Phosphate aquisition by red clover and black mustard on a humic podzol

Recent investigations have shown that phosphate (P) mobilization by root exudates is an important feature of genotypes to acquire P even in soils of low‐P availability. We, therefore, investigated P mobilization processes in the rhizosphere of red clover (Trifolium pratense L.) and black mustard (Brassica nigra L.) on a humic podzol. As measured by the Kuchenbuch‐Jungk method (Kuchenbuch and Jungk, 1982), both species accumulated similar quantities of citrate (12 μmol/g soil) in the rhizosphere in about 1 mm distance from the soil‐root interface. Despite of similar concentrations of P‐mobilizing citrate in the rhizosphere of both species, red clover took up nearly the two‐fold of P compared to black mustard. Differences in rhizosphere pH were determined between both species. Black mustard did not acidify the rhizosphere, whereas red clover decreased the pH in the rhizosphere from 5.8 to about 4.0 (in 0.01M CaCl₂). The simultaneous acidification and excretion of citrate compared to citrate excretion alone had consequences for P mobilization processes in the rhizosphere. Phosphate mobilization from the soil solid phase was higher at higher pH. Thus, the citrate‐induced P desorption was not the limiting step in P acquisition by red clover and black mustard. Calculations of P distribution in the soil solution between free ortho‐P and humic‐associated P showed that at higher pH most of the P was associated with dissolved humic substances, whereas at pH < 5, most of the P was present as free ortho‐P. These P species can readily be taken up by the roots whereas humic‐associated P must probably be desorbed from the humic surface before uptake. Phosphate species calculations, therefore, explained the higher P uptake of red clover compared to black mustard. Aluminum species distribution calculations in the soil solution further show that even at pH < 5.0 in the soil solution, citrate strongly complex Al and thereby reduce the activity of monomeric Al species. The excretion of citrate can, therefore, counteract the root induced acidification of the rhizosphere with respect to Al toxicity.     

The excretion of citric and malic acid by proteoid roots of Lupinus albus L.; effects on soil solution concentrations of phosphate,iron, and aluminum in the proteoid rhizosphere in samples of an oxisol and a luvisol

Organic acid concentration in the proteoid rhizosphere of White Lupin in different soil samples (Oxisol-A_p = Ox, Luvisol-A_p and Luvisol-C = L_A and L_C) was determined in order to study the influence of root-released carboxylates on the mobilization of phosphate, aluminum, and iron in the rhizosphere. In the L_C, organic acids were accumulated as Casalts extractable with water. In the proteoid rhizosphere of this soil sample 55 μmol citrate and 8 μmol malate per g soil were found. In the Ox, no water extractable organic acids were present. However, determination of citrate in the solid phase of this soil by Diffuse Reflectance Infrared Fourier Transform Spectroscopy gave concentrations of 88 and 68 μmol citrate per g soil without and with P application, respectively. Displaced soil solution from the proteoid root rhizosphere of the Ox and the L_A increased in Fe and Al concentrations from <50 μmol/L (soil from reference pots without plants) to more than 600 μmol Fe+Al/L. The concentration of P was increased by a factor of 2 despite of P uptake by the proteoid roots. The mobilization of Al, Fe, and P is attributed to ligand exchange of phosphate against citrate and to the solubilization of Al and Fe as carboxylate complexes.     

Biodegradation of low molecular weight organic acids in rhizosphere soils from a tropical montane rain forest

Root exudation of organic acids could be an important strategy for plant acquisition of phosphorus (P) from P-deficient soils in tropical rain forests. However, the efficacy of organic acids on P mobilization in the rhizosphere could be reduced due to their rapid biodegradation by rhizosphere microorganisms. To assess the dynamics and function of organic acids in the rhizosphere soils in tropical rain forests, we examined the concentrations of oxalate, citrate, and malate in soil solution and the mineralization kinetics of ¹⁴C-radiolabelled oxalate and citrate in the rhizosphere and bulk soil fractions. We compared two tropical montane rain forests from Mt. Kinabalu, Borneo that share similar parent material (i.e., sedimentary rocks) and climate but differ in terms of soil age. The older soil (Tertiary age materials) was affected by podzolization and had less inorganic labile P compared to the younger soil (Quaternary colluvial deposits). In the P-deficient older soil, the rhizosphere soil solution contained markedly higher concentrations of oxalate, citrate, and malate than did the bulk soil, whereas in the P-rich younger soil, the levels of organic acids in the rhizosphere were lower. The higher levels of organic acids in the rhizosphere of P-deficient soils are caused by greater root exudation and the lower sorption capacity for organic acids. The results of mineralization kinetics showed that oxalate and citrate in soil solution were rapidly mineralized in both rhizosphere and bulk fractions of both P-rich and P-deficient soils, having short mean residence times (2.3–13.1 h for oxalate and 0.8–1.6 h for citrate). The mineralization rates of oxalate and citrate were highest in the rhizosphere fraction of the P-deficient soil, where the pool of organic acids was largest and rapidly replenished by root exudation. Our data indicate that consumption as well as production of organic acids in the rhizosphere could be enhanced in P-deficient soil. The efficacy of organic acids on P mobilization in the rhizosphere in tropical montane rain forests appears to vary depending on the level of soil P availability and the anion sorption capacity, attributable to soil aging with podzolization.     

The acquisition of phosphate by higher plants: Effect of carboxylate release by the roots. A critical review.

Phosphorus is one of the most limiting macronutrients for plant productivity in agriculture worldwide. The main reasons are the limited rock phosphate reserves and the high affinity of phosphate (P) to the soil solid phase, restricting the P availability to the plant roots. Plants can adapt to soils low in available P by changing morphological or/and physiological root features. Morphological changes include the formation of longer root hairs and a higher root : shoot ratio both parameters increasing the root surface which provides the shoot with P. This may be successful if the P availability in soil, i.e., the P concentration of the soil solution is not extremely low (> 1–2 µM P). If the P concentration of the soil solution is lower, the diffusive flux to the root surface will be very low and may not satisfy the P demand of the shoots. Under these conditions plants have developed strategies to increase the rhizosphere soil solution concentration by secreting mobilizing agents. The most effective way of P mobilization is the release of di‐ and tricarboxylic acid anions, especially oxalate and citrate. Citrate can accumulate in the rhizosphere up to concentrations up to 80 µmol g^?1 soil. Cluster root formation is an efficient way of carboxylate accumulation in the cluster root rhizosphere improving P mobilization. Cluster roots strongly improve the acquisition of the mobilized P. Considering a single root, around 80–90% of the mobilized P diffuses away from the root. From the rhizosphere of cluster roots, most of the mobilized P is taken up by the cluster roots. Both, the strong accumulation of carboxylates in and the effective P uptake from the cluster‐root rhizosphere are the basis of the unique ability of P acquisition by cluster root‐forming plants. Plants that do not form cluster roots, e.g., red clover, can also accumulate carboxylates in the rhizosphere. Red clover accumulates high quantities of citrate in the rhizosphere soil. Model calculations show that the release of citrate by red clover roots and its accumulation in the rhizosphere strongly improve P acquisition by this plant species in various soils. Similar results are obtained with alfalfa. In sugar beet, oxalate release can strongly contribute to P acquisition. In summary, P acquisition can be strongly improved by the release of carboxylates and should be taken as a challenge for basic and applied research.     

Availability to plants of phosphate adsorbed on goethite: experiment and simulation

A model has been developed to predict the availability of sorbed phosphate to plants. It is based on uptake by a growing root system. Phosphate sorption is assumed to be reversible and the rate of desorption is assumed not to limit uptake. The model was tested against observed availability of sorbed phosphate estimated from uptake by plants growing in phosphorus-deficient conditions. The plants were grown on quartz sand coated with goethite on which phosphate was adsorbed, and which was mixed with nutrient solution. The influence of the phosphate loading of goethite and the goethite content of the sand on available phosphate was examined. Phosphate uptake was predicted well by the model for treatments with high phosphate loading and different goethite content of the sand. However, for a treatment with low phosphate loading, uptake was underestimated by the model. In this treatment the pH of the 'soil’ solution decreased from 5·5 to 4·2 during the experiment. The phosphate concentration in solution increased with a lowering of the pH in this pH range, resulting in increased phosphate uptake.     

Phosphate rock dissolution and transformation in the rhizosphere of tea (Camellia sinensis L.) compared with other plant species

Tea (Camellia sinensis L.) is generally grown in highly weathered acidic Ultisols of the humid tropics. The low pH, large P fixing capacity and moisture content of these soils favour the dissolution of phosphate rock. Plant species differ widely in their ability to take up P from phosphate rock, and we have compared phosphate mobilization in the rhizosphere of tea with that under calliandra (Calliandra calothyrsus L.), Guinea grass (Panicum maximum L.) and bean (Phaseolus vulgaris L.) by studying the changes in the concentration of P fractions at known distances from the root surface in an acidic (pH in water 4.5) Ultisol from Sri Lanka treated with a phosphate rock. Plants were grown in the top compartment of a two-compartment device, comprising two PVC cylinders physically separated by a 24-μm pore-diameter polyester mesh. A planar mat of roots was formed on the mesh in the top compartment, and the soil on the other side of the mesh in the lower compartment was cut into thin slices parallel to the rhizosphere and analysed for pH and P fractions. All plant species acidified the rhizosphere (pH [water] difference between bulk and rhizosphere soils was 0.17-0.26) and caused more rock to dissolve in the rhizosphere (10–18%) than in the bulk soil (8–11%). Guinea grass was most effective, though the rate of acidification per unit root surface area was least (0.02μmol H⁺ cm^?2) among the four species. Tea produced the largest rate of acidification per unit root surface area (0.08μmol H⁺ cm⁺²). All species depleted P extracted by a cation–anion exchange resin and inorganic P extracted by 0.1 M NaOH. All except tea depleted organic P extracted by 0.1M NaOH in the rhizosphere. The external P efficiencies (mg total P uptake) of Guinea grass, bean, tea and calliandra in soil fertilized with phosphate rock were 4.82 ± 0.42, 4.02 ± 0.32, 1.06 ± 0.02 and 0.62 ± 0.02, respectively, and the corresponding internal P efficiencies (mg shoot dry matter production per mg plant P) were 960 ± 75, 1623 ± 79, 826 ± 33 and 861 ± 44. This study showed that the various crops cultivated in tea lands differ in their rates of acidification, phosphate rock dissolution and P transformation in the rhizosphere. This requires testing under field conditions.     

Adsorption of glycerophosphate on goethite (α‐FeOOH): A macroscopic and infrared spectroscopic study

Adsorption, desorption, and precipitation reactions at environmental interfaces govern the bioavailability, mobility, and fate of organic phosphates in terrestrial and aquatic environments. Glycerophosphate (GP) is a common environmental organic phosphate, however, surface adsorption reactions of GP on soil minerals have not been well understood. The adsorption characteristics of GP on goethite were studied using batch adsorption experiments, zeta (ζ) potential measurements, and in situ attenuated total reflectance‐Fourier transform infrared spectroscopy (ATR‐FTIR). GP exhibited fast initial adsorption kinetics on goethite, followed by a slow adsorption. The maximum adsorption densities of GP on goethite were 2.00, 1.95, and 1.44 μmol m^?2 at pH 3, 5, and 7, respectively. Batch experiments showed decreased adsorption of GP with increasing pH from 3 to 10. Zeta potential measurements showed a remarkable decrease in the goethite isoelectric point upon GP adsorption (from 9.2 to 5.5), suggesting the formation of inner‐sphere surface complexes. In addition, the ATR‐FTIR spectra of GP sorbed on goethite were different from those of free GP at various pH values. These results suggested that GP was bound to goethite through the phosphate group by forming inner‐sphere surface complexes.     

Desorption Characteristics of Three Mineral Oxides and a Non-crystalline Aluminosilicate for Supplying Phosphate in Soilless Root Media

The objective was to evaluate phosphate desorption characteristics of synthetic hematite, goethite, and allophane and commercial alumina after loading at maximum adsorbed phosphate levels to determine their potential to release phosphate at a constant, low level to sustain plant growth in soilless media and reduce phosphate leaching. Desorption isotherms were measured at pH 6.4 ± 0.1 using a continuously stirred-flow reactor. The time period during which dissolved phosphate was maintained within the range of 5–0.2 mg·L^?1 phosphate-P decreased in the order: allophane (12.4 d) > alumina (4.6 d) > goethite (3.6 d) > hematite (1.9 d). Allophane released the most phosphate during the desorption process (40% of maximum adsorbed phosphate; 12.7 mg?g^?1) followed by alumina and goethite (19–20%; ≈2.5 mg?g^?1) and lastly hematite (5%; 0.1 mg?g^?1). Allophane demonstrated the greatest potential as a phosphate-charged source for soilless root media, in amount and duration of phosphate release.     

Effects of crystallinity of goethite: II. Rates of sorption and desorption of phosphate

Eight samples of goethite ranging in surface area from 18 to 132 m² g^-1 were mixed with phosphate at a range of pH values for periods which ranged from 0·5 h to 6 weeks. The sample with a surface area of 18 m² g^-1 had been hydrothermally treated to improve its crystallinity. Its rate of reaction with phosphate depended on pH but was complete within a day. Its maximum observed reaction was close to the theoretical maximum for surface adsorption of 2·5 μmol m^?2. For the other samples, phosphate continued to react for up to 3 weeks and exceeded the value of 2·5 μmole m^?2. The duration and extent of the reaction depended on the crystallinity of the goethite. The results were closely described by a model in which the phosphate ions were initially adsorbed on to charged external surfaces. The phosphate ions then diffused into the particles. This was closely described using equations for diffusion into a cylinder. Samples of goethite which had been loaded with phosphate dissolved more slowly in HCl, and had a longer lag phase, than phosphate-free goethite. For the hydrothermally treated goethite, HCl removed much of the phosphate when only a small proportion of the iron had been dissolved. For a poorly crystallized goethite, it was necessary to dissolve much more of the iron to obtain a similar removal of phosphate. Brief treatment with NaOH removed most of the phosphate from the hydrothermally treated goethite but only half the phosphate from a poorly crystallized goethite. These results are consistent with the idea that phosphate ions were not only bound on external surface sites but had also penetrated into meso- and micro-pores between the domains of the goethite crystals and were then adsorbed on internal surface sites. This penetration tied the domains together more firmly thus increasing the lag phase for dissolution. Differences between sites for phosphate adsorption are therefore caused mainly by their location on either external or internal sites. Models that ignore this are incomplete.     

碳对微生物–根系介导的蔬菜作物磷吸收的影响

  【目的】  碳是微生物代谢活动的能量来源,解析碳驱动的微生物磷周转对根系/根际属性以及作物磷吸收的影响,对探索提高磷利用效率的根际调控措施具有重要的指导意义。  【方法】  以绿叶蔬菜上海青(Brassica chinensis L., Xiaqing 3)为供试作物进行盆栽试验,供试碳源为葡萄糖。设置添加葡萄糖(+G)和不添加葡萄糖(?G,对照)两个处理,在添加葡萄糖后第7天和第21天,测定土壤微生物量磷与Olsen-P含量、根际酸性磷酸酶活性以及柠檬酸和苹果酸含量、根系形态(生物量、根冠比、根长、根系直径、比根长和根系组织密度)与根际生理(酸性磷酸酶、柠檬酸和苹果酸)指标和作物磷吸收量。  【结果】  添加葡萄糖后第7天,土壤微生物量磷增加,Olsen-P含量降低;上海青根系生物量和根冠比显著高于对照,另外,与不加葡萄糖处理相比,添加葡萄糖导致上海青总根长降低33%,根系平均直径增加27%,比根长降低46%,根际柠檬酸含量增加106%。从第7天到第21天,添加葡萄糖处理土壤微生物量磷降低,Olsen-P含量增加,上海青根系生长速率显著提高。葡萄糖添加后第21天,添加葡萄糖处理土壤Olsen-P含量高于对照土壤;与不加葡萄糖的处理相比,根际酸性磷酸酶和柠檬酸的分泌降低,上海青根系总根长增加,其相对增加量为31%。添加葡萄糖对第7天和第21天上海青地上部磷吸收没有显著影响。  【结论】  添加葡萄糖提高了前期(添加葡萄糖后第7天)根际微生物量磷,降低了Olsen-P含量,促进根际柠檬酸的分泌满足作物生长对磷的需求。后期(添加葡萄糖后第21天),微生物量磷的降低促进土壤有效磷含量的增加,刺激根系快速伸长。微生物介导磷周转诱导作物调节根系形态和根际分泌物响应土壤磷环境的变化,维持地上部磷营养。     

Effect of buckwheat (Fagopyrum esculentum) on soil‐phosphorus availability and organic acids

As a cover crop, buckwheat (Fagopyrum esculentum) may increase soil‐P availability. Buckwheat was grown in low‐P and P‐fertilized field plots, and organic anions were measured in rhizosphere soil. Soil‐P availability was not affected by buckwheat, but the concentration of rhizosphere tartrate^2– was significantly higher (p < 0.005) in low‐P vs. P‐fertilized plots. This suggests that organic‐anion root exudation may have a role in buckwheat‐rhizosphere P dynamics.     

Root-induced changes in the rhizosphere: Importance for the mineral nutrition of plants

Root-induced changes in the rhizosphere may affect mineral nutrition of plants in various ways. Examples for this are changes in rhizosphere pH in response to the source of nitrogen (NH₄-N versus NO₃-N), and iron and phosphorus deficiency. These pH changes can readily be demonstrated by infiltration of the soil with agar containing a pH indicator. The rhizosphere pH may be as much as 2 units higher or lower than the pH of the bulk soil. Also along the roots distinct differences in rhizosphere pH exist. In response to iron deficiency most plant species in their apical root zones increase the rate of H⁺ net excretion (acidification), the reducing capacity, the rate of Fe^III reduction and iron uptake. Also manganese reduction and uptake is increased several-fold, leading to high manganese concentrations in iron deficient plants. Low-molecular-weight root exudates may enhance mobilization of mineral nutrients in the rhizosphere. In response to iron deficiency, roots of grass species release non-proteinogenic amino acids (?phytosiderophores”?) which dissolve inorganic iron compounds by chelation of Fe^III and also mediate the plasma membrane transport of this chelated iron into the roots. A particular mechanism of mobilization of phosphorus in the rhizosphere exists in white lupin (Lupinus albus L.). In this species, phosphorus deficiency induces the formation of so-called proteoid roots. In these root zones sparingly soluble iron and aluminium phosphates are mobilized by the exudation of chelating substances (probably citrate), net excretion of H⁺ and increase in the reducing capacity. In mixed culture with white lupin, phosphorus uptake per unit root length of wheat (Triticum aestivum L.) plants from a soil low in available P is increased, indicating that wheat can take up phosphorus mobilized in the proteoid root zones of lupin. At the rhizoplane and in the root (root homogenates) of several plant species grown in different soils, of the total number of bacteria less than 1 % are N₂-fixing (diazotrophe) bacteria, mainly Enterobacter and Klebsiella. The proportion of the diazotroph bacteria is higher in the rhizosphere soil. This discrimination of diazotroph bacteria in the rhizosphere is increased with foliar application of combined nitrogen. Inoculation with the diazotroph bacteria Azospirillum increases root length and enhances formation of lateral roots and root hairs similarly as does application of auxin (IAA). Thus rhizosphere bacteria such as Azospirillum may affect mineral nutrition and plant growth indirectly rather than by supply of nitrogen.     

Kinetics of soil phosphate desorption as affected by citric acid

Citrate can accumulate in the rhizosphere of phosphate (P) deficient plants. Therefore, P desorption was investigated in relation to citrate concentrations applied in a single or sequential mode to a Luvisol and a Podzol. Cumulative P desorption increased up to 20 times that of the control, the absolute increase being higher in the Podzol. The rate of P release was closely related to the rate of Fe+Al release indicating that desorption originated mainly from Fe/Al associated P. The mode of citrate loading affected the P desorption in an ambiguous manner. P-diffusion coefficients were calculated from the desorption data. The coefficients strongly increased with citrate loading, the increase being more pronounced in the Podzol.     

Phosphate Speciation and Citrate-Induced Mobilization of P in an Acric Oxisol

Billions of dollars are spent annually in Brazil on imported phosphorus (P) fertilizers for agricultural crops produced on Oxisols. Phosphate fixation by Oxisols is a main limitation to crop productivity because these highly weathered soils have a high phosphate adsorption capacity. The objective was to determine whether reaction of an Acric Oxisol with citrate increases P availability. Columns of P-amended subsoil material were leached with 1 mM citrate solution, and effluent was monitored. Solid-phase speciation was measured for different stages of leaching using X-ray absorption near edge structure (XANES) spectroscopy at the P K-edge. When soil columns were leached with up to 56 column pore volumes of 1 mM citrate solutions, no P was detected in column effluent samples, whereas aluminum (Al) and iron (Fe) increased up to 30 and 1.4 µmol L^?1, respectively. The XANES analysis suggested that with increased leaching, a minor proportion of phosphate adsorbed on Fe-oxides increased as phosphate adsorbed on Al-oxides decreased. The results indicated that citrate tends to induce minor species redistribution of phosphate between Al- and Fe-oxide bound forms, but P mobilization was trivial compared with complexometric mobilization of Fe and Al.     

Presence of bacteria reduced phosphate adsorption on goethite

Despite extensive studies, the information obtained from pure iron and aluminum (hydr)oxides cannot fully explain phosphate fixation in natural soils because of the ubiquitous interactions between (hydr)oxides and bacteria in soil. The effect of bacteria (Bacillus subtilis subsp. and Pseudomonas fluorescens) on phosphate adsorption on goethite (α‐FeOOH) was systematically examined under varying reaction times, phosphate concentrations, pH, ionic strength and bacteria dosage. Batch experiments in all cases showed significantly less adsorption on bacteria–goethite complexes than on pure goethite, demonstrating an inhibitory effect of bacteria. The inhibition of phosphate adsorption increased with bacterial loading, and showed a significant, non‐linear correlation with the decrease in the goethite positive charge induced by the bacteria. Moreover, in both the desorption experiment and in situ, the attenuated total reflectance Fourier‐transform infrared (ATR‐FTIR) spectra suggested a competition of bacteria surface groups (phosphate and carboxyl) with solution phosphate for hydroxyl on goethite. Therefore, the negative influences of bacteria on phosphate adsorption on goethite were probably caused by the surface charge modification and the competitive adsorption induced by the bacteria. Under most conditions, the effects of B. subtilis subsp. were conspicuous, while only slight influences were found for P. fluorescens. This difference between the two bacteria species was explained by differences in their surface charge, group content and chemical interaction with goethite. These findings uncover an important role of bacteria in phosphate phyto‐availability and mobility in natural environments.     

Distribution of exudates of lupin roots in the rhizosphere under phosphorus deficient conditions

Abstract

The distribution of secretory acid phosphatase and organic acids enhanced by phosphorus deficiency in lupin rhizosphere was investigated using a rhizobox system which separated the rhizosphere soil into 0.5 mm fractions. In the soil fraction closest to the root surface, the lupin exudates displayed an acid phosphatase activity of 0.73 u g^?1 dry soil and citrate concentration of 85.2 μmol g^?1 dry soil, respectively. The increase of the acid phosphatase activity-induced an appreciable depletion of organic P in the rhizosphere, indicating that lupin efficiently utilized the organic P from soil through the enzyme activitye The sterile treatments demonstrated that the acid phosphatase in the rhizosphere was mainly derived from lupin root secretions. The secretory organic acids enhanced considerably the solubility of the inorganic P in three types of soil and a sludge. However, the secretory acid phosphatase and organic acids from lupin roots were only detected in a considerable amount in 0-2.5 mm soil fractions from root surface.     

Organic acid behaviour in a calcareous soil implications for rhizosphere nutrient cycling

Calcareous soils are frequently characterized by the low bioavailability of plant nutrients. Consequently, many vascular plant species are unable to successfully colonize calcareous sites and the floristic composition of calcareous and acid silicate soils has been shown to differ markedly. The root exudation of oxalate and citrate has been suggested to play a pivotal role in same nutrient acquisition mechanisms operating in calcareous soils. The aim of this study was therefore to investigate the nutrient extraction efficiency of three individual organic acids commonly identified in root exudates, i.e. citric, malic and oxalic acid. Our results clearly demonstrate the context dependent nature of nutrient release by organic acids. The degree of P extraction was highly dependent on which organic acid was added, their concentration and pH, and their contact time with the soil. P is generally more efficiently extracted by organic acids at a high pH and follows the series oxalate>citrate>malate. The opposite relationship between pH and extraction efficiency was apparent for most other cations examined (e.g. Zn, Fe), which are more efficiently extracted by organic acids at low pH. A serious constraint to the ecological importance of organic acid exudation in response to P deficiency is, however, their very low P mobilization efficiency. For every mol of soil P mobilized, 1000 mol of organic acid has to be added. It can, however, be speculated that in a calcareous soil with extremely low P concentrations it is still beneficial to the plants to exude organic acids in spite of the seemingly high costs in terms of carbon.     

Phosphorus Efficiency of Buckwheat (Fagopyrum esculentum)

The ability of buckwheat (Fagopyrum esculentum) roots to acquire phosphorus (P) was characterized by investigating P uptake, morphological features, and chemical changes in the rhizosphere. Over a range of nutrient solution P concentrations (5–500 μmol · L^?1), maximum shoot growth was achieved with a P supply between 5 and 100 μmol · L^?1. Root weight and root length, as well as length and frequency of root hairs, were higher at low P levels. Root surface and the root surface/shoot dry weight ratio reached high values. Though P uptake rates were only moderate (0.15 pmol · cm^?1 root · sec^?1), shoot P concentrations were high (1.8% of dry weight with 100 μM P) predominantly being inorganic (80%). Phosphorus efficiency was characterized by a high specific absorption rate (810 mmol P · kg^?1 root dry wt · d^?1) rather than by an efficient utilization for dry weight production. Root exudates of low-P plants had lower pH values than exudates of high-P plants and increased the solubility of FePO₄ and MnO₂ to a greater extent. Amounts of exuded organic acids and phenolics were low and could not account for the observed solubilization of FePO₄ and MnO₂. Enhanced hydrolysis of glucose-6-phosphate by exudates from low-P plants was due to an increased “soluble” acid phosphatase activity, and root surface phosphatase activity was also slightly enhanced with P deficiency. In the rhizosphere soil of buckwheat, some depletion of organic P forms was observed, and in pot trials with quartz sand, buckwheat utilized glucose- 6-phosphate as a P source at the same rate as inorganic P.     

铝和镉胁迫对两个大麦品种矿质营养和根系分泌物的影响

A hydroponic experiment was carried out to study the effect of aluminum （Al） and cadmium （Cd） on Al and mineral nutrient contents in plants and Al-induced organic acid exudation in two barley varieties with different Al tolerance. Al- sensitive cv. Shang 70-119 had significantly higher Al content and accumulation in plants than Al-tolerant cv. Gebeina, especially in roots, when subjected to low pH （4.0） and Al treatments （100 μmol L^-1 Al and 100 μmol L^-1 Al ＋1.0 μmol L^-1 Cd）. Cd addition increased Al content in plants exposed to Al stress. Both low pH and Al treatments caused marked reduction in Ca and Mg contents in all plant parts, P and K contents in the shoots and leaves, Fe, Zn and Mo contents in the leaves, Zn and B contents in the shoots, and Mn contents both in the roots and leaves. Moreover, changes in nutrient concentrations were greater in the plants exposed to both Al and Cd than in those exposed only to Al treatment. A dramatic enhancement of malate, citrate, and succinate was found in the plants exposed to 100 μmol L^-1 Al relative to the control, and the Al-tolerant cultivar had a considerable higher exudation of these organic acids than the Al-sensitive one, indicating that Al-induced enhancement of these organic acids is very likely to be associated with Al tolerance.     

Assessment of diffusive phosphate supply in soils by microdialysis

Standard procedures to assess P availability in soils are based on batch experiments with various extractants. However, in most soils P nutrition is less limited by bulk stocks but by strong adsorption and transport limitation. The basic principle of root‐phosphate uptake is to strip phosphate locally from the solid phase by forming a radial depletion zone in the soil solution, optionally enhanced by release of mobilizing substances. Microdialysis (MD), a well‐established method in pharmacokinetics, is capable to mimic important characteristics of P root uptake. The sampling is by diffusional exchange through a semipermeable membrane covering the probes with their sub‐mm tubular structure. Additionally, the direct environment of the probe can be chemically modified by adding, e.g ., carboxylates to the perfusate. This study is the first approach to test the applicability of MD in assessing plant available phosphate in soils and to develop a framework for its appropriate use.We used MD in stirred solutions to quantify the effect of pumping rate, concomitant ions, and pH value on phosphate recovery. Furthermore, we measured phosphate yield of top‐soil material from a beech forest, a non‐fertilized grassland, and from a fertilized corn field. Three perfusates have been used based on a 1 mM KNO₃ solution: pure (1), with 0.1 mM citric acid (2), and with 1 mM citric acid (3). Additionally, a radial diffusion model has been parametrized for the stirred solutions and the beech forest soil.Results from the tests in stirred solutions were in good agreement with reported observations obtained for other ionic species. This shows the principal suitability of the experimental setup for phosphate tests. We observed a significant dependency of phosphate uptake into the MD probes on dialysate pumping rate and on ionic strength of the outside solution. In the soils, we observed uptake rates of the probes between 1.5 × 10⁻¹⁵ and 6.7 × 10⁻¹⁴ mol s⁻¹ cm⁻¹ in case of no citrate addition. Surprisingly, median uptake rates were mostly independent of the bulk soil stocks, but the P‐fertilized soil revealed a strong tailing towards higher values. This indicates the occurrence of hot P spots in soils. Citrate addition increased P yields only in the higher concentration but not in the forest soil. The order of magnitude of MD uptake rates from the soil samples matched root‐length related uptake rates from other studies. The micro‐radial citrate release in MD reflects the processes controlling phosphate mobilization in the rhizosphere better than measurements based on “flooding” of soil samples with citric acid in batch experiments. Important challenges in MD with phosphate are small volumes of dialysate with extremely low concentrations and a high variability of results due to soil heterogeneity and between‐probe variability. We conclude that MD is a promising tool to complement existing P‐analytical procedures, especially when spatial aspects or the release of mobilizing substances are in focus.