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.     

Ability of various water‐insoluble fertilizers to supply available phosphorus in hydroponics to plant species with diverse phosphorus‐acquisition efficiency: Involvement of organic acid accumulation in plant tissues and root exudates

Previous studies describe the suitability of a new type of phosphorus (P) fertilizer, called “rhizosphere‐controlled fertilizer” (RCF), to supply available P to plants while reducing soil phosphorus fixation. In order to explore the involvement of organic acid root exudation in P uptake from RCF, we investigated the relationship between shoot and root P concentrations, and the concentration of the main polycarboxylic organic acids in roots, shoots, and plant exudates. Plant species with different P‐acquisition efficiency (low: maize; medium: chickpea; high: lupin) were grown in hydroponics with three different P fertilizers: The water‐insoluble P fraction of RCF (RCF); Phospal, a slow‐release source of phosphate composed of calcium and aluminum phosphates (PH); monopotassiumphosphate (KP), and a control treatment without P (P–). RCF was as efficient as KP in supplying P to plants in the case of chickpea and lupin, and slightly less efficient than KP in maize. However, P from PH was not available for maize and less available compared to KP and RCF in chickpea and lupin. This variation reflects the different efficiencies in P acquisition for the three plant species. Except in the case of maize, plants receiving KP presented the lowest concentration of organic acids in roots and exudates, while those plants suffering severe P deficiency (P– and PH) showed the highest organic acid concentration. However, RCF had a high concentration of organic acids in roots and exudates, as well as a high P concentration in the shoot indicating that P uptake from RCF is enhanced due to root release and action of specific organic acids.     

The quantitative effect of chemical phosphate mobilization by carboxylate anions on P uptake by a single root. II. The importance of soil and plant parameters for uptake of mobilized P

Model calculations were made in order to quantify the effect of carboxylate excretion on phosphate (P) uptake by a single root. The uptake of chemically mobilized P increased exponentially with increasing concentration of adsorbed citrate or oxalate in soil because of the exponential relationship between adsorbed carboxylate and the solubilizing effect of carboxylate on P. The effect of local citrate excretion compared with uniform citrate excretion along the whole root was also calculated. Local exudation increased the uptake of chemically mobilized P because the higher concentration of citrate increases the solubilization of P. Additionally the effect of citrate excretion by root clusters e.g. proteoid roots was evaluated. Uptake of chemically mobilized P by root clusters was much higher than that of single roots, especially if the ratio of P buffering to citrate buffering was high. This is often the case in P fixing soils where by definition P buffering is high and citrate buffering is low because of the short time of reaction between root excreted citrate and rhizosphere soil. The reason for the superiority of cluster roots lies in the fact that most of the mobilized P is transported away from a single root to be absorbed by neighbouring roots in the clusters. This phenomenon demonstrates the strong ecological significance of cluster roots in relation to nutrient mobilization. The calculations on the effect of oxalate excretion by sugar beet roots on the uptake of mobilized P show that under P fixing conditions the influx of mobilized P will exceed that of P transported by diffusion to the root surface by a factor of 1.5—6.0.     

Plant responses to drought and phosphorus deficiency: contribution of phytohormones in root‐related processes

Environmental stresses are one of the most limiting factors in agricultural productivity. A large portion of the annual crop yield is lost to pathogens (biotic stress) or the detrimental effects of abiotic‐stress conditions. There are numerous reports about chemical characterization of quantitatively significant substrate fluxes in plant responses to stress factors in the root‐rhizosphere system, e.g., nutrient mobilization, heavy‐metal and aluminum immobilization, or establishment of plant‐growth‐promoting rhizobacteria (PGPR) by exudation of organic anions, phytosiderophores, or carbohydrates into the soil, respectively. The hormonal regulation of these responses is not well understood. This paper highlights this complex process, stressing the involvement of phytohormones in plant responses to drought and phosphorus deficiency as examples. Beside ethylene, abscisic acid (ABA) plays an important role in drought‐stress adaptation of plants. This hormone causes morphological and chemical changes in plants, ensuring plant survival under water‐limited conditions. For example, ABA induces stomata closure, reduction in leaf surface, and increase in root : shoot ratio and, thus, reduction in transpiration and increase in soil volume for water uptake. Furthermore, it supports water uptake in soil with decreasing water potential by osmotic adjustment. Suitability of hormonal parameters in the selection for improving stress resistance is discussed. Auxins, ethylene, and cytokinins are involved in morphological adaption processes to phosphorus (P) deficiency (increase in root surface, e.g., by the formation of more dense root hairs or cluster roots). Furthermore, indole‐3‐acetic acid increases root exudation for direct and indirect phosphorus mobilization in soil. Nevertheless, the direct use of the trait “hormone content” of a particular plant organ or tissue, for example the use of the drought‐stress‐induced ABA content of detached leaves in plant breeding for drought‐stress‐resistant crops, seems to be questionable, because this procedure does not consider the systemic principle of hormonal regulation in plants.     

The Function of a Maize-Derived Phosphoenol pyruvate Carboxylase (PEPC) in Phosphorus-Deficient Transgenic Rice

Transgenic rice ( Oryza sativa L., a C₃ plant) lines carrying a complete phospho enol pyruvate carboxylase (PEPC) gene from maize (a C₄ plant) were tested for their performance in terms of organic acid synthesis and organic acid exudation into the rhizosphere under phosphorus (P)-deficient conditions. High PEPC activity increased the fraction of photosynthetically fixed carbon allocated to the organic acid pool, and P deficiency enhanced oxalate exudation from the roots of the transgenic plants. There was no evidence that the transformed PEPC was involved in internal P recycling in the plant. However, the root PEPC activity was positively correlated with the oxalate exudation and negatively correlated with the root P concentration, and a higher root PEPC activity led to a higher oxalate exudation. Thus, it is suggested that C₄-PEPC transgenic rice plants had acquired the ability to exude oxalate, which enhanced their capacity to adapt to low P soil conditions.     

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.     

Phosphorus uptake by rice from soil that is flooded, drained or flooded then drained

To investigate the mechanisms by which rice plants growing in alternately flooded and drained soils absorb soil phosphate, we grew rice in moist, flooded and flooded then moist soils, and compared the measured uptake of phosphorus (P) with that calculated using a mathematical model of uptake allowing for solubilization by various means. The theory and equations for the model are given, together with a method for solving diffusion equations near roots in a root system of increasing density. The diffusion coefficients and buffer powers of P in the soil under the different water regimes are measured by following diffusion of P to a resin sink, and the parameters describing solubilization are estimated from previously published results. In all the water regimes studied, the plants relied upon solubilization for most of their P. The roots were not mycorrhizal, as they will often not be in intermittently flooded soils. In the flooded soil, uptake was three times that in the moist soil, and was consistent with solubilization by acidification caused by roots as a result of oxidation of iron and imbalance between the intake of cations and anions. In the moist soil, the uptake was consistent with solubilization by excretion of organic anions from the roots. In the flooded then moist soil, uptake declined sharply as the soil dried because P became immobilized in the soil. However, the final uptake was similar to that in the continuously moist soil, indicating that some of the immobilized P was re‐solubilized by roots, possibly by excretion of organic anions.     

Root Morphology,Proton Release,and Carboxylate Exudation in Lupin in Response to Phosphorus Deficiency

Narrow-leafed lupin (Lupinus angustifolius L.) is widely planted in infertile acidic soils where phosphorus (P) deficiency is one of the major limiting factors for plant growth. A hydroponic experiment was conducted to examine the morphological and physiological responses of roots of narrow-leafed lupin in response to altered P supply at 0, 1, 10, 25 or 75 μ M P as monopotassium phosphate (KH₂PO₄). Low P (P₀ and P₁) significantly decreased the plant biomass, but the supply of 10 μ M P was sufficient to produce similar plant biomass as the maximal P supply (P₇₅), indicating an efficient P acquisition by narrow-leafed lupin. Phosphorus deficiency did not enhance rates of carboxylate exudation and proton release by plant roots, indicating that carboxylate exudation and proton release are not the mechanisms for efficient P acquisition. In contrast, low P supply evidently modified the root morphology by increasing the primary root elongation, and developing a large number of cluster-like first-order lateral roots with dense root hairs, thus allowing efficient P acquisition by narrow-leafed lupin under low P supply.     

Mobilization of phosphorus contributes to positive rotational effects of leguminous cover crops on maize grown on soils from northern Nigeria

Previous research has demonstrated a positive rotational effect of tropical leguminous cover crops on maize growth on a luvisol from Nigeria. This effect could not be explained by a better N supply. The objective of the present work was to further clarify whether improved P nutrition has been a contributing factor. Nine cover crops and maize were studied in nutrient solution‐culture with 1 and 20 μM P and with NO₃‐N as N source for root physiological parameters that may affect P mobilization. Zea mays, Lablab purpureus, and Centrosema pubescens responded to P deficiency by higher rates of proton excretion. Clitoria ternatea excreted OH^— with only small differences due to P nutrition. At low P supply, Chamaecrista rotundifolia, Clitoria, and Centrosema had the highest exudation rates of organic acid anions, especially citrate and malate. A major difference between plant species was found in root‐surface acid phosphatase activity. Cajanus cajan expressed the highest phosphatase activity. V_max of P uptake increased markedly under P deficiency, particularly for maize. Compared to the other plant species Cajanus, Chamaecrista, and Clitoria were characterized by a greater capability to absorb P at low external P concentrations. The nine cover crops and maize were also grown in pot experiments using two soils from northern Nigeria low in available P. The ultisol from Jos had a high P fixation capacity and was more acidic than the alfisol from Zaria. All plant species were precultured in the first season at 100 (Zaria) and 250 (Jos) mg P per pot. In the subsequent season, maize was uniformly seeded into all pots containing the remaining roots and the incorporated shoot dry mass (according to 15 mg P per pot) of the pre‐crop. No P and 250 (Zaria) or 500 (Jos) mg P per pot were freshly applied. Maize growth and P uptake were enhanced after legumes in Zaria soil. Cajanus showed the highest residual effect, and also Clitoria, Chamaecrista, and Lablab showed effects superior to the mean. In Jos soil, Clitoria, Cajanus, and Lablab enhanced maize growth above average. Also, a highly significant positive correlation between P uptake and biomass production was obtained. These results indicate that enhancement of maize growth after leguminous cover crops, at the low P supply of the soil used, mainly depends on P mobilization capacity of the cover crop.     

Rhizosphere interactions between microorganisms and plants govern iron and phosphorus acquisition along the root axis - model and research methods

Iron and phosphorus availability is low in many soils; hence, microorganisms and plants have evolved mechanisms to acquire these nutrients by altering the chemical conditions that affect their solubility. In plants, this includes exudation of organic acid anions and acidification of the rhizosphere by release of protons in response to iron and phosphorus deficiency. Grasses (family Poaceae) and microorganisms further respond to Fe deficiency by production and release of specific chelators (phytosiderophores and siderophores, respectively) that complex Fe to enhance its diffusion to the cell surface. In the rhizosphere, the mutual demand for Fe and P results in competition between plants and microorganisms with the latter being more competitive due to their ability to decompose plant-derived chelators and their proximity to the root surface; however microbial competitiveness is strongly affected by carbon availability. On the other hand, plants are able to avoid direct competition with microorganisms due to the spatial and temporal variability in the amount and composition of exudates they release into the rhizosphere. In this review, we present a model of the interactions that occur between microorganisms and roots along the root axis, and discuss advantages and limitations of methods that can be used to study these interactions at nanometre to centimetre scales. Our analysis suggests mechanisms such as increasing turnover of microbial biomass or enhanced nutrient uptake capacity of mature root zones that may enhance plant competitiveness could be used to develop plant genotypes with enhanced efficiency in nutrient acquisition. Our model of interactions between plants and microorganisms in the rhizosphere will be useful for understanding the biogeochemistry of P and Fe and for enhancing the effectiveness of fertilization.     

含磷物质对铅耐性与敏感性植物生长及磷铅吸收的影响

将耐性植物芥蓝（Brassicacapitata）与敏感植物小白菜（Brassica chinenzis）暴露于含Pb的营养液中,通过添加磷灰石矿尾料（PR）、重过磷酸钙（TSP）及二者混合物（P＋T）处理,研究含磷物质对两种植物吸收Ph的影响以及Pb在植物根表面形态变化过程。结果显示：添加含磷物质降低了耐性植物芥蓝根部Pb的含量,但对地上部Ph吸收的影响不显著;对敏感植物小白菜来说,PR和P＋T处理的植物根部及地上部Pb的含量与对照基本一致,而TSP处理促进了Pb的吸收,表现出小白菜根部及地上部Pb的含量显著高于对照组。含磷物质添加诱导了Pb在植物根表面形成Pb5（PO4）3Cl、Pb5（PO4）3OH的沉淀,但这并没有直接导致两种植物对Pb吸收量的减少。     

铝胁迫下大豆根系脱落酸累积与柠檬酸分泌的关系研究

为探讨铝(Al)胁迫条件下脱落酸(ABA)调控植物根系有机酸分泌的机制,进行了ABA与Al诱导大豆根系柠檬酸分泌的关系试验。结果表明:1)外源ABA和ABA合成抑制剂fluridone分别提高和降低了Al诱导的大豆根尖ABA含量的增加,但对根系柠檬酸分泌量均无影响,ABA对根系内源柠檬酸含量和柠檬酸合成酶的活性也没有影响;2)分根试验表明,与Al直接接触的根部(Part A)内源ABA含量发生变化,且有柠檬酸的分泌,而不与Al直接接触的根部(Part B)内源ABA含量也发生变化,但没有柠檬酸分泌;3)Al胁迫下,大豆耐Al基因型柠檬酸分泌量远高于敏感基因型,但二者的内源ABA含量却没有差异;4)30μmol AlCl3处理,在0~12 h柠檬酸分泌速率和内源ABA含量随Al处理时间增加而增加,去除Al胁迫时(12~18 h),柠檬酸分泌速率继续增加,但内源ABA含量则迅速下降。综合以上结果,推测ABA不是通过提高Al诱导柠檬酸分泌来调控大豆耐Al性。     

Differential mobilization of P in the maize rhizosphere by citric acid and potassium citrate

The release of organic acid anions from plant roots into soil has been hypothesized to be a mechanism for enhancing phosphorus availability in the rhizosphere. Although these compounds are excreted from the cytoplasm as organic acid anions (e.g. citrate, malate), when the H⁺-ATPase is also upregulated there is evidence to suggest that they enter the soil as organic acids (e.g. citric acid, malic acid). The aim of this study was to evaluate the role of citric acid (H-citrate) and potassium citrate (K-citrate) in the mobilization and plant uptake of P from two acid soils contrasting in their P availability. Our results indicated that the mobilization of P from a labelled patch of soil was soil type dependent, was controlled by its intrinsic P status, and that more P was made available by K-citrate than H-citrate. Similarly, the uptake of ³³P from the rhizosphere by Zea mays L. was greatest in the presence of K-citrate in comparison to H-citrate. However, a significant increase in shoot ³³P content was only observed in the more acidic soil with high P sorption potential (Haplic podzol) while no significant increase was observed in the less acidic soil with low P sorption potential (Eutric cambisol). We conclude that the chemical form of organic acid anion excretion may have a significant impact on its P mobilization capability. The contrasting results with the two acid soils indicate that organic acids may not provide a universal mechanism for enhancing P uptake from soil.     

铝胁迫下萹蓄与饭豆根系有机酸分泌特性比较

以蓄和饭豆为试验材料,从对Al胁迫的响应时间、蛋白质合成抑制剂的影响、体内有机酸含量变化及阴离子通道抑制剂的影响等方面,比较研究了Al诱导根系分泌有机酸的差异,以进一步明确铝(Al)诱导植物根系有机酸分泌的过程。结果表明,萹蓄根系在Al胁迫后30.min内分泌出草酸,而Al胁迫至少4.h后饭豆根系才开始分泌柠檬酸;蛋白质合成抑制剂环己亚酰胺(Cycloheximide,CHM)不影响萹蓄根系草酸的分泌,但抑制了84%Al诱导的饭豆根系柠檬酸的分泌,表明前者不需要新蛋白质的诱导合成,却是后者所必需的。Al处理不改变萹蓄根尖草酸的含量,但明显提高了饭豆根尖柠檬酸的含量;阴离子通道抑制剂苯甲酰甲醛(Phenylglyoxal,PG)和蒽-9-羧酸(Anthrancene-9-carboxylic.acid,A-9-C)分别有效抑制Al诱导萹蓄根系草酸的分泌及饭豆根系柠檬酸的分泌,再次证明两者有机酸的分泌是通过某种被Al所诱导或激活的阴离子通道所实现的。     

Differential Aluminum Resistance and Organic Acid Anions Secretion in Triticale

Abstract

Triticale (X Triticosecale Wittmack), a hybrid of wheat and rye, shows a high degree of aluminum (Al) tolerance, but variation in Al resistance between cultivars does exist. The mechanisms responsible for differential Al resistance in 10 triticale cultivars were investigated in this study. Triticale roots secreted both malate and citrate in response to Al stress. The amount of organic acid anions secreted was correlated positively to the relative root elongation (an index for Al resistance) and negatively to the Al content in root apices under Al stress, suggesting that the secretion of malate and citrate seems to be involved in the exclusion of Al from root tip. The Al‐induced secretion of malate and citrate was characterized using an Al‐resistant cultivar (ZC 237) and an Al‐sensitive cultivar (OH 1621). Root elongation was significantly inhibited in both ZC 237 and OH 1621 after 24 h of exposure to 30, 50, or 100 µM Al but was more strongly in OH 1621 than in ZC 237 at all Al concentrations tested. A marked lag phase (3 h) between the addition of Al and the secretion of organic acid anions was observed in both triticale cultivars, and the secretion increased with increasing external Al concentration. The two anion‐channel inhibitors, phenylglyoxal and niflumic acid, significantly inhibited the secretion of malate and citrate in ZC 237, with the degree of the inhibition of niflumic acid greater than that of phenylglyoxal. The Al‐induced secretion of malate and citrate decreased to a very low level at low temperature (4°C) in both cultivars. These results indicate that Al‐induced malate and citrate secretion from roots play important roles in excluding Al and thereby detoxifying Al in triticale. The Al‐induced organic acid anions were inhibited by anion‐channel inhibitors and were dependent on temperature.     

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.     

Organic acids in root exudates from Picea abies seedlings influenced by mycorrhiza and aluminum

Organic acid anions exuded from roots of Picea abies (Norway spruce) seedlings grown on glass beads in the presence and absence of mycorrhiza (Laccaria bicolor) and aluminum (Al) at pH 3.9 were measured. We wanted to test if the roots exuded more organic acid anions when exposed to Al and if mycorrhization influenced the exudation. Oxalate was exuded in far higher amounts than any other organic acid anion, with a maximum rate of 1.7 nmol (mg root DW)^–1 d^–1. Mycorrhizal roots exuded significantly more oxalate than nonmycorrhizal roots. The presence of Al did not enhance oxalate exudation. We conclude that the oxalate exuded constitutively by Picea abies/Laccaria bicolor may lead to rhizosphere oxalate concentrations that are relevant for Al resistance.     

P and N deficiency change the relative abundance and function of rhizosphere microorganisms during cluster root development of white lupin (Lupinus albus L.)

We studied microbe-plant interactions of white lupin, a cluster root-forming plant, under low P and N conditions to examine increased nutrient acquisition by plants either by a shift to a more specialized microbial community or changes in microbial enzyme production. White lupin plants were grown in rhizoboxes filled with either P- or N-deficient soil; fertilized soil was used as control. After cultivation of plants in a glasshouse for 41 d, plant growth (shoot and roots) and P and N accumulation in shoots were measured. Microbial functions were analyzed by P- and N-cycling enzymes. The microbial community structure was estimated by fingerprinting (denaturing gradient gel electrophoresis) and sequencing techniques. P deficiency induced the released citrate and acid phosphomonoesterases from cluster roots and stimulated the production of microbe-derived alkaline phosphomonoesterase in the rhizosphere. P deficiency decreased microbial diversity in the cluster root rhizosphere. Increased relative abundance of Burkholderiales in the rhizosphere of P deficient plants might be responsible for the degradation of different organic P fractions such as phytates. N deficiency induced an increase of the number of nodules and P concentration in shoot as well as roots of white lupin. We clarified that high release of citrate from cluster roots might be the preferred mechanisms to meet the P demand of nodulated plants under N deficiency. In addition, the high abundance of Rhizobiales and Rhodospirillales in the rhizosphere of cluster roots showed that the importance of N-fixing microorganisms under N deficiency. The contribution of rhizosphere microorganisms due to similar activities of N-cycling enzymes under the two different N treatments is less important for N nutrition of plants. Further understanding of the regulation of cluster roots under N-deficiency will provide new information on the interactions between P and N nutrition.     

Adsorption and desorption dynamics of citric acid anions in soil

The functional role of organic acid anions in soil has been intensively investigated, with special focus on (i) microbial respiration and soil carbon dynamics, (ii) nutrient solubilization or (iii) metal detoxification and reduction of plant metal uptake. Little is known about the interaction dynamics of organic acid anions with the soil matrix and the potential impact of adsorption and desorption processes on the functional significance of these effects. The aim of this study was to characterize experimentally the adsorption and desorption dynamics of organic acid anions in five agricultural soils differing in iron and aluminium oxide contents and using citrate as a model carboxylate. Results showed that both adsorption and desorption processes were fast in all soils, reaching a steady state within approximately 1 hour. However, for a given total soil citrate concentration (c_t) the steady state was critically dependent on the starting conditions of the experiment, whether most of the citrate was initially present in solution (c_l) or held on the solid phase (c_s). Specifically, desorption‐led processes resulted in significantly smaller steady‐state solution concentrations than adsorption‐led processes, indicating that hysteresis occurred. As it is not possible to distinguish between different adsorption and desorption pools in soil experimentally, a new dynamic hysteresis model that relies only on measured soil solution concentrations was developed. The model satisfactorily explained experimental data and was able to predict dynamic adsorption and desorption behaviour. To demonstrate its use, we applied the model to two relevant situations involving exudation and microbial degradation. The study highlighted the complex nature of citrate adsorption and desorption dynamics in soil. We conclude that existing models need to incorporate both temporal and hysteresis components to describe realistically the role and fate of organic acids in soil processes.