Modelling root-induced solubilization of nutrients

Theory for coupled diffusion processes in soil is briefly described and three examples of its application to understand root-induced solubilization of nutrients given. The examples are: (1) solubilization of P through root-induced pH changes in the rhizosphere of rice plants growing in flooded soil; (2) solubilization of P through excretion of organic chelating agents from rice roots growing in aerobic soil; and (3) the effects of root geometry on P solubilization, particularly cylindrical versus planar geometry and the effect of excretion of a solubilizing agent being localized along the root axis. The theory is tested by comparing measured concentration profiles of P near roots with the predictions of the theory made using independently measured parameter values. In the examples given, the agreement between the observed and predicted concentration profiles is very good, indicating that the theory is sound and the processes involved well understood.     

Citric acid excretion and precipitation of calcium citrate in the rhizosphere of white lupin (Lupinus albus L.)

Abstract. White lupin ( Lupinus albus L.) was grown for 13 weeks in a phosphorus (P) deficient calcareous soil (20% CaCO₃, pH(H₂O)7.5) which had been sterilized prior to planting and fertilized with nitrate as source of nitrogen. In response to P deficiency, proteoid roots developed which accounted for about 50% of the root dry weight. In the rhizosphere soil of the proteoid root zones, the pH dropped to 4.8 and abundant white precipitates became visible. X-ray spectroscopy and chemical analysis showed that these precipitates consisted of calcium citrate. The amount of citrate released as root exudate by 13-week-old plants was about 1 g plant⁻¹, representing about 23% of the total plant dry weight at harvest. In the rhizosphere soil of the proteoid root zones the concentrations of available P decreased and of available Fe, Mn and Zn increased. The strong acidification of the rhizosphere and the cation/anion uptake ratio of the plants strongly suggests that proteoid roots of white lupin excrete citric acid, rather than citrate, into the rhizosphere leading to intensive chemical extraction of a limited soil volume. In a calcareous soil, citric acid excretion leads to dissolution of CaCO₃ and precipitation of calcium citrate in the zone of proteoid roots.     

The potential for nitrification and nitrate uptake in the rhizosphere of wetland plants: a modelling study

BACKGROUND AND AIMS: It has recently found that lowland rice grown hydroponically is exceptionally efficient in absorbing NO3-, raising the possibility that rice and other wetland plants growing in flooded soil may absorb significant amounts of NO3- formed by nitrification of NH4+ in the rhizosphere. This is important because (a) this NO3- is otherwise lost through denitrification in the soil bulk; and (b) plant growth and yield are generally improved when plants absorb their nitrogen as a mixture of NO3- and NH4+ compared with growth on either N source on its own. A mathematical model is developed here with which to assess the extent of NO3- absorption from the rhizosphere by wetland plants growing in flooded soil, considering the important plant and soil processes operating. METHODS: The model considers rates of O2 transport away from an individual root and simultaneous O2 consumption in microbial and non-microbial processes; transport of NH4+ towards the root and its consumption in nitrification and uptake at the root surface; and transport of NO3- formed from NH4+ towards the root and its consumption in denitrification and uptake by the root. The sensitivity of the model's predictions to its input parameters is tested over the range of conditions in which wetland plants grow. KEY RESULTS: The model calculations show that substantial quantities of NO3- can be produced in the rhizosphere of wetland plants through nitrification and taken up by the roots under field conditions. The rates of NO3- uptake can be comparable with those of NH4+. The model also shows that rates of denitrification and subsequent loss of N from the soil remain small even where NO3- production and uptake are considerable. CONCLUSIONS: Nitrate uptake by wetland plants may be far more important than thought hitherto. This has implications for managing wetland soils and water, as discussed in this paper.     

The relationship between silicon availability,and growth and silicon concentration of the salt marsh halophyte Spartina anglica

Incorporation of cover crops into cropping systems may contribute to a more efficient utilization of soil and fertilizer P by less P-efficient crops through exudation of P-mobilizing compounds by the roots of P-efficient plant species. The main objective of the present work was to test this hypothesis. First a method has been developed which allows the quantification of organic anion exudation from individual cluster roots formed by P-deficient white lupin (Lupinus albus L.). Lupin plants were grown in nutrient solution at 1 μM P and in a low P loess in small rhizotrons. Organic anions exuded from intact plants grown in nutrient solution were collected from individual cluster roots and root tips sealed in small compartments by an anion-exchange resin placed in nylon bags (resin-bags). Succinate was the dominant organic anion exuded followed by citrate and malate. The mean of citrate exudation-rate was 0.06 pmol mm⁻¹ s⁻¹ with exudation highly dependent on the citrate concentration and on the age of the cluster roots. Exudates from cluster roots and root tips grown at the soil surface (rhizotron-grown plants) were collected using overlayered resin–agar (resin mixed with agar). Citrate exudation from cluster roots was 10 times higher than that from root tips. Fractionation of P in the cluster root rhizosphere-soil indicates that white lupin can mobilize P not only from the available and acid-soluble P, but also from the stable residual soil P fractions. In pot experiments with an acid luvisol derived from loess low in available P, growth of wheat was significantly improved when mixed-cropped with white lupin due to improved P uptake. Both in mixed culture and in rotation wheat could benefit from the P mobilization capacity of white lupin, supporting the hypothesis above. Nine tropical leguminous cover crops and maize were grown in a pot experiment using a luvisol from Northern Nigeria low in available P. All plant species derived most of their P from the resin and bicarbonate-extractable inorganic P. Organic P (P_o) accumulated particularly in the rhizosphere of all plant species. There was a significant negative correlation between the species-specific rhizosphere acid phosphatase activity and P_o accumulation. Growth and P uptake of maize grown in rotation after legumes were enhanced indicating that improved P nutrition was a contributing factor. This revised version was published online in June 2006 with corrections to the Cover Date.     

Phosphorus efficiency and the forms of soil phosphorus utilized by upland rice cultivars

Experimental measurements of phosphorus (P) uptake and the forms of soil P depleted from an Ultisol by 6 upland rice cultivars are reported. In both P-fertilized and-unfertilized soil, the majority of P taken up was solubilized from a 0.1 M NaOH-soluble pool by root-induced changes. The soil pH within 4 mm of the roots was lowered by up to 0.5 units (from 4.6), but this by itself could not account for the P solubilized, and nor could increased phosphatase activity near the roots. The possible role of root-released low molecular weight organic acid anions in P solubilization is discussed. No significant differences in the extent of solubilization by a given root mass could be detected between cultivars. In P-unfertilized soil, but not in P-fertilized soil, there were significant differences between cultivars in internal P efficiency as measured by shoot dry weight per unit total plant P. In unfertilized soil, root growth and P uptake were strongly correlated with the P content of the seeds from which the plants were grown.     

Enhanced zinc uptake by rice through phytosiderophore secretion: a modelling study

Rice (Oryza sativa L.) secretes far smaller amounts of metal-complexing phytosiderophores (PS) than other grasses. But there is increasing evidence that it relies on PS secretion for its zinc (Zn) uptake. After nitrogen, Zn deficiency is the most common nutrient disorder in rice, affecting up to 50% of lowland rice soils globally. We developed a mathematical model of PS secretion from roots and resulting solubilization and uptake of Zn, allowing for root growth, diurnal variation in secretion, decomposition of the PS in the soil, and the transport and interaction of the PS and Zn in the soil. A sensitivity analysis showed that with realistic parameter values for rice in submerged soil, the typically observed rates of PS secretion from rice are sufficient and necessary to explain observed rates of Zn uptake. There is little effect of diurnal variation in secretion on cumulative Zn uptake, irrespective of other model parameter values, indicating that the observed diurnal variation is not causally related to Zn uptake efficiency. Rooting density has a large effect on uptake per unit PS secretion as a result of overlap of the zones of influence of neighbouring roots. The effects of other complications in the rice rhizosphere are discussed.     

Spatial and temporal variation in organic acid anion exudation and nutrient anion uptake in the rhizosphere of Lupinus albus L.

We investigated in situ the temporal patterns and spatial extent of organic acid anion exudation into the rhizosphere solution of Lupinus albus, and its relation with the nutrient anions phosphate, nitrate and sulfate by means of a rhizobox micro suction cup method under P sufficient conditions. We compared the soil solution in the rhizosphere of cluster roots with that in the vicinity of normal roots, nodules and bulk soil. Compared to the other rhizosphere and soil compartments, concentrations of organic acid anions were higher in the vicinity of cluster roots during the exudative burst (citrate, oxalate) and nodules (acetate, malate), while concentrations of inorganic nutrient anions were highest in the bulk soil. Both active cluster roots and nodules were most efficient in taking up nitrate and phosphate. The intensity of citrate exudation by cluster roots was highly variable. The overall temporal patterns during the lifetime of cluster roots were overlaid by a diurnal pattern, i.e. in most cases, the exudation burst consisted of one or more peaks occurring in the afternoon. Multiple exudation peaks occurred daily or were separated by 1 or 2 days. Although citrate concentrations decreased with distance from the cluster root apex, they were still significantly higher at a distance of 6 to 10 mm than in the bulk soil. Phosphate concentrations were extremely variable in the proximity of cluster roots. While our results indicate that under P sufficient conditions cluster roots take up phosphate during their entire life time, the influence of citrate exudation on phosphate mobilization from soil could not be assessed conclusively because of the complex interactions between P uptake, organic acid anion exudation and P mobilization. However, we observed indications of P mobilization concurrent with the highest measured citrate concentrations. In conclusion, this study provides semiquantitative in situ data on the reactivity of different root segments of L. albus L. in terms of root exudation and nutrient uptake under nutrient sufficient conditions, in particular on the temporal variability during the lifetime of cluster roots.     

The supply of nutrient ions by diffusion to plant roots in soil

Summary A single-root technique is used to measure the rate of supply of potassium by diffusion to 1-cm portions of cylindrical roots of onion and leek plants growing in soils containing different levels of exchangeable potassium. The relation between uptake and characteristics of the plant and soil is interpreted on the basis of a diffusion supply model. Uptake is accounted for in terms of the geometry of the absorbing root surface, the physiologically controlled absorbing power of the root, and the diffusion through the soil. The different uptakes of potassium by roots of comparable absorbing power from different soils can be predicted with some success from calculations using the root dimensions and either diffusion coefficients of potassium in soil, derived from flux to a cation exchange resin paper, or the form of the potassium scorption isotherm relating the concentration of labile ions to those in the soil solution. It is calculated that diffusion through the soil has reduced potassium uptake by the roots to between 87 and 39 per cent of that expected for roots of the same absorbing power in a stirred culture solution at the same initial soil solution concentration.     

大豆和水稻对铝胁迫响应的生理机制

通过水培方式,研究了铝处理对双子叶植物大豆和单子叶植物水稻根系生长、养分吸收、根系内含物及根分泌物的影响.结果表明,低铝 (10 μmol·L^-1)浸种刺激大豆种子萌发和根系的生长,对水稻无明显促进作用.铝处理增加了两种作物根系对P的吸收,降低K、Ca、Mg的吸收.水稻比大豆根系积累较少的Al和更多的P.铝胁迫条件下,大豆和水稻根系内源可溶性蛋白含量升高、可溶性酚含量下降、可溶性糖含量先上升后下降,且大豆根系内源柠檬酸含量下降明显.与大豆相比,水稻积累较低的柠檬酸和较高的可溶性蛋白、可溶性酚,但两者可溶性糖没有差异.铝处理增加大豆根系柠檬酸、可溶性蛋白、可溶性酚、可溶性糖的分泌量,且大豆分泌量显著高于水稻.在铝处理条件下,大豆根系具有较高的阳离子交换量(CEC),而水稻的CEC较低.这说明大豆和水稻对铝胁迫具有不同的生理反应,水稻的高耐铝性可能与其较高的磷吸收和较低的CEC有关,而与其根系分泌物的关系不大.     

Variation in phosphorus uptake efficiency by genotypes of cowpea (Vigna unguiculata) due to differences in root and root hair length and induced rhizosphere processes

The lengths of roots and root hairs and the extent of root-induced processes affect phosphorus (P) uptake efficiency by plants. To assess the influence of variation in the lengths of roots and root hairs and rhizosphere processes on the efficiency of soil phosphorus (P) uptake, a pot experiment with a low-P soil and eight selected genotypes of cowpea (Vigna unguiculata (L) WALP) was conducted. Root length, root diameter and root hair length were measured to estimate the soil volume exploited by roots and root hairs. The total soil P was considered as a pool of Olsen-P, extractable with 0.5 M NaHCO₃ at pH 8.5, and a pool of non-Olsen-P. Model calculations were made to estimate P uptake originated from Olsen-P in the root hair zone and the Olsen-P moving by diffusion into the root hair cylinder and non-Olsen-P uptake. The mean uptake rate of P and the mean rate of non-Olsen-P depletion were also estimated. The genotypes differed significantly in lengths of roots and root hairs, and in P uptake, P uptake rates and growth. From 6 to 85% of total P uptake in the soil volume exploited by roots and root hairs was absorbed from the pool of non-Olsen-P. This indicates a considerable activity of root-induced rhizosphere processes. Hence the large differences show that traits for more P uptake-efficient plants exist in the tested cowpea genotypes. This opens the possibility to breed for more P uptake-efficient varieties as a way to bring more sparingly soluble soil P into cycling in crop production and obtain capitalisation of soil P reserves.     

The chemistry of the lowland rice rhizosphere

Models and experimental studies of the rhizosphere of rice plants growing in anaerobic soil show that two major processes lead to considerable acidification (1–2 pH units) of the rhizosphere over a wide range of root and soil conditions. One is generation of H⁺ in the oxidation of ferrous iron by O₂ released from the roots. The other is release of H⁺ from roots to balance excess intake of cations over anions, N being taken up chiefly as NH₄ ⁺. CO₂ exchange between the roots and soil has a much smaller effect. The zone of root-influence extends a few mm from the root surface. There are substantial differences along the root length and with time. The acidification and oxidation cause increased sorption of NH₄ ⁺ ions on soil solids, thereby impeding the movement of N to absorbing root surfaces. But they also cause solubilization and enhanced uptake of soil phosphate.     

Impact of phosphorus mineral source (Al–P or Fe–P) and pH on cluster-root formation and carboxylate exudation in Lupinus albus L.

Lupinus albus L. were grown in rhizoboxes containing a soil amended with sparingly available Fe–P or Al–P (100 μg P g⁻¹ soil/resin mixture). Root halves of individual plants were supplied with nutrient solution (minus P) buffered at either pH 5.5 or 7.5, to assess whether the source of mineral-bound P and/or pH influence cluster-root growth and carboxylate exudation. The P-amended soil was mixed 3:1 (w/w) with anion-exchange resins to allow rapid fixation of carboxylates. Treatments lasted 10 weeks. Forty percent and 30% of the root mass developed as cluster roots in plants grown on Fe–P and Al–P respectively, but cluster-root growth was the same on root-halves grown at pH 5.5 or 7.5. Mineral-bound P source (Al– or Fe–P) had no influence on the types of carboxylates measured in soil associated with cluster roots—citrate (and trace amounts of malate and fumarate) was the only major carboxylate detected. The [citrate] in the rhizosphere of cluster roots decreased with increased shoot P status (suggesting a systemic effect) and also, only for plants grown on Al–P, with decreased pH in the root environment (suggesting a local effect). In a separate experiment using anion exchange resins pre-loaded with malate or citrate, we measured malate (50%) and citrate (79%) recovery after 30 days in soil. We therefore, also conclude that measurements of [citrate] and [malate] at the root surface may be underestimated and would be greater than the 40- and 1.6-μmol g⁻¹ root DM, respectively estimated by us and others because of decomposition of carboxylates around roots prior to sampling.     

The dependence of methane transport in rice plants on the root zone temperature

Large diurnal and seasonal variations in methane flux from rice paddies have been found in many studies. Although these variations are considered to result from changes in methane formation rates in the soil and the transport capacity (e.g. biomass, physiological activities, and so on) of rice plants, the real reasons for such variations are as yet unclear. This study was conducted to clarify the effects of temperature on the rate of methane transport from the root zone to the atmosphere using hydroponically grown rice plants. Methane emission rates from the top of the rice plants whose roots were soaked in a solution with a high methane concentration were measured using a flow-through chamber method with the top or root of the rice plants being kept at various temperatures. The methane emission rates and methane concentrations in solution were analyzed using a diffusion model which assumes that the methane emission from a rice paddy is driven by molecular diffusion through rice plants by a concentration gradient. In the experiment where the temperature around the root was changed, the conductance for methane diffusion was typically 2.0-2.2 times larger when the solution temperature was changed from 15 to 30 °C. When the air temperature surrounding the top of the rice plant was changed, the change in conductance was much less. In addition, from measurements of methane flux and methane concentration in soil water in a lysimeter rice paddy during the 2 growing seasons of rice, it was found that the conductance for methane transport was correlated with the soil temperature at 5 cm depth. These results suggest that the temperature around the root greatly affects the methane transport process in rice plants, and that the process of passing through the root is important in determining the rate of methane transport through rice plants.     

Simulation of nutrient uptake by a growing root system considering increasing root density and inter-root competition

A simulation model is presented which describes uptake of a growth limiting nutrient from soil by a growing root system. The root surface is supposed to behave like a zero-sink. Uptake of the nutrient is therefore determined by the rate of nutrient supply to the root surface by mass flow and diffusion. Inter-root competition and time dependent root density are accounted for by assigning to each root a finite cylindrical soil volume that delivers nutrients. The radius of these cylinders declines with increasing root density. Experiments with rape plants grown on quartz sand were used to evaluate the model. Simulated nitrogen uptake agreed well with observed uptake under nitrogen limiting conditions. In case no nitrogen limitation occurred nitrogen uptake was overestimated by the model, probably because the roots did not behave like a zero-sink any more.     

The effects of water regime on phosphorus responses of rainfed lowland rice cultivars

Background and Aims

Soil phosphorus (P) solubility declines sharply when a flooded soil drains, and an important component of rice (Oryza sativa) adaptation to rainfed lowland environments is the ability to absorb and utilize P under such conditions. The aim of this study was to test the hypothesis that rice cultivars differ in their P responses between water regimes because P uptake mechanisms differ.

Methods

Six lowland rice cultivars (three considered tolerant of low P soils, three sensitive) were grown in a factorial experiment with three water regimes (flooded, moist and flooded-then-moist) and four soil P levels, and growth and P uptake were measured. Small volumes of soil were used to maximize inter-root competition and uptake per unit root surface. The results were compared with the predictions of a model allowing for the effects of water regime on P solubility and diffusion.

Key Results

The plants were P stressed but not water stressed in all the water regimes at all P levels except the higher P additions in the flooded soil. The cultivar rankings scarcely differed between the water regimes and P additions. In all the treatments, the soil P concentrations required to explain the measured uptake were several times the concentration of freely available P in the soil.

Conclusions

The cultivar rankings were driven more by differences in growth habit than specific P uptake mechanisms, so the hypothesis cannot be corroborated with these data. Evidently all the plants could tap sparingly soluble forms of P by releasing a solubilizing agent or producing a greater root length than measured, or both. However, any cultivar differences in this were not apparent in greater net P uptake, possibly because the restricted rooting volume meant that additional P uptake could not be converted into new root growth to explore new soil volumes.Key words: Oryza sativa, rainfed lowland, phosphorus efficiency, root morphology, solubilization, rice cultivar     

The influence of rhizosphere in rice crop on resin-extractable phosphate in flooded soils at various levels of phosphate applications

Summary Poor or lack of response of lowland rice to P fertilization is a well-known fact. Several studies were conducted in this direction however, our understanding regarding the underlying mechanism has been far from clear. A remarkable influence of rice plants on P transformation in submerged soil is identified in this experiment which may shed light on this problem. Accordingly, in presence of rice plants P was mobilized during the initial growing period followed by immobilization. The increased microbial activity in presence of physiologically active roots was responsible for P mobilization, while capacity of rice plants to reoxidize the rhizosphere, by secretion of oxygen from roots, during later growing period was responsible for P immobilization.The extent of P mobilization decreased while that of immobilization increased with increasing P levels in different soils. At a given P level the ratio of P mobilization to immobilization was higher in a soil where crop growth was better and P uptake was higher as compared to another soil where crop growth was poor with lower P uptake.Thus, lowland rice plants appear to possess an unique physiological mechanism, to regulate the contrasting changes in P availability in the rhizosphere depending on P requirement by the plants or P availability in soil, which in turn is responsible for the poor or lack of response to P fertilization.The experiment was conducted in a growth chamber. Two soils with widely varying properties were used.     

Root growth,aerenchyma development,and oxygen transport in rice genotypes subjected to drought and waterlogging

Soils under field conditions may experience fluctuating soil water regimes ranging from drought to waterlogging. The inability of roots to acclimate to such changes in soil water regimes may result in reduced growth and function thereby, dry matter production. This study compared the root and shoot growth, root aerenchyma development, and associated root oxygen transport of aerobic and irrigated lowland rice genotypes grown under well-watered (control), waterlogged, and droughted soil conditions for 30 days. The aerobic genotypes were as tolerant as the irrigated lowland genotypes under waterlogging because of their comparable abilities to enhance aerenchyma that effectively facilitated O₂ diffusion to the roots for maintaining root growth and dry matter production. Under drought, aerobic genotypes were more tolerant than the irrigated lowland genotypes due to their higher ability to maintain nodal root production, elongation, and branching, thus, less reduction in dry matter production. Aerenchyma was also formed in droughted roots regardless of genotypes, but was resistant to internal O₂ transport under O₂ deficiency. The ability of roots to resist temporal variations in drought and waterlogging stresses might have strong implications for the adaptation of rice growing in environments with fluctuating soil water regimes.     

Citrate-permeable channels in the plasma membrane of cluster roots from white lupin

White lupin (Lupinus albus) is well adapted to phosphorus deficiency by developing cluster roots that release large amounts of citrate into the rhizosphere to mobilize the sparingly soluble phosphorus. To determine the mechanism underlying citrate release from cluster roots, we isolated protoplasts from different types of roots of white lupin plants grown in phosphorus-replete (+P) and phosphorus-deficient (-P) conditions and used the patch-clamp technique to measure the whole-cell currents flowing across plasma membrane of these protoplasts. Two main types of anion conductance were observed in protoplasts prepared from cluster root tissue: (1) an inwardly rectifying anion conductance (IRAC) activated by membrane hyperpolarization, and (2) an outwardly rectifying anion conductance (ORAC) that became more activated with membrane depolarization. Although ORAC was an outward rectifier, it did allow substantial inward current (anion efflux) to occur. Both conductances showed citrate permeability, with IRAC being more selective for citrate3- than Cl- (PCit/PCl = 26.3), while ORAC was selective for Cl- over citrate (PCl/PCit = 3.7). Both IRAC and ORAC were sensitive to the anion channel blocker anthracene-9-carboxylic acid. These currents were also detected in protoplasts derived from noncluster roots of -P plants, as well as from normal (noncluster) roots of plants grown with 25 microm phosphorus (+P). No differences were observed in the magnitude or frequency of IRAC and ORAC currents between the cluster roots and noncluster roots of -P plants. However, the IRAC current from +P plants occurred less frequently than in the -P plants. IRAC was unaffected by external phosphate, but ORAC had reduced inward current (anion efflux) when phosphate was present in the external medium. Our data suggest that IRAC is the main pathway for citrate efflux from white lupin roots, but ORAC may also contribute to citrate efflux.     

Root Carbon Dioxide Fixation by Phosphorus-Deficient Lupinus albus (Contribution to Organic Acid Exudation by Proteoid Roots)

When white lupin (Lupinus albus L.) is subjected to P deficiency lateral root development is altered and densely clustered, tertiary lateral roots (proteoid roots) are initiated. These proteoid roots exude large amounts of citrate, which increases P solubilization. In the current study plants were grown with either 1 mM P (+P-treated) or without P (-P-treated). Shoots or roots of intact plants from both P treatments were labeled independently with 14CO2 to compare the relative contribution of C fixed in each with the C exuded from roots as citrate and other organic acids. About 25-fold more acid-stable 14C, primarily in citrate and malate, was recovered in exudates from the roots of -P-treated plants compared with +P-treated plants. The rate of in vivo C fixation in roots was about 4-fold higher in -P-treated plants than in +P-treated plants. Evidence from labeling intact shoots or roots indicates that synthesis of citrate exuded by -P-treated roots is directly related to nonphotosynthetic C fixation in roots. C fixed in roots of -P-treated plants contributed about 25 and 34% of the C exuded as citrate and malate, respectively. Nonphotosynthetic C fixation in white lupin roots is an integral component in the exudation of large amounts of citrate and malate, thus increasing the P available to the plant.