max (low affinity) and Km (low affinity). Rates of P efflux into 1–1000 mmoles m^?3 Pi were 1.7 to two times higher from High-P than Low-P plants (12–22 vs 7–11 μmoles g fr wt^?1 h^?1). Below 1 mmole m^?3 Pi, uptake and efflux rates were similar: the equilibrium concentration, at which net uptake was zero, was 0.22 mmoles m^?3 for High-P plants and 0.05 mmoles m^?3 for Low-P plants. Similar results were obtained with A. filiculoides. P transport characteristics of Azolla, a fern, are closely comparable with those of higher plants. Its high P requirement in the field arises from its ecological rather than physiological behaviour. We interpret the field behaviour by exploring the relationship between Azolla growth rate in the field, plant P concentration in the field, Pi transport rates required to support such growth, and Pi concentrations in pond waters. The transport characteristics which must operate in the field match those of Low-P plants in the laboratory, not High-P plants. Thus, Pi uptake in High-P plants should be interpreted as repressed from the normal state, instead of that in Low-P plants being induced.     

Phosphate acquisition heterosis in Arabidopsis thaliana: a morphological and physiological analysis

Although phosphate acquisition efficiency (PAE) is of considerable agricultural importance, little is known about its inheritance. The objective of this study was to determine the inheritance of PAE-related morphological and physiological traits in Arabidopsis thaliana. C24 and Col-O, two Arabidopsis accessions differing in their abilities to acquire phosphate from hydroxyl phosphate, were crossed. The resulting hybrid showed superior acquisition of phosphate from hydroxylapatite when compared with either parent. The data suggest that the superiority of the F1 hybrid is due to the accumulation of favourable dominant genes at numerous loci. The hybrid inherited the long root hair length trait from C24 and the long root length trait of Col-O. In addition, the hybrid inherited enhanced phosphate transporter expression from C24. The analysis of morphological and physiological traits in this hybrid will be useful for evaluating and predicting PAE performance in other plant species.     

Human SNPs reveal no evidence of frequent positive selection

We compared the single-nucleotide polymorphisms (SNPs) in humans in 182 housekeeping and 148 tissue-specific genes. SNPs were divided into rare and common polymorphisms based on their frequencies. We found that housekeeping genes tend to be less polymorphic than tissue-specific genes for both rare and common SNPs. Using mouse as a second species for computing sequence divergences, we found no evidence of positive selection: for both housekeeping and tissue-specific genes, the ratio of nonsynonymous to synonymous common SNPs per site showed no significant difference from that of divergence. Similarly, we observed no evidence of positive selection for the 289 and 149 genes that have orthologs available for divergence calculation between humans and chimpanzees and between humans and Old World monkeys, respectively. A comparison with previous SNP studies suggests that approximately 20% of the nonsynonymous SNPs in the human population are nearly neutral and that positive selection in the human genome might not be as frequent as previously thought.     

Phosphate determination during hypokinesia and ambulation in establishing phosphate changes in trained and untrained subjects

Hypokinesia (diminished movement) induces phosphate (P) changes; however, it is not known if P change is greater in trained than untrained subjects. Measuring P balance and P retention during hypokinesia (HK) and P load, we studied if changes in P retention and P depletion were significantly (p<0.05) greater in trained than untrained subjects. Studies were done during a 30-d pre-HK period and a 364-d HK period. Forty male trained and untrained healthy individuals aged 24.5±5.4 yr were chosen as subjects. All volunteers were equally divided into four groups: trained ambulatory control subjects (TACS), trained hypokinetic subjects (THKS), untrained ambulatory control subjects (UACS), and untrained hypokinetic subjects (UHKS). All THKS and UHKS were limited to an average walking distance of 0.3 km/d, and TACS and UACS were on an average running distance of 9.8 and 1.8 km/d, respectively. Subjects took daily 12.7-mmol dicalcium-phosphate/kg body weight in the form of supplementation. Negative P balance, fecal P loss, urinary P and calcium (Ca) excretion, serum P, and total Ca (Ca_t) levels increased significantly (p<0.05), whereas P retention, serum 1,25-dihydroxyvitamin D [1,25 (OH)₂D₃] and intact parathyroid hormone (iPTH) level decreased significantly (p<0.05) in THKS and UHKS when compared with their pre-HK values and their respective ambulatory controls (TACS and UACS). However, P retention, P balance, serum, urinary, and fecal P, and serum hormone level changed significantly (p<0.05) more in THKS than UHKS. Retention of P, fecal P, urinary P and Ca loss, serum P and Ca_t level, P balance, 1,25(OH)₂D₃, and iPTH level change insignificantly (p>0.05) in TACS and UACS when compared with their pre-HK control values. It was concluded that significant negative P balance may indicate P depletion, whereas significant P loss in spite of negative P balance and P load may suggest P retention incapacity; however, P depletion was greater in THKS than UHKS. Clearly, P is wasted much more in THKS than UHKS.     

Phosphate solubilization by organic anion excretion from rice (Oryza sativa L.) growing in aerobic soil

A mathematical model of P solubilization by organic anion excretion from roots is described and used to account for P solubilization by rice (Oryza sativa L.) plants growing in aerobic soil. In previous experiments with rice in an aerobic, highly-weathered, P-deficient soil, we found that the plants were able to solubilize P from an alkali-soluble pool and thereby increase their P uptake. The solubilization could not be explained by pH changes nor by the release of phosphatases. In subsequent experiments we found excretion of citrate from rice roots into nutrient solutions, and the synthesis and excretion of citrate tended to increase under P starvation. The model allows for the diffusion of the organic anion away from a root, its decomposition by soil microbes, its reaction with the soil in solubilizing P, and diffusion of the solubilized P back towards the root as well as away from it. We calculated the rate of citrate excretion from rice roots growing in soil based on measured steady-state citrate concentrations in the rhizosphere and calculated rates of decomposition. Calculations using these and other model parameters obtained independently showed that the observed solubilization and increased P uptake by rice growing in soil could be accounted for. A sensitivity analysis of the model is given. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of freezing on rhizosphere and root nutrient content using two soil sampling methods

This study investigated how deep freezing affects the mineral nutrient content of roots and rhizosphere soil. Two different methods of separating the rhizosphere from the roots were used: i) a brushing method, where the rhizosphere soil was brushed off and ii) a washing method, where the rhizosphere was extracted together with the roots.When unfrozen material was used, the concentrations of K and Fe were significantly higher in the washing method as compared with the brushing method. When the material had been deep frozen, significantly higher concentrations of K, Fe, Mg, and Al were found in the extract from the washing method, indicating a considerable leakage from the roots. No significant differences were found between frozen and unfrozen material with the brushing method. In bulk soil, freezing resulted in increased concentrations of Mn, Al and Fe, even when no roots were present. The brushing method can be used for both frozen and fresh material, although fresh material is preferable. Extraction of soil plus roots cannot be recommended for deep frozen soil.     

Genetic diversity for mycorrhizal symbiosis and phosphate transporters in rice

Phosphorus(P) is a major plant nutrient and developing crops with higher P-use efficiency is an important breeding goal.In this context we have conducted a comparative study of irrigated and rainfed rice varieties to assess genotypic differences in colonization with arbuscular mycorrhizal(AM) fungi and expression of different P transporter genes.Plants were grown in three different soil samples from a rice farm in the Philippines.The data show that AM symbiosis in all varieties was established after 4 weeks of growth under aerobic conditions and that,in soil derived from a rice paddy,natural AM populations recovered within6 weeks.The analysis of AM marker genes(AM1,AM3,AM14) and P transporter genes for the direct Pi uptake(PT2,PT6) and AM-mediated pathway(PT11,PT13) were largely in agreement with the observed root AM colonization providing a useful tool for diversity studies.Interestingly,delayed AM colonization was observed in the aus-type rice varieties which might be due to their different root structure and might confer an advantage for weed competition in the field.The data further showed that P-starvation induced root growth and expression of the high-affinity P transporter PT6 was highest in the irrigated variety IR66 which also maintained grain yield under P-deficient field conditions.     

Phosphate (Pi) and arsenate uptake by two wheat (Triticum aestivum) cultivars and their doubled haploid lines

BACKGROUND AND AIMS: Arsenic accumulation in cereal crops represents an important pathway for human exposure to arsenic from the environment. The objectives of the present work were to find whether the relationship between arsenate and phosphate (Pi) uptake rate differs among genotypes and to select genotypes with a low arsenate uptake rate with the aim of improving food safety and human health. METHODS: A hydroponic experiment was conducted using two wheat (Triticum aestivum) cultivars (Hanxuan 10 and Lumai 14) and ten doubled haploid (DH) lines derived from them to investigate Pi and arsenate uptake over 48 h. Ten plants were transferred to bottles containing 50 mL of pre-treatment solution containing 0.5 mM CaCl2 and 5 mM MES set at pH 6.0 with 330 microM Pi as KH2PO4 and 7.33 microM arsenate. The solutions were aerated continuously. At 8, 24 and 48 h after uptake, 1 mL of test solution was sampled for determination of Pi and arsenate concentrations. KEY RESULTS AND CONCLUSIONS: For each wheat line, Pi and arsenate concentrations in the test solution decreased with uptake time. Exponential (for Pi) or polynomial (for arsenate) regression plots fitted the data closely. For all genotypes, net Pi uptake rates decreased with time (from 0 to 48 h). However, net arsenate uptake rates decreased with time for D5, changed little with time for the male parent, D4 and D6, and increased with time for the others. An inflexion of about 25 microm Pi was observed for the relationship between arsenate and Pi concentrations in the test solution, indicating that 25 microm could be the point where the high-affinity uptake system 'switches on', or dominates over low-affinity uptake. In addition, the male parent, D1, D6 and D10 were considered ideal genotypes because they possess Pi transporters that discriminate strongly against arsenate and are expected to accumulate less arsenate in the field.     

稻、麦根系H~ 的分泌与介质磷水平的关系

水稻、小麦根系H~ 的分泌量随供磷水平的降低而增加,并存在明显的昼夜变化。在自然光照下H~ 分泌量随光强度增加而增多,同时强光比黑暗时H~ 分泌对磷供应水平更为敏感。磷供应不足还诱导水稻根系柠檬酸分泌量增加,而苹果酸则差异不明显。难溶性磷的溶解率与根系H~ 和柠檬酸分泌所导致的根际pH下降有密切联系。因此,在有效磷不足的条件下可明显提高稻、麦根际土壤中难溶性磷的利用率,其中丰产型小麦和粳稻品种对土壤中磷利用的根际效应更为显著。