Qualitative and quantitative analysis of water‐soluble root exudates in relation to plant species and development

The composition of root‐derived substances is of great importance for the understanding of processes in the rhizosphere. Therefore, methods allowing a comprehensive collection and chemical analysis of the organic root exudates are necessary. In this study, we compare different methods with regard to their suitability to collect and characterize root exudates. Because the percolation or water logging method failed to quantitatively extract root exudates, a dipping method was developed which allowed an almost complete sampling of coldwater‐soluble root exudates. By ¹⁴CO₂ labeling of the shoots the composition of root exudates was found to be influenced by plant species and growth stage. In comparison to pea plants maize plants had a higher share of carboxylic acids and a lower share of sugars. Younger maize plants exuded considerably higher amounts of ¹⁴C labeled organic substances per g root dry matter than older ones. During plant development the relative amount of sugars decreased at the expense of carboxylic acids. The described methods are well suited for the elucidation of the influence of growth factors on root exudation.     

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.     

Phosphorus Influx and Root‐Shoot Relations as Indicators of Phosphorus Efficiency of Different Crops

Abstract

The large variation in phosphorus acquisition efficiency of different crops provides opportunities for screening crop species that perform well on low phosphorus (P) soil. To explain the differences in P efficiency of winter maize (Zea mays L.), wheat (Triticum aestivum L.), and chickpea (Cicer arietinum L.), a green house pot experiment was conducted by using P‐deficient Typic ustochrept loamy sand soil (0.5 M NaHCO₃‐extractable P 4.9 mg kg^?1, pH 7.5, and organic carbon 2.7 g kg^?1) treated with 0, 30, and 60 mg P kg^?1 soil. Under P deficiency conditions, winter maize produced 76% of its maximum shoot dry weight (SDW) with 0.2% P in shoot, whereas chickpea and wheat produced about 30% of their maximum SDW with more than 0.25% P in shoot. Root length (RL) of winter maize, wheat, and chickpea were 83, 48, and 19% of their maximum RL, respectively. Considering relative shoot yield as a measure of efficiency, winter maize was more P efficient than wheat and chickpea. Winter maize had lower RL/SDW ratio than that of wheat, but it was more P efficient because it could maintain 2.2 times higher P influx even under P deficiency conditions. In addition, winter maize had low internal P requirement and 3.3 times higher shoot demand (i.e., higher amount of shoot produced per cm of root per second). Even though chickpea had 1.2 times higher P influx than winter maize, it was less P efficient because of few roots (i.e., less RL per unit SDW). Nutrient uptake model (NST 3.0) calculations satisfactorily predicted P influxes by all the three crops under sufficient P supply conditions (C_Li 48 µM), and the calculated values of P influx were 81–99% of the measured values. However, in no‐P treatment (C_Li 3.9 µM), under prediction of measured P influx indicated the importance of root exudates and/or mycorrhizae that increase P solubility in the rhizosphere. Sensitivity analysis showed that in low P soils, the initial soil solution P concentration (C_Li) was the most sensitive factor controlling P influx in all the three crops.     

Amino acids exuded from axenic roots of lettuce and white lupin seedlings exposed to different cadmium concentrations

The objective of this study was to evaluate if amino acids in roots and/or in root exudates play a role in cadmium (Cd) stress. Lettuce (Lactuca sativa L. cv. Reine de Mai) and white lupin (Lupinus albus L. cv. lublanc) were grown for 19 to 21 days with axenic roots in a hydroponic system. After treatment with various concentrations of Cd (0, 0.01, 0.1, 1, 10, and 100 μM Cd) per nine days, roots and root exudates were collected. The stress did not result in significant dry weight (DW) differences between Cd‐treated and control plants, but Cd induced decreases in relative water content (RWC) and water potential (Pm). Amino acid levels and carbon (¹⁴C) incorporation into amino acids increased at low Cd concentrations in roots. However, 100 μM Cd induced a decrease of amino acid levels and an equally significant reduction of ¹⁴C incorporation, suggesting a decreased plant metabolism. Moreover, a higher Cd concentration induced increased levels of specific amino acids, for instance asparagine and lysine in lettuce and asparagine and hydroxylysine in lupin roots. Amino acids in root exudates corresponded less than 1% of the amounts found in root cells suggesting that amino acids could not be the major Cd chelators. Amino acid accumulation in root exudates differed than that found in roots except for asparagine. In conclusion, Cd induces in the root and root exudates increased levels of specific amino acids, such as Asn, Lys and HLys similarly to other environmental stresses. Although the amino acids could not participate in Cd chelation, lysine and its derivatives, such as hydroxylysine, could be used as stress markers for Cd in higher plants.     

Is occluded phosphate plant‐available?

Background: The phosphate concentration of the soil solution is generally low, allowing sufficient plant nutrition only for a few days. Therefore, supply from various fractions of bound phosphate is essential to meet plant demand. It is known that plants have developed strategies to acquire phosphorus (P) from phosphates adsorbed on clay minerals or oxides, from organically bound phosphates, and from calcium phosphates. However, it is generally assumed that occluded phosphate is not plant‐available. Results: In a pot experiment, two plant species, namely maize (Zea mays L.) and white lupin (Lupinus albus L.), differing in acquisition efficiency, were used to investigate whether Al oxide‐occluded and Fe oxide‐occluded phosphates can be acquired. Artificially prepared Al oxide‐occluded phosphate or Fe oxide‐occluded phosphate, respectively, was added to a subsoil low in available phosphates. It is shown that both plant species were not able to acquire P from Al oxide‐occluded phosphate. Also, maize was incapable of using Fe oxide‐occluded phosphate. In contrast, white lupin took up significant amounts of P from Fe oxide‐occluded phosphate. Conclusion: It is concluded that the strategy to form cluster roots together with their reducing power may allow white lupin to destabilize Fe oxides that occlude phosphates and to mine the soil for this additional phosphate fraction.     

Cu‐, Zn‐ und Cd‐Aneignungsvermögen von zwei Spinatgenotypen in Abhängigkeit von der P‐Versorgung und Wurzelexsudation

Cu, Zn, and Cd acquisition by two spinach cultivars depending on P nutrition and root exudation Within a spectrum of 11 spinach cultivars (cvs) differences in the Cu, Zn, and Cd contents of shoots had been noticed. The aim of this study was therefore to analyze in more detail the acquisition of Cu, Zn, and Cd by the most differing cultivars (Tabu and Monnopa) in dependence on P nutrition. The plants were grown in a low phosphorus Luvisol (pH 6.3; total contents Cu: 89, Zn: 297, Cd: 2.4 mg kg^—1) with two phosphorus levels in pots under natural conditions. For the determination of inflow, root length/shoot weight ratio and of the Cu, Zn, and Cd concentration in the soil solution (rhizosphere) plants were harvested 26 and 40 days after sowing. Root exudation of organic acids of the two cvs was measured 35 days after growing in quartz sand with different P supply. Both cultivars responded to P fertilizer by doubling their shoot weight. With increased P supply (0.68—0.77% P in shoot‐DM) both cultivars showed similar heavy metal contents in the shoot resulting from similar root length/shoot weight ratios (RSR) and net uptake rates of the three elements as well as the same element concentrations in the rhizosphere soil solution. Under P deficiency, however, cv. Tabu (0.52% P in shoot‐DM) showed in comparison with cv. Monnopa (0.48% P) higher Cu, Zn, and Cd contents of shoots although its RSR was smaller than that of cv. Monnopa. However, the inflow for Cu was higher and for Zn and Cd significantly higher compared with cv. Monnopa. This result of cv. Tabu corresponded with higher concentrations of Cu, Zn, and Cd of its rhizosphere soil solution, and its higher exudation rates of oxalate, citrate, and malate (3.9; 1.0; 0.7 nmol cm^—1 h^—1). The corresponding values for cv. Monnopa were: 1.7; 0.3; 0.4 nmol cm^—1 h^—1. The mobilization of Cu, Zn, and Cd by the excreted organic acids seems to be responsible for the higher Cu, Zn, and Cd inflow of cv. Tabu.     

Differences in Root Exudation Among Phosphorus‐Starved Genotypes of Maize and Green Gram and Its Relationship with Phosphorus Uptake

Abstract

Availability of phosphorus (P) in soil and its acquisition by plants is affected by the release of high and low molecular weight root exudates. A study was carried out to ascertain the qualitative and quantitative differences in root exudation among the genotypes of maize (Zea mays L.) and green gram (Vigna radiata L.) under P‐stress. Results showed that both inter‐ and intra‐species differences do exist among maize and green gram in terms of root exudation, P uptake, and shoot and root P content. In general, green gram, a legume crop, had greater root exudation compared to maize. However, the amino acid content of the total root exudates in maize was two‐fold as compared to green gram. The maize and green gram genotypes possessed genetic variability in root exudation. Irrespective of the species or genotypes, a positive relationship was found among P uptake rates, total root exudation, and shoot and root ³²P content. The amount of sugars and amino acid present in the root exudates of P‐starved seedlings also add to the variation in P uptake efficiency of genotypes.     

Root Exudates of Rice Cultivars Affect Rhizospheric Phosphorus Dynamics in Soils with Different Phosphorus Statuses

Exudation of organic acids by the roots of three rice cultivars grown in three soils of different phosphorus (P) statuses, and their impacts on the rhizospheric P dynamics and P uptake by the rice plants, were investigated. Quantum root exudates from all the rice cultivars were significantly greater at 21 days after transplantation than at panicle initiation or flowering stages. Malic acid was the most predominant organic acid present in the rice root exudates (10.3 to 89.5 μmol plant^?1 d^?1), followed by tartaric, citric, and acetic acids. Greater exudation of organic acids from rice grown in P-deficient soil by all the rice cultivars suggested response of rice plant to P stress. Results indicate that the release of organic acids in the root exudates of rice plants can extract P from strongly adsorbed soil P fraction, thereby increasing native soil P utilization efficiency and ensuring adequate P nutrition for the growing rice plants.     

Mineral‐nutrient synergism and dilution responses to nitrogen fertilizer in field‐grown maize

Nitrogen (N)‐fertilizer applications to field‐grown maize may result in a dilution response whereby essential mineral‐element concentrations in shoots would decrease as shoot‐dry‐matter accumulation increased. To investigate this, the effect of N‐fertilizer treatments (no N or fertilizer rate based upon 5.3 or 8.5 t ha^–1 yield goal) on maize (Zea mays L.) shoot dry weight and shoot mineral concentrations (N, P, K, S, Mg, Ca, and Mn) at the sixth leaf (V6), twelfth leaf (V12), and tassel (VT) development stages were investigated in a 2‐year study conducted at Brookings, South Dakota (USA). With increasing N‐fertilizer application rates, shoot dry weight was greater and shoot P and K concentrations decreased. A possible explanation of this dilution response is that planting‐time P and K fertilizers, which were applied in a band near the seed furrow, may have enhanced the uptake of P and K in a manner that was independent of N‐fertilizer treatments. Increased shoot‐dry‐weight production due to the application of N fertilizers, if P and K uptake were similar across N‐fertilizer treatments, would lead to decreased shoot P and K concentrations in N‐sufficient compared with N‐deficient plants. Conversely, N‐fertilizer‐induced increases in shoot dry weight were accompanied by increased shoot concentrations of N, Ca, and Mn. This synergistic response between dry‐weight accumulation and shoot N concentration was present at all leaf developmental stages studied, while that for Ca was present only at VT. Thus, N fertilizer applications that increase shoot dry weight can affect the dilution and synergistic responses of specific mineral nutrients in maize shoots. Crop developmental stage as well as the location of these specific mineral nutrients in the soil profile might play important roles in mediating these responses.     

Root exudation of sugars,amino acids,and organic acids by maize as affected by nitrogen,phosphorus, potassium,and iron deficiency

Root exudates play a major role in the mobilization of sparingly soluble nutrients in the rhizosphere. Since the amount and composition of major metabolites in root exudates from one plant species have not yet been systematically compared under different nutrient deficiencies, relations between exudation patterns and the type of nutrient being deficient remain poorly understood. Comparing root exudates from axenically grown maize plants exposed to N, K, P, or Fe deficiency showed a higher release of glutamate, glucose, ribitol, and citrate from Fe‐deficient plants, while P deficiency stimulated the release of γ‐aminobutyric acid and carbohydrates. Potassium‐starved plants released less sugars, in particular glycerol, ribitol, fructose, and maltose, while under N deficiency lower amounts of amino acids were found in root exudates. Principal‐component analysis revealed a clear separation in the variation of the root‐exudate composition between Fe or P deficiency versus N or K deficiency in the first principal component, which explained 46% of the variation in the data. In addition, a negative correlation was found between the amounts of sugars, organic and amino acids released under deficiency of a certain nutrient and the diffusion coefficient of the respective nutrient in soils. We thus hypothesize that the release of dominant root exudates such as sugars, amino acids, and organic acids by roots may reflect an ancient strategy to cope with limiting nutrient supply.     

Determination of organic acids in tissues and exudates of maize, lupin, and chickpea by high-performance liquid chromatography-tandem mass spectrometry

This article describes a fast and simple methodology for the extraction and determination of organic acids in tissues and root exudates of maize, lupin, and chickpea by LC/MS/MS. Its main advantage is that it does not require sample prepurification before HPLC analysis or sample derivatization to improve sensibility. The results obtained showed good precision and accuracy, a recovery close to 100%, and no significant matrix effect. Moreover, the sensibility of the method is in general better than that of previously described methodologies, with detection limits between 15 and 900 pg injected.     

Phosphatversorgung von Sommerweizen (Triticum aestivum L.) in Mischkultur mit Weißer Lupine (Lupinus albus L.)

Phosphorus nutrition of spring wheat (Triticum aestivum L.) in mixed culture with white lupin (Lupinus albus L.). Spring wheat (Triticum aestivum L. ?Schirokko”?) and white lupin (Lupinus albus L.) were grown in mixed culture in Mitscherlich pots with 20 kg of soil in a green house. The soil used was a B_t of a Parabraunerde-Pseudogley from loess low in available P and limed from pH 4.6 to pH 6.5. Phosphorus was added as phosphate rock. In half of the pots cylinders of stainless steel screen prevented intertwining of the roots of the plant species. Independent of P addition, white lupin had higher dry matter production and P uptake than wheat, even although wheat had thinner roots and higher root densities than lupin, factors which favour the utilization of soil and fertilizer P. The higher P efficiency of white lupin was due to higher P uptake rates per unit root length mainly through mobilization of P especially in the rhizosphere of the proteoid roots. When the roots of the two species were allowed to intertwine, shoot dry matter production of wheat was nearly double because of improved tillering. Higher P concentrations and a more than 2-fold higher P uptake indicated that the increase in dry matter production of wheat was due to improved P nutrition. Nitrogen concentrations, however, remained unaffected at sufficient levels. An increased P uptake rate per unit root length was responsible for the better utilization of P by wheat, rather than the increase in total root length, due to the extended root volume. White lupin was able to mobilize P in the rhizosphere in excess of its own requirements. Thus mobilized P may be available to less P-efficient plants grown in mixed culture.     

铅胁迫下小花南芥与玉米间作对根系分泌物 有机酸的影响

为了揭示Pb胁迫对间作和单作的超累积植物和作物根系分泌低分子有机酸的影响,研究设置400 mg·L?1Pb胁迫,采用水培曝气法试验,以玉米和小花南芥单作为对照处理,研究Pb胁迫下玉米和小花南芥间作对植物根系形态、根系分泌有机酸及Pb吸收的影响。结果表明:与单作相比,间作小花南芥情况下,玉米根系分泌物检测到乳酸;玉米分根条数、根表面积和根密度与单作相比分别增加60%、15%和42%,地下部和地上部干重生物量分别增加108%和75%,玉米地下部Pb含量下降44%;与单作相比,间作玉米条件下,小花南芥根系分泌物检测到乙酸和乳酸,小花南芥根系分泌物量与单作相比增加103%~1 700%,小花南芥地下部和地上部Pb累积量分别比单作增加49%和75%,转运系数增加22%。相关分析结果表明,单作小花南芥只有地上部Pb累积量与草酸显著相关,而间作小花南芥地下部和地上部Pb累积量与草酸、柠檬酸和苹果酸显著相关。研究表明超富集植物小花南芥与玉米间作体系,根系分泌的有机酸改变了Pb在小花南芥和玉米体内的累积特征,促进超累积植物小花南芥累积Pb,减少农作物玉米植株体内Pb含量。Pb胁迫下超累积植物小花南芥与玉米间作是一种可行的修复模式。     

Phosphorus uptake and rhizosphere properties of intercropped and monocropped maize, faba bean, and white lupin in acidic soil

Little information is available on phosphorus (P) uptake and rhizosphere processes in maize (Zea mays L.), faba bean (Vicia faba L.), and white lupin (Lupinus albus L.) when intercropped or grown alone in acidic soil. We studied P uptake and soil pH, carboxylate concentration, and microbial community structure in the rhizosphere of maize, faba bean, and white lupin in an acidic soil with 0–250 mg P (kg⁻¹ soil) as KH₂PO₄ (KP) or FePO₄ (FeP) with species grown alone or intercropped. All plant species increased the pH compared to unplanted control, particularly faba bean. High KP supply (>100 mg P kg⁻¹) significantly increased carboxylate concentration in the rhizosphere of maize. The carboxylate composition of the rhizosphere soil of maize and white lupin was significantly affected by P form (KP or FeP), whereas, this was not the case for faba bean. In maize, the carboxylate composition of the rhizosphere soil differed significantly between intercropping and monocropping. Yield and P uptake were similar in monocropping and intercropping. Monocropped faba bean had a greater concentration of phospholipid fatty acids in the rhizosphere than that in intercropping. Intercropping changed the microbial community structure in faba bean but not in the other corps. The results show that P supply and P form, as well as intercropping can affect carboxylate concentration and microbial community composition in the rhizosphere, but that the effect is plant species-specific. In contrast to previous studies in alkaline soils, intercropping of maize with legumes did not result in increased maize growth suggesting that the legumes did not increase P availability to maize in this acidic soil.     

Organic acid excretion from roots: a plant mechanism for enhancing phosphorus acquisition,enhancing aluminum tolerance,and recruiting beneficial rhizobacteria

The beneficial effects of organic acids (OAs) excretion from plant roots were first proposed as being associated with the mechanism of superior phosphorus utilization by the cluster roots of white lupin (Lupinus albus L.), and these effects are now widely accepted as pleiotropic effects associated with stress tolerance of plants. Excreted OAs detoxify rhizotoxic aluminum, recruit beneficial bacterium for induced systemic resistance, and modify root architecture to enhance phosphorus starvation. OA excretion is probably optimized in the carbon economy and is coordinately regulated with other traits that additively confer each stress factor. Here we present an overview of the molecular physiology of OA excretion from roots, how plants activate OA excretion, and how this excretion can be managed as a specific response.     

Phosphorus bioavailability of sewage sludge‐based recycled fertilizers in an organically managed field experiment

Recycled phosphorus (P) fertilizers from sewage sludge can contribute to the ongoing effort of closing the P cycle. Five recycled P fertilizers (Struvite SSL, Struvite AirPrex®, P‐RoC®, Pyrolysis coal, and Na‐SSA) were tested for their P availability in a two‐year field experiment with maize. The experiment was conducted on an organic certified research station at soil pH 6.5. Other P fertilizer treatments included: phosphate rock (PR), compost, and an unfertilized control. In addition, the rhizobacteria strain Bacillus sp. Proradix (Proradix®) was applied to test its ability to increase P bioavailability. Each year, shoot DM and P offtake of maize was measured and P use efficiency of the tested fertilizers was calculated. No significant differences in shoot DM were found among fertilized treatments and the unfertilized control in both years of experiment. Fertilization with recycled fertilizers increased P offtake by between 0% (Na‐SSA) and 27.5% (Struvite SSL) compared to the unfertilized control. Rhizobacteria application led to an increase in P offtake of maize from 25.9 to 38.7 kg P ha^?1 when combined with PR fertilization in the year of fertilizer application, while no significant effect was found for the recycled fertilizers. Some of the tested recycled fertilizers from urban waste water can be considered as effective fertilizers for their use in organic agriculture.     

METABOLITE PROFILING OF SHOOT EXTRACTS,ROOT EXTRACTS,AND ROOT EXUDATES OF RICE PLANT UNDER PHOSPHORUS DEFICIENCY

We applied a metabolite profiling technique to investigate root exudates under phosphorus (P) deficiency. Oryza sativa was grown in culture solution containing three P levels (0, 1, and 8 mg P L^?1). Shoot extracts, root extracts, and root exudates were obtained from 18 and 23-day-old plants and their metabolites were determined by capillary electrophoresis/time-of-flight mass spectrometry. Eighty, 90, and 65 metabolites were identified in shoot extracts, root extracts, and root exudates, respectively. Sixty-three to eighty-four percent of the metabolites were exuded to the rhizosphere. More than 33% of the metabolites in the root exudates showed higher concentration at low P than at high P. On the other hand, only 14% of the metabolites in the root extracts showed lower concentration at low P than at high P. These results suggest that rice roots actively release many metabolites in response to P deficiency.     

Metabolite profiling of shoot extract,root extract,and root exudate of rice under nitrogen and phosphorus deficiency

ABSTRACT

Root exudate is derived from plant metabolites and its composition is affected by plant nutrient status. A deficiency of mineral nutrients, such as nitrogen (N) and phosphorus (P), strongly affects the type and amount of plant metabolites. We applied a metabolite profiling technique to investigate root exudates of rice plants under N and P deficiency. Oryza sativa was grown in culture solution containing two N levels (0 and 60 mg N L^?1) or two P levels (0 and 8 mg P L^?1). Shoot extracts, root extracts, and root exudates were obtained from the rice plants 5 and 15 days after transplanting and their metabolites were determined by capillary electrophoresis/time-of-flight mass spectrometry. Shoot N concentration and dry weight of rice plants grown at ?N level were lower than those of plants grown at +N level. Shoot P concentration and dry weight of rice plants grown at ?P level were lower than those of plants grown at +P level. One hundred and thirty-two, 127, and 98 metabolites were identified in shoot extracts, root extracts, and root exudates, respectively, at the two N levels. One hundred and thirty-two, 128, and 99 metabolites were identified in shoot extracts, root extracts, and root exudates, respectively, at the two P levels. Seventy-seven percent of the metabolites were exuded to the rhizosphere. The concentrations of betaine, gamma-aminobutyric acid, and glutarate in root exudates were higher at both ?N and ?P levels than at their respective high levels. The concentration of spermidine in root exudates was lower at both ?N and ?P levels than at their respective high levels. The concentrations of the other metabolites in root exudates were affected differently by plant N or P status. These results suggest that rice roots actively release many metabolites in response to N and P deficiency.     

一种新型根系分泌物收集装置与收集方法的介绍

根系分泌物在养分活化、改善环境胁迫方面具有重要作用,很多科技工作者对根系分泌物的研究表现出极大兴趣,取得了一系列进展。但土壤栽培条件下,根系分泌物收集是一个难点。本文介绍了一种新型根系分泌物的收集装置与收集方法。该装置由根系生长箱和分泌物收集箱组成,植物在生长箱土壤中生长,通过定向引导作用,根系从生长箱穿过琼脂层进入收集箱中生长,待收集箱内积累一定根系后,通过淋洗收集箱内的介质,实现根系分泌物收集。研究发现,利用该装置收集分泌物,植物总根尖数的90%分布在收集箱。外源有机酸加样回收率可达70%以上。土壤栽培条件下,随生长时期延长,大豆有机酸分泌量逐渐增加,苹果酸分泌量高于柠檬酸。而且土壤栽培条件下大豆柠檬酸和苹果酸分泌量是溶液栽培时的11.4倍和6.7倍。上述研究表明,该装置可以用于土壤栽培条件下根系有机酸的分泌研究。     

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.