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.     

Studies on the adsorption and availability of copper in some arid soils

Summary Eight texturally different arid soils have been studied for the adsorption reactions of copper and thereby evaluated for their quantity, intensity and supply parameters for the copper. In these soils with the addition of increasing amounts of copper there was increase in the equilibrium concentration, adsorption, per cent saturation of adsorption capacity and supply parameter of copper. However, negative relationship of differential buffering capacity with quantity, intensity and supply parameters revealed that the sandy loam soils exhibited comparatively more resistance to change in the solution concentration of copper. Multiple regression analysis revealed that in all soils quantity, intensity and differential buffering capacity were the sole parameters accounting for the supply of the nutrient. Sandy loam soils having comparatively higher values for the adsorption maxima, bonding energy constant and differential buffering capacity of the soils required higher doses of applied copper to change in the solution concentration than in sandy soils.     

Iron,zinc and manganese nutrition of wetland rice (Oryza sativa L.) on a gypsum amended sodic soil

A field experiment was conducted to evaluate the effect of three levels of Fe and two levels of Zn, and their combinations, on the growth, yield and Fe, Zn, and Mn nutrition of rice on a zinc deficient sodic soil amended with gypsum. Iron and zinc were supplied as sulphates. Application of Zn significantly enhanced the yield of rice and available soil and plant Zn irrespective of Fe application. Maximum response of rice to Zn was obtained when Fe was applied at the highest rate. While Fe application brought about a significant improvement in available soil and plant Fe and Mn, it decreased significantly Zn content of the crop. After crop harvest, recovery of added Fe was 20% and Zn 12%. Results suggest that benefits of Fe application to rice in sodic soils can only be realised if it is applied along with Zn.     

The relative efficiency of zinc carriers on growth and zinc nutrition of corn

Summary A comparison of different zinc carriers showed that application of Zn-DTPA, Zn-EDTA, Zn-fulvate and ZnSO₄ significantly increased the dry matter yield and zinc uptake by corn over the control treatment where no zinc was applied. The chelates in particular enhanced to a greater extent the uptake of both native and applied sources than that observed with ZnSO₄ as the zinc carrier. Both the dry matter yield and zinc uptake by corn showed a positive and significant relationship with self-diffusion coefficient of zinc showing thereby that diffusion contributed mainly the supply of Zn from the ambient soil matrix to plant roots. The effectiveness of the chelates varied depending on their capacity to retain Zn in a soluble form in the soil solution.It is evident that zinc nutrition of plants in alkaline and calcareous soils can be more effectively regulated by both synthetic and natural chelates or organic manures which contain substantial amount of complexed zinc.Journal Paper No. 1 from the Department of Soil Science and Agric. Chemistry, Tirhut College of Agriculture, Dholi, Muzaffarpur, Bihar, India.     

Does increased zinc uptake enhance grain zinc mass concentration in rice?

Rice (Oryza sativa) is the worlds’ most important cereal and potentially an important source of zinc (Zn) for people who eat mainly rice. To improve Zn delivery by rice, plant Zn uptake and internal allocation need to be better understood. This study reports on within‐plant allocation and potential Zn accumulation in the rice grain in four so‐called aerobic rice cultivars (Handao297, K150, Handao502 and Baxiludao). Two controlled‐condition experiments were carried out, one with a wide range of constant Zn concentrations in the medium and one with a range of plant growth rate‐related supply rates. In both experiments, increased Zn supply induced increased plant Zn uptake rate throughout crop development, when expressed as daily Zn uptake (μg day^?1) or as daily Zn uptake per gram of plant dry matter (μg g^?1). Zinc mass concentration (ZnMC) in all plant organs increased with an increase in Zn supply but to various degrees. At higher uptake levels, the ZnMC in stems increased most, while the ZnMC in hulled grains (brown rice) increased least. The increase in leaf ZnMC was generally small, but at toxic levels in the medium, leaf ZnMC increased significantly. It appears that regulation of grain Zn loading differs from regulation of Zn loading to other organs. A milling test on seeds of Baxiludao and Handao502 showed that when ZnMC in brown rice increased from 13 to 45 mg kg^?1, ZnMC in polished rice grains (endosperm) also increased from 9 to 37 mg kg^?1 but remained three to five times lower than that in the bran. Irrespective of the ZnMC in the brown rice, around 75% of total grain Zn was present in the endosperm. In both cultivars, there was a major difference in ZnMC between bran and endosperm (120 and 37 mg kg^?1, respectively), suggesting a barrier for Zn transport between the two tissues. There seems to be a second barrier between stem and rachis, as their ZnMCs also differed greatly (300 and 100 mg kg^?1, respectively) in both cultivars at higher plant ZnMC. It is concluded that there is too little scope from a human nutrition perspective to enhance ZnMC in rice endosperm by simply increasing the Zn supply to rice plants because Zn allocation to the endosperm is limited, while observed genotypic differences indicate scope for improvement through breeding.     

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.     

Efficiency of zinc fertilization for flooded rice

Summary Several Zn-deficient soils from the major rice growing areas in the USA were characterized with respect to extractable and labile Zn and other parameters including available P, pH, organic matter, and cation exchange capacity. A greenhouse experiment was conducted to determine whether the above factors influence the response of several rice cultivars to ZnSO₄ or ZnEDTA applied as a mixed preplant treatment or to the floodwater at panicle initiation.Response of the rice to Zn varied widely among soils with labile Zn (L value) being a poor indicator of Zn availability and soil pH, and DTPA or 2 N MgCl₂ extraction of Zn being the most reliable indices. Soil incorporation prior to flooding was more effective than floodwater application for the initial crop, but both methods of placement were comparable for a subsequent ratoon crop. In either situation, ZnSO₄ was superior to ZnEDTA. Recovery of Zn from ZnSO₄ was generally less than 5 percent where Zn response was obtained and up to 14 percent from nonresponsive soils.     

From Flooded to Aerobic Conditions in Rice Cultivation: Consequences for Zinc Uptake

Scarcity of water causes a shift from flooded to aerobic conditions for rice production in zinc deficient areas in Northern China. This shift alters soil conditions that affect zinc availability to the crop. This paper concerns the effect of aerobic compared to flooded conditions on crop biomass production, grain yield and zinc content. A field experiment was done with six rice genotypes (Oryza sativa L.) grown on a calcareous soil, both with (23 kg Zn ha⁻¹) and without Zn fertilization. Sampling was conducted at tillering and physiological mature stage. Zn concentration in the shoots was significantly lower at both stages in plants grown in the aerobic field. At maturity, Zn uptake, biomass production, grain yield and Zn-harvest index [grain Zn/(shoot + grain Zn)] were lower under aerobic cultivation. Rice genotypes including aerobic rice and lowland rice differ in degree of response to low Zn supply. A twofold difference was found among aerobic genotypes in grain yield and Zn uptake. Also Zn-harvest index varied significantly. Zn application affected neither grain yield nor grain Zn content, although it significantly improved biomass production in both systems in most genotypes. These results demonstrate that introduction of aerobic rice systems on calcareous soils may increase Zn deficiency problems.     

Relative effectiveness of various types and methods of zinc application on rice and maize crops grown in calcareous soil

Summary Application of zinc sulphate mixed with compost/poultry manure proved to be equivalent to the effect of dipping the seedling roots in 4% ZnO suspension with respect to rice yields but Zn-amended organic manures were superior to other treatments with regards to total Zn uptake. A marked residual effect of soil applied treatments was recorded on the succeeding maize crop. Application of poultry manure alone was about one and a half times more effective than compost alone in increasing the rice and maize grain yields. Poultry manure surpassed compost in increasing zinc uptake by the crops and at the same time it built up more available Zn in soil than compost for the following crop. The magnitude of yields and Zn uptake response were magnified when zinc sulphate was applied along with organic manures. Application of 25 kg zinc sulphate/ha alone had the same effect as 50 quintals poultry manure alone or 12.5 kg zinc sulphate applied with 50 quintals compost/ha with respect to crop yields. A significant positive correlation was, recorded in both the crops between Zn concentration in grain or straw and their respective yields.     

有机物料对滨海盐渍土重金属根际效应的影响Ⅰ.土壤内源Zn形态分布

研究了排水条件下施用腐熟有机物料、种稻改良滨海盐渍土内源Ｚｎ形态分布．结果表明,单淹水使土中各形态Ｚｎ一定程度上向生物有效性较低的Ｚｎ形态转化,有效态Ｚｎ降低．土壤盐分量不同,明显影响无定形氧化铁结合态、紧结有机态以及硅酸盐矿物态Ｚｎ变化．种稻不施有机物料,根际交换态和硅酸盐矿物态Ｚｎ亏缺;碳酸盐结合态、氧化锰结合态和无定形氧化铁结合态Ｚｎ富集．根际Ｚｎ形态转化强度大于非根际,其有效态Ｚｎ量接近临界值并高于非根际．有机物料利于根际内外土壤中硅酸盐矿物态Ｚｎ的转化,低盐土壤根际更强烈．随有机物料用量增加,促使根际硅酸盐矿物态、碳酸盐结合态及氧化锰结合态Ｚｎ向交换态、紧结有机态和无定形氧化铁结合态Ｚｎ转化,低盐土壤较明显．     

Different performances of soil phosphate tests for reflecting the effects of buffering capacity on uptake of native phosphate with time

The ability of two sodium bicarbonate (Colwell and Olsen) and two ammonium fluoride (Bray I and Bray II) soil tests to reflect the effect of phosphate buffering capacity of the soil on plant growth through time was studied on ten Argentine soils. The soils were divided into three groups (low, medium and high buffering capacity) according to a buffering index calculated from the slope of the Freundlich equation. The relation between phosphate extracted by soil tests and both relative yield and phosphate uptake of rye grass plants was affected by the phosphate buffering capacity of the soil. The effect of buffering on that relation was more marked for the sodium bicarbonate tests (specially Colwell) than for the Bray tests. This effect was consistent with time. Hence, adjustment for buffering would be more important for the sodium bicarbonate tests than for the Bray tests. Soils with high buffering capacity were able to sustain a greater rate of phosphate uptake. The effect of buffering on the relation between soil tests and both relative yield and phosphate uptake was greatest when the plants were young and decreased with time. This effect would therefore be very important for the early nutrition of annual pasture or crop species.     

有机物料对滨海盐渍土重金属根际效应的影响 Ⅰ.土壤内源Zn形态分布

研究了排水条件下施用腐熟有机物料、种稻改良滨海盐渍土内源Zn形态分布。结果表明,单淹水使土中各形态Zn一定程度上向生物有效性较低的Zn形态转化,有效态Zn降低。土壤盐分量不同,明显影响无定形氧化铁结合态、紧结有机态以及硅酸盐矿物态Zn变化。种稻不施有机物料,根际交换态和硅酸盐矿物态Zn亏缺;碳酸盐结合态、氧化锰结合态和无定形氧化铁结合态Zn富集。根际Zn形态转化强度大于非根际,其有效态Zn量接近临界值并高于非根际。有机物料利于根际内外土壤中硅酸盐矿物态Zn的转化,低盐土壤根际更强烈。随有机物料用量增加,促使根际硅酸盐矿物态、碳酸盐结合态及氧化锰结合态Zn向交换态、紧结有机态和无定形氧化铁结合态Zn转化,低盐土壤较明显。     

Improved nitrogen nutrition enhances root uptake, root-to-shoot translocation and remobilization of zinc ((65) Zn) in wheat

This study focussed on the effect of increasing nitrogen (N) supply on root uptake and root-to-shoot translocation of zinc (Zn) as well as retranslocation of foliar-applied Zn in durum wheat (Triticum durum). Nutrient solution experiments were conducted to examine the root uptake and root-to-shoot translocation of (65) Zn in seedlings precultured with different N supplies. In additional experiments, the effect of varied N nutrition on retranslocation of foliar-applied (65) Zn was tested at both the vegetative and generative stages. When N supply was increased, the (65) Zn uptake by roots was enhanced by up to threefold and the (65) Zn translocation from roots to shoots increased by up to eightfold, while plant growth was affected to a much smaller degree. Retranslocation of (65) Zn from old into young leaves and from flag leaves to grains also showed marked positive responses to increasing N supply. The results demonstrate that the N-nutritional status of wheat affects major steps in the route of Zn from the growth medium to the grain, including its uptake, xylem transport and remobilization via phloem. Thus, N is a critical player in the uptake and accumulation of Zn in plants, which deserves special attention in biofortification of food crops with Zn.     

Enhancing phosphorus and zinc acquisition efficiency in rice: a critical review of root traits and their potential utility in rice breeding

Background

Rice is the world''s most important cereal crop and phosphorus (P) and zinc (Zn) deficiency are major constraints to its production. Where fertilizer is applied to overcome these nutritional constraints it comes at substantial cost to farmers and the efficiency of fertilizer use is low. Breeding crops that are efficient at acquiring P and Zn from native soil reserves or fertilizer sources has been advocated as a cost-effective solution, but would benefit from knowledge of genes and mechanisms that confer enhanced uptake of these nutrients by roots.

Scope

This review discusses root traits that have been linked to P and Zn uptake in rice, including traits that increase mobilization of P/Zn from soils, increase the volume of soil explored by roots or root surface area to recapture solubilized nutrients, enhance the rate of P/Zn uptake across the root membrane, and whole-plant traits that affect root growth and nutrient capture. In particular, this review focuses on the potential for these traits to be exploited through breeding programmes to produce nutrient-efficient crop cultivars.

Conclusions

Few root traits have so far been used successfully in plant breeding for enhanced P and Zn uptake in rice or any other crop. Insufficient genotypic variation for traits or the failure to enhance nutrient uptake under realistic field conditions are likely reasons for the limited success. More emphasis is needed on field studies in mapping populations or association panels to identify those traits and underlying genes that are able to enhance nutrient acquisition beyond the level already present in most cultivars.     

Silicon suppresses zinc uptake through down-regulating zinc transporter gene in rice

One of the beneficial effects of silicon (Si) is to improve nutrient imbalance including deficiency and excess of nutrients, however the molecular mechanisms underlying this effect are still poorly understood. In this study, we investigated the interaction between Si and zinc (Zn) in rice by using a mutant (lsi1) defective in Si uptake and its wild-type (WT, cv. Oochikara) at different Zn levels. High Zn inhibited the root elongation of both WT and lsi1 mutant, but Si did not alleviate this inhibition in both lines. By contrast, Si supply decreased Zn concentration in both the roots and shoots of the WT, but not in the lsi1 mutant. A short-term (24 h) labeling experiment with stable isotope ⁶⁷Zn showed that Si decreased ⁶⁷Zn uptake, but did not affect the root-to-shoot translocation and distribution ratio to different organs of ⁶⁷Zn in the WT. Furthermore, Si accumulated in the shoots, rather than Si in the external solution, is required for suppressing Zn uptake, but this was not caused by Si-decreased transpiration. A kinetic study showed that Si did not affect K_m value of root Zn uptake, but decreased V_max value in the WT. Analysis of genes related with Zn transport showed that among ZIP family genes, the expression of only OsZIP1 implicated in Zn uptake, was down-regulated by Si in the WT, but not in the lsi1 mutant. These results indicate that Si accumulated in the shoots suppresses the Zn uptake through down-regulating the transporter gene involved in Zn uptake in rice.     

Zinc deficiency-inducible OsZIP8 encodes a plasma membrane-localized zinc transporter in rice

Zinc is an essential micronutrient for several physiological and biochemical processes. To investigate its transport in rice, we characterized OsZIP8, a rice ZIP (Zrt, Irt-like Protein) gene that is strongly up-regulated in shoots and roots under Zn deficiency. OsZIP8 could complement the growth defect of Zn-uptake yeast mutant. The OsZIP8-GFP fusion proteins were localized to the plasma membrane, suggesting that OsZIP8 is a plasma membrane zinc transporter in rice. Activation and overexpression of this gene disturbed the zinc distribution in rice plants, resulting in lower levels in shoots and mature seeds, but an increase in the roots. Field-grown transgenic plants were shorter than the WT. Under treatment with excess zinc, transgenics contained less zinc in their shoots but accumulated more in the roots. Altogether, these results demonstrate that OsZIP8 is a zinc transporter that functions in Zn uptake and distribution. Furthermore, zinc homeostasis is important to the proper growth and development of rice.     

Rhizosphere pH as a result of nitrogen levels and NH4/NO3 ratio and its effect on zinc availability and on growth of rice flower (Ozothamnus diosmifolius)

The effect of modification of the rhizosphere pH, via solution-N concentration and source, on rice flower (Ozothamnus diosmifolius, Astraceae) growth was investigated in two different experiments. In order to simulate a wide range of pHs easily, the plants were grown in an inert artificial substrate (perlite). In the first the rhizosphere pH was modified through variation of N concentrations and the NH₄/NO₃-N ratio in the irrigation water. In the second the rhizosphere pH was modified solely by altering the NH₄/NO₃-N ratio while irrigation-N concentration was held at the level found to be optimal in the first experiment. Cultivation of rice flower, a new crop in Israel, is hampered by lack of knowledge on its Zn nutrition. Because availability of soil Zn largely depends on pH we investigated in the second experiment the effect of Zn foliar application. The growth of rice flower plants under low-N fertilization or low NH₄/NO₃-N ratio was poor and the plants exhibited growth disorders such as tipburn, severe chlorosis and necrosis. These growth disorders could not be ascribed to any direct effect of N nutrition therefore it was suggested that the indirect effect of the treatments, e.g., the rhizosphere pH dominates rice flower growth through its effect on nutrient availability. The only nutrient that was significantly correlated with pH and yield parameters in both experiments was Zn. All irrigation-nutrients concentrations were within the recommended range for hydroponically grown plants; however, the leaf-Mn concentration of plants grown in pH above 7.5 was in the toxic range while that of Zn was deficient. The high preferential uptake of Mn over Zn by rice flower plants and the question of whether high Mn uptake induced Zn deficiency remain open.     

Understanding zinc reactions in vertisols to predict zinc responses by wheat

Since Zn availability to plants growing in a soil is governed by quantity, intensity, buffer power, rate of Zn desorption and diffusion, an improved understanding of a number of these factors in Vertisols would facilitate a more reliable prediction of crop requirements for Zn. The DTPA-extractable Zn, a quantity factor, together with initial Zn desorption rate coefficients, accounted for 80% of the variation in relative dry matter yield of wheat grown to anthesis. In combination with these factors, desorption (buffer) power explained 92% of the variation in Zn concentration in the young mature leaf blade (YMB) of wheat. Thus, the combination of the quantity, rate of Zn desorption and buffer power better predict growth responses of wheat to applied Zn in Vertisols than the commonly-used single extraction with DTPA alone (quantity), which provides only a static measure of Zn availability.     

Effect of zinc fertilization on the mineral nutrition of rices differing in tolerance to zinc deficiency

Summary The effects of four Zn levels on the electrochemical and chemical properties of the soil solution, and on the growth and mineral nutrition of two rice varieties (IR26 and IR34) differing in tolerance to Zn deficiency were studied in the greenhouse using Zn-deficient soils from two locations. A similar experiment was conducted in culture solution to check how Zn addition affects translocation of other nutrients.In both soil and culture solution, plant Zn concentrations alone was not enough to account for varietal tolerance to Zn deficiency. Comparison of nutrient to Zn and shoot to root ratios of nutrients was more useful in determining the possible mechanism of varietal tolerance. IR 34 appeared to tolerate the disorder due to its lower Zn requirement, more efficient Zn translocation and ability to maintain lower Fe/Zn, Cu/Zn, Mg/Zn and P/Zn ratios in the shoot than the more susceptible variety, IR26. This was shown to be due to decreased translocation of Fe, Mg and P to shoots and decreased absorption of Cu by the root in IR34 in culture solution studies. Adding Zn further reduces translocation or absorption of these nutrients and depending on the nutrient supply of the soil, could cause deficiencies or mineral imbalances, especially of Fe, Cu, and P.These observed varietal differences regarding Zn requirement and the interaction of Zn with absorption and translocation of plant nutrients necessitates revision of recommendations for Zn fertilization. There is an inevitable need for Zn application in severely Zn-deficient soils regardless of rice variety. But on marginally Zn-deficient soils especially those low in Fe, Cu, or P, Zn fertilization is not advisable when resistant rice varieties are used.     

The role of arbuscular mycorrhizas in improving plant zinc nutrition under low soil zinc concentrations: a review

Many of the world’s soils are zinc (Zn) deficient. Consequently, many crops experience reduced growth, yield and tissue Zn concentrations. Reduced concentrations of Zn in the edible portions of crops have important implications for human Zn nutrition; this is a cause of global concern. Most terrestrial plant species form arbuscular mycorrhizas (AM) with a relatively limited number of specialized soil fungi. Arbuscular mycorrhizal fungi (AMF) can take up nutrients, including Zn, and transfer them to the plant, thereby enhancing plant nutrition. Under high soil Zn concentrations the formation of AM can also ‘protect’ against the accumulation of Zn in plant tissues to high concentrations. Here, a short review focusing on the role of AM in enhancing plant Zn nutrition, principally under low soil Zn concentrations, is presented. Effects of Zn on the colonisation of roots by AMF, direct uptake of Zn by AMF, the role of AM in the Zn nutrition of field grown plants, and emerging aspects of Zn molecular physiology of AM, are explored. Emergent knowledge gaps are identified and discussed in the context of potential future research.