Effects of Zn-complexes on zinc uptake by wheat (Triticum aestivum) roots: a comprehensive consideration of physical, chemical and biological processes on biouptake

Commonly used equilibrium models for metal biouptake, such as the Free Ion Activity Model (FIAM) and the Biotic Ligand Model (BLM), are limited to the cases in which mass diffusive transport is not the flux-determining step. In analyses of metal biouptake from a complexing medium, all the physical (diffusion), chemical (dissociation kinetics of metal complexes), and biological (transport and internalization) processes have to be taken into account. A short-term zinc uptake by wheat (Triticum aestivum) roots from culture solutions in the absence or presence of synthetic ligands (NTA, nitrilotriacetic acid, and EDTA, ethylenediaminetetraacetate) was studied. At the same free Zn²⁺ concentration $\left( {\left\{ {{\text{Zn}}^{{\text{2 + }}} } \right\} = 1.5 \times 10^{ - 8} {\text{M}}} \right)$ , the uptake of Zn was significantly enhanced in the presence of ligands and was larger when Zn complexes have a quicker dissociation rate. The diffusional fluxes in the same culture solution were determined with the differential pulse anodic stripping voltammetry (DPASV) method, and the diffusive gels in thin film (DGT) technique. The contribution from Zn complexes to root Zn uptake was in better agreement with the degree of Zn complex labilities measured with DPASV than with DGT. The diffusion of free Zn²⁺ ion to the root surface is a rate-controlling step for Zinc biouptake when the free Zn²⁺ concentration is low. Based on the comprehensive consideration of the diffusion and dissociation processes of Zn²⁺ ion and Zn complexes and the existence of high- and low-affinity uptake systems in the root surface, a two-pathway Zn uptake model was developed to predict the resulting Zn uptake fluxes into roots in the overall range of exposure.     

Modelling zinc uptake by rice crop using a Barber-Cushman approach

The Barber-Cushman mechanistic nutrient uptake model which has been utilized extensively to describe and predict nutrient uptake by crop plants at different stages of crop growth was evaluated for its ability to predict the Zn uptake by rice seedlings. 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 soil volume that delivers nutrients. The radii of these cylinders decline with increasing density. Since mass flow and diffusion each supply zinc to the root, the process can be described mathematically using the model of Barber-Cushman (1984). The 11 parameters of the model for the uptake by rice cultivars were measured by established experimental techniques. Zinc uptake at different growth stages predicted by the model was compared to measured zinc uptake by rice cultivars grown on sandy loam soil in a green house. Predicted zinc uptake was significantly correlated with observed uptake r ²=0.99^**. Sensitivity analysis was also used to investigate the impact of changes in soil nutrient supply, root morphological and root uptake kinetic parameters on simulated nutrient uptake. Overall results of sensitivity analysis indicate that the half distance between root axes, rate of root growth and water flux affect the uptake of zinc particularly at their higher values rather than at lower values and DaZn is the most sensitive parameter for zinc uptake at its lower values.     

Uptake of zinc by rye,bread wheat and durum wheat cultivars differing in zinc efficiency

Effect of zinc (Zn) nutritional status on uptake of inorganic ⁶⁵Zn was studied in rye (Secale cereale, cv. Aslim), three bread wheat (Triticum aestivum, cvs. Dagdas, Bezostaja, BDME-10) and durum wheat (Triticum durum, cv. Kunduru-1149) cultivars grown for 13 days in nutrient solution under controlled environmental conditions. The cultivars were selected based on their response to Zn deficiency and to Zn fertilization in calcareous soils under field conditions. When grown in Zn-deficient calcareous soil in the field, the rye cultivar had the highest, and the durum wheat the lowest Zn efficiency. Among the bread wheats, BDME-10 showed higher susceptibility to Zn deficiency and Bezostaja and Dagdas were less affected by Zn deficiency. Similarly to field conditions, in nutrient solution visual Zn deficiency symptoms (i.e. necrotic lesions on leaf blade) appeared to be more severe in Kunduru-1149 and BDME-10 and less severe in rye cultivar Aslim. Under Zn deficiency, shoot concentrations of Zn were similar between all cultivars. Cultivars with adequate Zn supply did not differ in uptake and root-to-shoot translocation rate of ⁶⁵Zn, but under Zn deficiency there were distinct differences; rye showed the highest rate of Zn uptake and the durum wheat the lowest. In the case of bread wheat cultivars, ⁶⁵Zn uptake rate was about the same and not related to their differential Zn efficiency. Under Zn deficiency, rye had the highest rate of root-to-shoot translocation of ⁶⁵Zn, while all bread and durum wheat cultivars were similar in their capacity to translocate ⁶⁵Zn from roots to shoots. When Zn²⁺ activity in uptake solution ranged between 117 p M and 34550 pM, Zn-efficient and Zn-inefficient bread wheat genotypes were again similar in uptake and root-to-shoot translocation rate of ⁶⁵Zn. The results indicate that high Zn efficiency of rye can be attributed to its greater Zn uptake capacity from soils. The inability of the durum wheat cultivar Kunduru-1149 to have a high Zn uptake capacity seems to be an important reason for its Zn inefficiency. Differential Zn efficiency between the bread wheat cultivars used in this study is not related to their capacity to take up inorganic Zn. This revised version was published online in June 2006 with corrections to the Cover Date.     

Root Morphology and Zn2+ Uptake Kinetics of the Zn Hyperaccumulator of Sedum alfredii Hance

Root morphology and Zn2 uptake kinetics of the hyperaccumulating ecotype (HE) and nonhyperaccumulating ecotype (NHE) of Sedum alfredii Hance were investigated using hydroponic methods and the radiotracer flux technique. The results indicate that root length, root surface area, and root volume of NHE decreased significantly with increasing Zn2 concentration in growth media, whereas the root growth of HE was not adversely affected, and was even promoted, by 500 μmol/L Zn2 . The concentrations of Zn2 in both ecotypes of S. alfredii were positively correlated with root length, root surface area and root volumes, but no such correlation was found for root diameter. The uptake kinetics for 65Zn2 in roots of both ecotypes of S. alfredii were characterized by a rapid linear phase during the first 6 h and a slower linear phase during the subsequent period of investigation. The concentration-dependent uptake kinetics of the two ecotypes of S. alfredii could be characterized by the Michaelis-Menten equation, with the Vmax for 65Zn2 influx being threefold greater in HE compared with NHE, indicating that enhanced absorption into the root was one of the mechanisms involved in Zn hyperaccumulation. A significantly larger Vmax value suggested that there was a higher density of Zn transporters per unit membrane area in HE roots.     

Kinetics of zinc release from ground tire rubber and rubber ash in a calcareous soil as alternatives to Zn fertilizers

Ground rubber contains 15?C20 g Zn kg^?1 but very low levels of Cd and could serve as an inexpensive byproduct Zn fertilizer. The aim of this investigation was to test Zn release in a soil treated with ground tire rubber and rubber ash compared with commercial Zn fertilizer and a laboratory grade zinc sulfate. A Zn-deficient soil was chosen from wheat fields in Isfahan province, central Iran, and the ground rubber, rubber ash and fertilizer-Zn and laboratory ZnSO₄ were added at 0.5 and 2 mg Zn kg^?1; 0.5 kg ha^?1 would usually correct Zn deficiency in such pot tests. The soil DTPA-extractable Zn was then measured with time and the results were described examining first order, Elovich, power function and parabolic diffusion kinetics models. In the pot experiment, corn (Zea mays L.) plants were exposed to three rates of Zn (0, 20, 40 mg Zn kg^?1) from two different sources (ZnSO₄ and ground rubber). Ground rubber was applied as 2?C3 mm and <1 mm diameter particles. Zinc treatments were mixed with the soils before planting. At harvest, concentrations of Zn, Pb, and Cd in roots and shoots of corn were measured. Results showed that ground rubber and rubber ash significantly increased the concentration of DTPA-Zn in the soil and this increase was higher than achieved with the commercial Zn fertilizer. At the lower Zn application rate, Zn release followed parabolic diffusion, while at the higher rate the kinetics of release followed power function and Elovich models. There was an increase in Zn concentration of corn shoot and roots by adding of Zn regardless the source of applied Zn. With increase in the rate of rubber used, the shoot Zn uptake increased. The Pb concentration of shoot and Cd concentrations of shoot and roots were low (less than 0.02 mg kg^?1) in all treatments. The results showed that the soil DTPA Zn decreases over time if the soil is amended with a soluble form of Zn whereas the reverse was observed if the Zn is added as ground rubber which only gradually transforms. Thus ground rubber and rubber ash offer strong value as Zn fertilizer for Zn deficient soils.     

Predicting zinc bioavailability to wheat improves by integrating pH dependent nonlinear root surface adsorption

Aim

Our aim was to improve the prediction of Zn bioavailability to wheat grown on low-Zn soils. The classical approach that directly relates Zn in a certain soil extract to Zn uptake has been shown to be inadequate in many cases. We tested a stepwise approach where the steps of the uptake process are characterized with, respectively, Zn solid-solution distribution, adsorption of Zn to root surface, Zn uptake into root and Zn translocation to shoot.

Methods

Two pot experiments were done with wheat grown on nine low-Zn soils varying widely in pH, clay and organic matter content. Soluble Zn concentrations in two soil extracts (DTPA and CaCl₂) were measured. Free Zn ion concentrations in CaCl₂ soil extracts were determined with the Donnan Membrane Technique. These Zn concentrations were then related to plant Zn uptake following both the direct and the stepwise approach.

Results

In the direct approach, Zn in the DTPA extract was a better predictor for shoot Zn uptake than Zn in the CaCl₂ extract. In the stepwise approach, the relationship between Zn in CaCl₂ extracts and the root surface adsorbed Zn was pH-dependent and nonlinear. Root surface adsorbed Zn was linearly related to root Zn uptake, and the latter was linearly related to the shoot Zn uptake. The stepwise approach improved the Zn uptake prediction compared to the direct approach and was also validated for different wheat cultivars.

Conclusions

The adsorption of Zn on the root surface is pH dependent and nonlinear with respect to the soil Zn concentration, and a useful proxy for bioavailable Zn over a wide range of soils.     

Characterization of Zinc Uptake, Binding, and Translocation in Intact Seedlings of Bread and Durum Wheat Cultivars

Durum wheat (Triticum turgidum L. var durum) cultivars exhibit lower Zn efficiency than comparable bread wheat (Triticum aestivum L.) cultivars. To understand the physiological mechanism(s) that confers Zn efficiency, this study used ⁶⁵Zn to investigate ionic Zn²⁺ root uptake, binding, and translocation to shoots in seedlings of bread and durum wheat cultivars. Time-dependent Zn²⁺ accumulation during 90 min was greater in roots of the bread wheat cultivar. Zn²⁺ cell wall binding was not different in the two cultivars. In each cultivar, concentration-dependent Zn²⁺ influx was characterized by a smooth, saturating curve, suggesting a carrier-mediated uptake system. At very low solution Zn²⁺ activities, Zn²⁺ uptake rates were higher in the bread wheat cultivar. As a result, the Michaelis constant for Zn²⁺ uptake was lower in the bread wheat cultivar (2.3 μm) than in the durum wheat cultivar (3.9 μm). Low temperature decreased the rate of Zn²⁺ influx, suggesting that metabolism plays a role in Zn²⁺ uptake. Ca inhibited Zn²⁺ uptake equally in both cultivars. Translocation of Zn to shoots was greater in the bread wheat cultivar, reflecting the higher root uptake rates. The study suggests that lower root Zn²⁺ uptake rates may contribute to reduced Zn efficiency in durum wheat varieties under Zn-limiting conditions.Soils that contain insufficient levels of the essential plant micronutrient Zn are common throughout the world. As a result, Zn deficiency is a widespread problem in crop plants, especially cereals (Graham et al., 1992). The importance of plant foods as sources of Zn, particularly in the marginal diets of developing countries, is well established (Welch, 1993). The development of crop plants that are efficient Zn accumulators is therefore a potentially important endeavor. In addition to its effects on nutrition, Zn deficiency in crops is relevant to other areas of human health. Another consequence of Zn-deficient soils is the tendency for plants grown in such soils to accumulate heavy metals. For example, in the Great Plains region of North America, where soil Zn levels are low and naturally occurring Cd is present, durum wheat (Triticum turgidum L. var durum) grains accumulate Cd to relatively high concentrations (Wolnik et al., 1983). The presence of Cd in food represents a potential human health hazard and, in response, international trade standards have been proposed to limit the levels of Cd in exported grain (Codex Alimentarius Commission, 1993). Thus, there is a need to understand the physiological processes that control acquisition of Zn from soil solution by roots and mobilization of Zn within plants.It has been demonstrated in recent years that crop plants vary in their ability to take up Zn, particularly when its availability to roots is limited. Zn efficiency, defined as the ability of a plant to grow and yield well in Zn-deficient soils, varies among wheat cultivars (Graham and Rengel, 1993). In field trials, durum wheat cultivars have been shown to be consistently less Zn efficient than bread wheat (Triticum aestivum L.) cultivars (Graham et al., 1992). Similarly, durum wheat varieties were reported to be less Zn efficient than bread wheat varieties when grown in chelate-buffered hydroponic nutrient culture (Rengel and Graham, 1995a).The physiological mechanism(s) that confers Zn efficiency has not been identified. Processes that could influence the ability of a plant to tolerate limited amounts of available Zn include higher root uptake, more efficient utilization of Zn, and enhanced Zn translocation within the plant. Cakmak et al. (1994) showed that a Zn-inefficient durum wheat cultivar exhibited Zn-deficiency symptoms earlier and more intensely than a Zn-efficient bread wheat cultivar even though the Zn tissue concentrations were similar in both lines, suggesting differential utilization of Zn in the two cultivars. Rates of Zn translocation to shoots were shown to vary among sorghum cultivars, although correlations with Zn efficiency were not established (Ramani and Kannan, 1985). Root uptake kinetics have been reported to vary between rice cultivars having different Zn requirements, with high-Zn-requiring cultivars exhibiting consistently higher root uptake rates (Bowen, 1986). In contrast, a correlation between Zn efficiency and rates of root Zn uptake in bread and durum wheat cultivars could not be demonstrated (Rengel and Graham, 1995b).In grasses Zn influx into the root symplasm has been hypothesized to occur as the free Zn²⁺ ion (Halvorson and Lindsay, 1977), as well as in the form of Zn complexes with nonprotein amino acids known as phytosiderophores (Tagaki et al., 1984) or phytometallophores (Welch, 1993). Concentration-dependent uptake of free Zn²⁺ ions has been shown to be saturable in several species, including maize (Mullins and Sommers, 1986), barley (Veltrup, 1978), and wheat (Chaudhry and Loneragan, 1972), suggesting that ionic uptake in grasses occurs via a carrier-mediated system. However, several of these studies have been criticized on the basis that excessively high (and physiologically unrealistic) Zn²⁺ concentrations were used (Kochian, 1993).This study was undertaken to examine unidirectional Zn²⁺ influx and translocation to shoots in Zn-efficient bread wheat lines and Zn-inefficient durum wheat lines. Experiments were performed in the absence of added phytometallophores and results are presumed to represent influx of ionic Zn²⁺. Zn activities in the nanomolar range were used to more closely mimic free Zn²⁺ levels occurring naturally in soil solution. The results presented here indicate that a Zn-efficient bread wheat cultivar maintained higher rates of Zn uptake than a Zn-inefficient durum wheat cultivar, particularly at low (and physiologically relevant) solution Zn²⁺ activities.     

Importance of seed Zn content for wheat growth on Zn-deficient soil

Seed nutrient reserves may be important for an early establishment of crops on low-fertility soils. This glasshouse pot study evaluated effects of seed Zn content on vegetative growth of two wheat (Triticum aestivum L.) genotypes differing in Zn efficiency. Low-Zn (around 250 ng Zn per seed) and high-Zn seed (around 700 ng Zn per seed on average) of Excalibur (Zn efficient) and Gatcher (Zn inefficient) wheats were sown in a Zn-deficient siliceous sand fertilised with 0, 0.05, 0.2, 0.8 or 3.2 mg Zn kg^-1 soil. After 3 weeks, plants derived from the high-Zn seed had better root and shoot growth; the cv. Excalibur accumulated more shoot dry matter than the cv. Gatcher. After 6 weeks, greater root and shoot growth of plants grown from the high-Zn seed compared to those from the low-Zn seed was obvious only at nil Zn fertilisation. A fertilisation rate of 0.2 mg Zn kg^-1 soil was required for achieving 90% of the maximum yield for plants grown from the high-Zn seed compared to 0.8 mg Zn kg^-1 soil for plants derived from the low Zn seed. The critical Zn level in youngest expanded leaves for 90% maximum yield was 16 mg Zn kg^-1 dry matter for both genotypes. Zn-efficient Excalibur had greater net Zn uptake rates compared to Zn-inefficient Gatcher after 3 weeks but they were not different at the 6-week harvest. Zinc-deficient plants had greater net uptake rates of Cu, Mn, B, P, and K but a reduced uptake rate of Fe. It is concluded that higher seed Zn content acted similar to a starter-fertiliser effect by improving vegetative growth and dissipating differences in Zn efficiency of wheat genotypes.     

Soil solution dynamics and plant uptake of cadmium and zinc by durum wheat following phosphate fertilization

A growth chamber study was conducted to evaluate the effect of application of phosphate fertilizer on soil solution dynamics of cadmium (Cd) and Cd accumulation in durum wheat (Triticum turgidum L. var. durum). Treatments consisted of three phosphate fertilizer sources containing 3.4, 75.2, and 232 mg Cd kg^?1 applied at three rates (20, 40 and 80 mg P kg^?1) plus a no fertilization control. An unplanted treatment at 40 mg P kg^?1 was included to separate the effects on soil solution Cd dynamics of the crop from that of the fertilizer. Soil solution samples were obtained using soil moisture samplers every 10 days after germination. The experimental results indicated that plant biomass significantly increased with P application rates and decreased with increased Cd concentration in the phosphate fertilizers. Total cadmium concentration in soil solution was not consistently affected by phosphate fertilization rate and fertilizer sources, and therefore Cd concentration in the fertilizer. Application of phosphate fertilizer, however, increased the concentration and accumulation of Cd and shoot Cd/Zn ratio, and decreased shoot Zn concentration in durum wheat. Phosphate sources had a marginally significant effect (P?=?0.05) on shoot Cd concentration and did not affect Cd accumulation in durum wheat. Concentration of Cd in soil solution was unrelated to Cd concentration in durum wheat. These results suggest that the immediate increase in Cd concentration and Cd accumulation in durum wheat with phosphate application is due more to competition between Zn and Cd for absorption into plants, enhanced root to shoot translocation and enhanced root development, than to a direct addition effect from Cd contained in phosphate fertilizer. In the short term, application of phosphate fertilizers can increase Cd concentration in the crops, regardless of the Cd concentration of the fertilizer. An optimal P fertilization, possibly in combination with Zn application, may offer an important strategy for decreasing Cd concentration and accumulation in crops.     

Root Morphology and Zn^2＋ Uptake Kinetics of the Zn Hyperaccumulator of Sedum alfredii Hance

Root morphology and Zn^2＋ uptake kinetics of the hyperaccumulating ecotype （HE） and nonhyperaccumulating ecotype （NHE） of Sedum alfredii Hance were investigated using hydroponic methods and the radiotracer flux technique. The results indicate that root length, root surface area, and root volume of NHE decreased significantly with increasing Zn^2＋ concentration in growth media, whereas the root growth of HE was not adversely affected, and was even promoted, by 500μmol/L Zn^2＋. The concentrations of Zn^2＋ in both ecotypes of S. alfredii were positively correlated with root length, root surface area and root volumes, but no such correlation was found for root diameter. The uptake kinetics for ^65Zn^2＋ in roots of both ecotypes of S. alfredii were characterized by a rapid linear phase during the first 6 h and a slower linear phase during the subsequent period of investigation. The concentration-dependent uptake kinetics of the two ecotypes of S. alfredii could be characterized by the Michaelis-Menten equation, with the Vmax for ^65Zn^2＋ influx being threefold greater in HE compared with NHE, indicating that enhanced absorption into the root was one of the mechanisms involved in Zn hyperaccumulation. A significantly larger Vmax value suggested that there was a higher density of Zn transporters per unit membrane area in HE roots.     

Applying a solute transfer model to phytoextraction: Zinc acquisition by Thlaspi caerulescens

Phytoextraction is the removal of metals from contaminated soils into harvested plant tissues. The rate of phytoextraction is governed by both soil and plant characteristics. Most effort has focused on identifying appropriate plants for phytoextraction, but the benefits from this effort will be marginal unless the metals are in phytoavailable forms in the rhizosphere. The concentration of a metal in the rhizosphere can be estimated using solute transfer models that incorporate: the metal concentration in the bulk soil solution, the buffer power of the soil, diffusion coefficient for the metal, water movement, root size and morphology, and the rate of entry of metal into the roots. Here a solute transfer model is developed to predict the concentration of Zn in the rhizosphere solution ([Zn]_ext) of Thlaspi caerulescens, a hyperaccumulator species that could be exploited for Zn phytoextraction. The model predicts that Zn accumulation by T. caerulescens is sub-optimal when the Zn concentration in the bulk soil solution is <27 M. Such a high [Zn]_ext is rare in contaminated agricultural soils, but is possible in the metalliferous substrates where T. caerulescens is endemic. Sensitivity analyses indicate that Zn diffusion is more important than transpiration-driven mass flow for Zn delivery to the root, implying that management of soil physical and hydrological properties will improve phytoextraction. Sensitivity analyses also imply that strategies to enhance the Zn absorption power of the root will not necessarily be successful for enhancing phytoextraction per se. Thus, research into enhancing Zn availability and mobility in soil will be as important as understanding and manipulating Zn uptake by plants. In general, such models can be used to identify constraints to efficient phytoextraction (whether plant or soil) and to determine whether commercial phytoextraction is feasible.     

The effects of cropping corn on the extractability of zinc added to a calcareous soil

The effects of cropping corn on the decrease in the extractability of Zn added to a calcareous soil were studied by a pot experiment and chemical extractions. The results show that the concentrations of Zn exchangeable with MgCl₂ (EXC-Zn) and extractable with DTPA (DTPA-Zn) in the soils with added Zn decreased with time. The processes associated with the decrease in extractability in DTPA of Zn added to soil can be described aptly by a diffusion equation which gives the proportion of added Zn in the non-DTPA fraction as a function of the square root of incubation time. This result suggests that the diffusion of Zn cations into microporous solids is a rate-limiting reaction. The relative diffusion rate coefficients (D/r²) were found to be 1.95×10^-10 and 3.34×10^-10 sec^-1 in the soils with added Zn of 20 and 60 mg kg^-1, respectively. Compared with uncropped soil, the concentrations of DTPA-Zn in the soils with added Zn were decreased by cropping. The decrease of DTPA-Zn in the soils in the presence of corn can be attributed to both its acquisition by corn and other processes associated with the growing of corn. The activity of plant roots would appear to enhance the process of decrease in the extractability in DTPA of Zn added to the soil. The source of Zn uptake by corn was affected by the loading or activity of Zn in soil. In the soil with low available Zn, the DTPA non-extractable Zn (non-DTPA-Zn) was mobilized and taken up by corn. In the soils with high available Zn, e.g. the recently added Zn, only EXC-Zn and a part of the DTPA-Zn were taken up by corn.     

Uptake and transport of foliar applied zinc (65Zn) in bread and durum wheat cultivars differing in zinc efficiency

Using two bread wheat (Triticum aestivum) and two durum wheat (Triticum durum) cultivars differing in zinc (Zn) efficiency, uptake and translocation of foliar-applied ⁶⁵Zn were studied to characterize the role of Zn nutritional status of plants on the extent of phloem mobility of Zn and to determine the relationship between phloem mobility of Zn and Zn efficiency of the used wheat cultivars. Irrespective of leaf age and Zn nutritional status of plants, all cultivars showed similar Zn uptake rates with application of ⁶⁵ZnSO₄ to leaf strips in a short-term experiment. Also with supply of ⁶⁵ZnSO₄ by immersing the tip (3 cm) of the oldest leaf of intact plants, no differences in Zn uptake were observed among and within both wheat species. Further, Zn nutritional status did not affect total uptake of foliar applied Zn. However, Zn-deficient plants translocated more ⁶⁵Zn from the treated leaf to the roots and remainder parts of shoots. In Zn-deficient plants about 40% of the total absorbed ⁶⁵Zn was translocated from the treated leaf to the roots and remainder parts of shoots within 8 days while in Zn-sufficient plants the proportion of the translocated ⁶⁵Zn of the total absorbed ⁶⁵Zn was about 25%. Although differences in Zn efficiency existed between the cultivars did not affect the translocation and distribution of ⁶⁵Zn between roots and shoots. Bread wheats compared to durum wheats, tended to accumulate more ⁶⁵Zn in shoots and less ⁶⁵Zn in roots, particularly under Zn-deficient conditions. The results indicate that differences in expression of Zn efficiency between and within durum and bread wheats are not related to translocation or distribution of foliar-applied ⁶⁵Zn within plants. Differential compartementation of Zn at the cellular levels is discussed as a possible factor determining genotypic variation in Zn efficiency within wheat.     

Effects of nutrient solution zinc activity on net uptake,translocation, and root export of cadmium and zinc by separated sections of intact durum wheat (Triticum turgidum L. var durum) seedling roots

Cd accumulation in durum wheat presents a potential health risk to consumers. In an effort to understand the physiological mechanisms involved with Cd accumulation, this study examined the effects of Zn on Cd root uptake and phloem translocation in a split– root system. Durum wheat seedlings were grown in chelate-buffered nutrient solution with intact root systems divided into two sections. Each root section grew in a separate 1 l pot, one of which contained 0.2 μM CdSO₄. In addition, each two-pot system contained ZnSO₄ in the following combinations (in μm) (for -cd root system: +cd root system): 1:1, 1:10, 10:1,10:10, 1:19, and 19:1. Harvested plant material was analyzed for Cd and Zn. In addition, rates of Cd and Zn net uptake, translocation to the shoot, and root export (translocation from one root segment to the other) between days 8 and 22 were calculated. Results show that Zn was not translocated from one root section to its connected root section. Uptake rates of Cd increased as solution Zn concentrations increased. Cd translocation from one root section to the other decreased significantly when the Zn concentration in either pot was greater than 1 μM. These results show the potential of Zn to inhibit movement of Cd via the phloem, and suggests that providing adequate Zn levels may limit phloem loading of Cd into wheat grain. Increasing the rhizosphere activity of Zn²⁺ in Cd-containing soils may therefore result in reduced Cd accumulation in grain even while net Cd uptake is slightly enhanced. This revised version was published online in June 2006 with corrections to the Cover Date.     

Phosphorus uptake by root systems: mathematical modelling in ecosys

The uptake of P by plant root systems is believed to be controlled by the concentration of soluble orthophosphate at the root surface. If a P transformation model in which this concentration is calculated were coupled to a root and mycorrhizal growth model in which this concentration is used to calculate P uptake, then it should be possible to simulate P uptake under different soil and climate conditions if soil properties relevant to the control of P concentration are known. To test this idea, models for the transformation and transport of inorganic and organic P were coupled to ones for root growth and nutrient uptake as part of the ecosys modelling program. Seasonal estimates of soluble P concentration, root growth and P uptake from the combined models were tested with data measured from barley under fertilized and unfertilized treatments in a long term P fertilizer experiment conducted on two different soils. In both soils the fertilizer treatment increased simulated and measured soluble P concentrations from 0.1-0.2 to 0.2-0.4 g m^-3, annual P uptake from 0.6-0.7 to 1.2-1.4 g m^-2, and annual DM accumulation from 400-500 to 700-800 g m^-2. Increases in soluble P concentrations caused by fertilizer P were reproduced in the model from changes in the balance between the desorption and dissolution of solid P on one hand, and the uptake of P by root and mycorrhizal systems on the other. Increases in P uptake caused by fertilizer P were reproduced in the model from higher solution P concentrations, root uptake kinetics, and from functional equilibria for C and P exchange simulated among mycorrhizal, root and shoot components of the plant. There was a tendency in the model to overestimate P uptake later in the growing season in the unfertilized treatment which could be corrected if parameters for root uptake kinetics were reduced after anthesis. Because the model is constructed independently of data for P uptake, and avoids the use of site-specific parameters, it may provide a means of estimating uptake under different managements and climates from soils of known properties.     

Nitric oxide flux from soil during the growing season of wheat by continuous measurements of the NO soil–atmosphere concentration gradient: A process study

The surface flux of nitric oxide from a wheat field was investigated from 23 March to 29 May 1997 in the Kerzersmoos, Switzerland. A plot fertilised with 19 kg N ha^-1 in cattle slurry and 40 kg N ha^-1 in mineral NH₄NO₃ fertiliser and a plot receiving no nitrogen containing fertiliser were compared. The flux was calculated based on hourly measurements of the NO soil–atmosphere concentration gradient using the one-dimensional soil diffusion model of Galbally and Johansson (1989). The soil bulk diffusion coefficient was determined from measurements of the ²²²Rn surface flux and the activity gradient between 10 cm depth and the surface. It ranged between 79% and 0.3% of the NO diffusion coefficient in air and was parameterised by air filled soil pore space. The indirectly determined NO flux agreed well with standard flux measurements using dynamic chambers. The largest NO emission was found following fertiliser application and irrigation. The emission occurred in pulses, which lasted for 4 days up to 3 weeks coinciding with elevated soil ammonium concentrations. Nitric oxide emission in 5 days following application of cattle slurry were 31 g NO-N ha^-1 and 5 g NO-N ha^-1 from the non-fertilised plot, respectively. Nitric oxide emission in 15 days following application of NH₄NO₃ was 95 g NO-N ha^-1 and 10 g NO-N ha^-1 from the non-fertilised plot, respectively. NO emission in 4 days following irrigation on 21 April were 36 g N ha^-1 from the fertilised and 39 g N ha^-1 from the non-fertilised plot. The daily NO emission before and after fertiliser and irrigation pulses was between 0.3 and 0.7 g NO-N ha^-1 d^-1. NO production and NO uptake of the soil was measured regularly. No systematic influence of management or climate on NO uptake was found. NO production was strongly stimulated by fertiliser input and soil moisture content. The simulation of NO production could be reproduced using a nitrification algorithm (Riedo et al., 1998) driven by soil temperature, moisture and ammonium concentration. A NO production rate constant of 1.1ċ10^-3 h^-1 at 15 °C was derived from a linear regression between nitrification and NO production. Introducing the parameterisation of NO production into the model of Galbally and Johansson (1989) the duration and the strength of the NO emission pulses could be reproduced and the total NO emission during the experiment was approximated within a factor of two. This revised version was published online in June 2006 with corrections to the Cover Date.     

Upscaling from Rhizosphere to Whole Root System: Modelling the Effects of Phospholipid Surfactants on Water and Nutrient Uptake

While the rhizosphere presents a different chemical, physical and biological environment to bulk soil, most experimental and modelling investigations of plant growth and productivity are based on bulk soil parameters. In this study, water and nutrient acquisition by wheat (Triticum aestivum L.) roots was investigated using rhizosphere- and root-system-scale modelling. The physical and chemical properties of rhizosphere soil could be influenced by phospholipid surfactants in the root mucilage. Two models were compared: a 2-dimensional (2D) Finite Element Method rhizosphere model, and a 3-dimensional (3D) root architecture model, ROOTMAP. ROOTMAP was parameterised to reproduce the results of the detailed 2D model, and was modified to include a rhizosphere soil volume. Lecithin (a phospholipid surfactant) could be exuded into the rhizosphere soil volume, decreasing soil water content and hydraulic conductivity at any given soil water potential, and decreasing phosphate adsorption to soil particles. The rhizosphere-scale modelling (5 × 5 mm² soil area, 10 mm root length, uptake over 12 h) predicted a reduction in water uptake (up to 16% at 30 kPa) and an increase in phosphate uptake (up to 4%) with lecithin exudation into the rhizosphere, but little effect on nitrate uptake, with only a small reduction in dry soil (1.6% at 200 kPa). The 3D root model reproduced the water (y = 1.013x, R² = 0.996), nitrate (y = 1x, R² = 1) and phosphate (y = 0.978x, R² = 0.998) uptake predictions of the rhizosphere model, providing confidence that a whole root system model could reproduce the dynamics simulated by a Finite Element Method rhizosphere model. The 3D root architecture model was then used to scale-up the rhizosphere dynamics, simulating the effect of lecithin exudation on water, nitrate and phosphate acquisition by a wheat root system, growing over 41 d. When applied to growing and responsive roots, lecithin exudation increased P acquisition by up to 13% in nutrient-rich, and 49% in relatively nutrient-poor soil. A comparison of wheat (Triticum aestivum L.) and lupin (Lupinus angustifolius L.) root architectures, suggested an interaction between the P acquisition benefit of rhizosphere lecithin and root architecture, with the more highly-branched wheat root structure acquiring relatively more P in the presence of lecithin than the sparsely-branched lupin root system.     

Quantitative relationship between zinc transport and plant uptake

Summary The importance of the Zn buffer power and its influence on the Zn concentration in soil solution was investigated in a simulatory experiment where the soil in question, previously treated with Zn and compacted to known bulk density, was eluted with 0.01M CaCl₂ under constant hydraulic head. The data so obtained were correlated with Zn, uptake by maize. The correlation coefficient for effluent Znvs total Zn uptake improved, remarkably when the corresponding Zn buffer power was also incorporated into the computations. It is concluded from this and the earlier investigation² that the Zn buffer power is the most important parameter governing Zn uptake by maize.     

The chelating effect of citric acid,oxalic acid,amino acids and Pseudomonas fluorescens bacteria on phytoremediation of Cu,Zn, and Cr from soil using Suaeda vera

Phytoextraction is a green technique for the removal of soil contaminants by plants uptake with the subsequent elimination of the generated biomass. The halophytic plant Suaeda vera Forssk. ex J.F.Gmel. is an native Mediterranean species able to tolerate and accumulate salts and heavy metals in their tissues. The objective of this study was to explore the potential use of S. vera for soil metal phytoextraction and to assess the impact of different chelating agents such as natural organic acids (oxalic acid [OA], citric acid [CA]), amino acids (AA) and Pseudomonas fluorescens bacteria (PFB) on the metal uptake and translocation. After 12 months, the highest accumulation of Cu was observed in the root/stem of PFB plots (17.62/8.19 mg/kg), in the root/stem of CA plots for Zn (31.16/23.52 mg/kg) and in the root of OA plots for Cr (10.53 mg/kg). The highest accumulation of metals occurred in the roots (27.33–50.76 mg/kg). Zn was the metal that accumulated at the highest rates in most cases. The phytoextraction percentages were higher for Cu and Zn (～2%) with respect to Cr (～1%). The percentages of metal removal from soil indicate the need to monitor soil properties, to recognize the influence of each treatment and to increase the concentration of bioavailable metals by the use of agricultural management practices aimed at promoting plant growth.