Zinc nutrition in rice production systems: a review

Background

Zinc (Zn) deficiency is one of the important abiotic factors limiting rice productivity worldwide and also a widespread nutritional disorder affecting human health. Given that rice is a staple for populations in many countries, studies of Zn dynamics and management in rice soils is of great importance.

Scope

Changing climate is forcing the growers to switch from conventional rice transplanting in flooded soils to water-saving cultivation, including aerobic rice culture and alternate wetting and drying system. As soil properties are changed with altered soil and water management, which is likely to affect Zn solubility and plant availability and should be considered before Zn management in rice. In this review, we critically appraise the role of Zn in plant biology and its dynamics in soil and rice production systems. Strategies and options to improve Zn uptake and partitioning efficiency in rice by using agronomic, breeding and biotechnological tools are also discussed.

Conclusions

Although soil application of inorganic Zn fertilizers is widely used, organic and chelated sources are better from economic and environmental perspectives. Use of other methods of Zn application (such as seed treatment, foliar application etc., in association with mycorrhizal fungi) may improve Zn-use efficiency in rice. Conventional breeding together with modern genomic and biotechnological tools may result in development of Zn-efficient rice genotypes that should be used in conjunction with judicious fertilization to optimize rice yield and grain Zn content.     

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.     

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.     

Bioavailability of zinc and phosphorus in calcareous soils as affected by citrate exudation

Aims

Zinc (Zn) and phosphorus (P) deficiency often occurs at the same time and limits crop production in many soils. It has been suggested that citrate root exudation is a response of plants to both deficiencies. We used white lupin (Lupinus albus L.) as a model plant to clarify if citrate exuded by roots could increase the bioavailability of Zn and P in calcareous soils.

Methods

White lupin was grown in nutrient solution and in two calcareous soils in a rhizobox. Rhizosphere soil solution was sampled to determine citrate, metals and P. Based on the measured citrate concentrations, a soil extraction experiment with citrate as extractant was done.

Results

Absence of Zn triggered neither cluster root formation nor citrate exudation of white lupin grown in nutrient solution, whereas low P supply did. The maximum citrate concentration (～1.5?mM) found in the cluster rhizosphere soil solution of one soil mobilized P, but not Zn. In the other soil the highest citrate concentration (～0.5?mM) mobilized both elements.

Conclusions

White lupin does not respond to low Zn bioavailability by increasing citrate exudation. Such a response was observed at low P supply only. Whether Zn and P can be mobilized by citrate is soil-dependent and the possible controlling mechanisms are discussed.     

Isotopic fractionation of Zn in tomato plants suggests the role of root exudates on Zn uptake

Aims

Phytosiderophore-chelated Zn can be absorbed in grasses. Root exudates of dicotyledonous plants can mobilize soil Zn but it is unclear how this affects Zn bioavailability. Stable Zn isotope shifts can indicate exudate-facilitated Zn uptake, since complexation of Zn²⁺ by organic ligands in solution yields a small, but detectable, enrichment of the heavy Zn isotope due to thermodynamic fractionation.

Methods

Tomato seedlings were grown in resin-buffered nutrient solution in which free Zn²⁺ concentrations are buffered, in a factorial design of two Zn levels and two solution volumes. The latter factor allowed altering the exudate concentrations in the solution. Dissolved Cu concentrations in the resin buffered system were used as a sensitive index of metal mobilization resulting from root activity. In addition, seedlings were grown in Zn deficient soil with and without Zn addition.

Results

The dissolved Cu concentration increased with Zn deficiency and was highest at the lowest solution volume, suggesting metal mobilization by root exudates. At low Zn supply, Zn in the plant was enriched in heavy Zn (⁶⁶Zn) and this was most pronounced at small solution volume. Similarly, Zn deficiency in soil enriched tomato shoot Zn with heavy isotope in this plant.

Interpretation

Zinc deficiency increases the contribution of Zn-exudate complexes, which are enriched in the heavy isotope compared to the free ion, to Zn uptake by transporting Zn from the bulk solution or soil to the roots where they likely dissociate and release Zn²⁺.     

Dynamics of plant metal uptake and metal changes in whole soil and soil particle fractions during repeated phytoextraction

Aims

Phytoextration of metal polluted soils using hyperaccumulators is a promising technology but requires long term successive cropping. This study investigated the dynamics of plant metal uptake and changes in soil metals over a long remediation time.

Methods

A soil slightly polluted with metals (S1) was mixed with highly polluted soil (S4) to give two intermediate pollution levels (S2, S3). The four resulting soils were repeatedly phyto-extracted using nine successive crops of Cd/Zn-hyperaccumulator Sedum plumbizincicola over a period of 4 years.

Results

Shoot Cd concentration decreased with harvest time in all soils but shoot Zn declined in S1 only. Similar shoot Zn concentrations were found in S2, S3 and S4 although these soils differed markedly in metal availability, and their available metals decreased during phytoextraction. A possible explanation is that plant active acquisition ability served to maintain plant metal uptake. Plant uptake resulted in the largest decrease in the acid-soluble metal fraction followed by reducible metals. Oxidisable and residual fractions were less available to plants. The coarse soil particle fractions made the major contribution to metal decline overall than the fine fractions.

Conclusion

Sedum plumbizincicola maintained long term metal uptake and the coarse soil particles played the most important role in phytoextraction.     

Silicon-mediated amelioration of zinc toxicity in rice (Oryza sativa L.) seedlings

Background and aims

Silicon (Si) was suggested to enhance plant resistance to toxic elements, and its beneficial role was mainly based on external and internal plant mechanisms. This work aimed at investigating the internal effect of Si on zinc (Zn) detoxification to rice (Oryza sativa L., cv. Tian You 116) seedlings.

Methods

In a hydroponic experiment, we examined the uptake, xylem loading and localization of Zn in rice seedlings under the condition of 200?μM Zn contamination with the additional silicate supply at three levels ( 0, 0.5 and 1.8?mM).

Results

The silicate addition significantly increased the seedling biomass, and decreased Zn concentration in both root and shoot of seedlings and in xylem sap flow. Zinpyr-1 fluorescence test and Energy-dispersive X-ray spectroscopy analysis showed the concentration of biologically active Zn²⁺ decreased, and Zn and Si co-localized in the cell wall of metabolically less active tissues, especially in sclerenchyma of root. The fractionation analysis further supported silicate supply increased about 10% the cell wall bound fraction of Zn.

Conclusions

This study suggests the Si-assisted Zn tolerance of rice is mainly due to the reduction of uptake and translocation of excess Zn, and a stronger binding of Zn in the cell wall of less bioactive tissues might also contribute to some degree.     

Comparison of As sequestration in iron plaque and uptake by different genotypes of rice plants grown in As-contaminated paddy soils

Background and aims

Limited information is available on comparing the iron plaque formation capabilities and their effect on arsenic (As) uptake by different rice plant genotypes grown in As-contaminated soils. This study investigates the effect of iron plaque on As uptake in different rice genotypes grown in As-contaminated soils from the Guandu Plain of northern Taiwan.

Methods

Twenty-eight rice genotypes including 14 japonica and 14 indica genotypes were used in this study. Rice seedlings were grown in As-contaminated soils for 38 days. The iron plaque formed on the rice roots were extracted using dithionite–citrate–bicarbonate. The concentrations of As, Fe, and P in soil solutions, iron plaque, and plants were measured. The speciation of As in the root’s iron plaque was determined by As K-edge X-ray absorption near-edge structure spectroscopy (XANES).

Results

The amounts of iron plaque formation on roots were significantly different among 28 tested rice genotypes, and 75.7–92.8 % of As uptake from soils could be sequestered in iron plaque. However, there were no significant negative correlations between the amounts of Fe or As in the iron plaque and the content of As accumulated in rice plants of tested genotypes. XANES data showed that arsenate was the predominant As species in iron plaque, and there were difference in the distribution of As species among different rice genotypes.

Conclusions

The iron plaque can sequester most of As uptake from soils no matter what rice genotypes used in this study. However, the iron plaque alone did not control the extent of As accumulation in rice plants from As-contaminated soils among 28 tested rice genotypes. Low As uptake genotypes of rice selected from this study can be recommended to be grown in the As-contaminated soils.     

Root porosity, radial oxygen loss and iron plaque on roots of wetland plants in relation to zinc tolerance and accumulation

Background and aims

Wetland plants have been widely used in constructed wetlands for the clean-up of metal-contaminated waters. This study investigated the relationship between rate of radial oxygen loss (ROL), root porosity, Zn uptake and tolerance, Fe plaque formation in wetland plants.

Methods

A hydroponic experiment and a pot trial with Zn-contaminated soil were conducted to apply different Zn level treatments to various emergent wetland plants.

Results

Significant differences were found between plants in their root porosities, rates of ROL, Zn uptake and Zn tolerance indices in the hydroponic experiment, and concentrations of Fe and Mn on roots and in the rhizosphere in the pot trial. There were significant positive correlations between root porosities, ROL rates, Zn tolerance, Zn, Fe and Mn concentrations on roots and in the rhizosphere. Wetland plants with higher root porosities and ROL tended to have more Fe plaque, higher Zn concentrations on roots and in their rhizospheres, and were more tolerant of Zn toxicity.

Conclusions

Our results suggest that ROL and root porosity play very important roles in Fe plaque formation, Zn uptake and tolerance, and are useful criteria for selecting wetland plants for the phytoremediation of Zn-contaminated waters and soils/sediments.     

<Emphasis Type="Bold">Zinc nutrition in wheat-based cropping systems</Emphasis>

Background

Zinc (Zn) deficiency is one of the most important micronutrient disorders affecting human health. Wheat is the staple food for 35% of the world’s population and is inherently low in Zn, which increases the incidence of Zn deficiency in humans. Major wheat-based cropping systems viz. rice–wheat, cotton–wheat and maize–wheat are prone to Zn deficiency due to the high Zn demand of these crops.

Methods

This review highlights the role of Zn in plant biology and its effect on wheat-based cropping systems. Agronomic, breeding and molecular approaches to improve Zn nutrition and biofortification of wheat grain are discussed.

Results

Zinc is most often applied to crops through soil and foliar methods. The application of Zn through seed treatments has improved grain yield and grain Zn status in wheat. In cropping systems where legumes are cultivated in rotation with wheat, microorganisms can improve the available Zn pool in soil for the wheat crop. Breeding and molecular approaches have been used to develop wheat genotypes with high grain Zn density.

Conclusions

Options for improving grain yield and grain Zn concentration in wheat include screening wheat genotypes for higher root Zn uptake and grain translocation efficiency, the inclusion of these Zn-efficient genotypes in breeding programs, and Zn fertilization through soil, foliar and seed treatments.

     

Effects of biochar amendment on root traits and contaminant availability of maize plants in a copper and arsenic impacted soil

Background and aims

Biochar has been proposed as a tool to enhance phytostabilisation of contaminated soils but little data are available to illustrate the direct effect on roots in contaminated soils. This work aimed to investigate specific root traits and to assess the effect of biochar amendment on contaminant availability.

Methods

Amendment with two different types of biochar, pine woodchip and olive tree pruning, was assessed in a rhizobox experiment with maize planted in a soil contaminated with significant levels of copper and arsenic.

Results

Amendment was found to significantly improve root traits compared to the control soil, particularly root mass density and root length density. Copper uptake to plants and ammonium sulphate extractable copper was significantly less in the biochar amended soils. Arsenic uptake and extractability varied with type of biochar used but was not considered to be the limiting factor affecting root and shoot development.

Conclusions

Root establishment in contaminated soils can be enhanced by biochar amendment but choice of biochar is key to maximising soil improvement and controlling contaminant availability.     

Distribution of cutin and suberin biomarkers under forest trees with different root systems

Background and aims

The rhizosphere, the soil immediately surrounding roots, provides a critical bridge for water and nutrient uptake. The rhizosphere is influenced by various forms of root–soil interactions of which mechanical deformation due to root growth and its effects on the hydraulics of the rhizosphere are the least studied. In this work, we focus on developing new experimental and numerical tools to assess these changes.

Methods

This study combines X-ray micro-tomography (XMT) with coupled numerical simulation of fluid and soil deformation in the rhizosphere. The study provides a new set of tools to mechanistically investigate root-induced rhizosphere compaction and its effect on root water uptake. The numerical simulator was tested on highly deformable soil to document its ability to handle a large degree of strain.

Results

Our experimental results indicate that measured rhizosphere compaction by roots via localized soil compaction increased the simulated water flow to the roots by 27 % as compared to an uncompacted fine-textured soil of low bulk density characteristic of seed beds or forest topsoils. This increased water flow primarily occurred due to local deformation of the soil aggregates as seen in the XMT images, which increased hydraulic conductivity of the soil. Further simulated root growth and deformation beyond that observed in the XMT images led to water uptake enhancement of ~50 % beyond that due to root diameter increase alone and demonstrated the positive benefits of root compaction in low density soils.

Conclusions

The development of numerical models to quantify the coupling of root driven compaction and fluid flow provides new tools to improve the understanding of plant water uptake, nutrient availability and agricultural efficiency. This study demonstrated that plants, particularly during early growth in highly deformable low density soils, are involved in active mechanical management of their surroundings. These modeling approaches may now be used to quantify compaction and root growth impacts in a wide range of soils.     

Arbuscular mycorrhizas are beneficial under both deficient and toxic soil zinc conditions

Background and aims

Arbuscular mycorrhizas (AM) play different roles in plant Zn nutrition depending on whether the soil is Zn-deficient (AM enhancement of plant Zn uptake) or Zn-toxic (AM protection of plant from excessive Zn uptake). In addition, soil P concentration modifies the response of AM to soil Zn conditions. We undertook a glasshouse experiment to study the interactive effects of P and Zn on AM colonisation, plant growth and nutrition, focusing on the two extremes of soil Zn concentration—deficient and toxic.

Methods

We used a mycorrhiza-defective tomato (Solanum lycopersicum) genotype (rmc) and compared it to its wild-type counterpart (76R). Plants were grown in pots amended with five soil P addition treatments, and two soil Zn addition treatments.

Results

The mycorrhizal genotype generally thrived better than the non-mycorrhizal genotype, in terms of biomass and tissue P and Zn concentrations. This was especially true under low soil Zn and P conditions, however there was evidence of the ‘protective effect’ of mycorrhizas when soil was Zn-contaminated. Above- and below-ground allocation of biomass, P and Zn were significantly affected by AM colonisation, and toxic soil Zn conditions.

Conclusions

The relationship between soil Zn and soil P was highly interactive, and heavily influenced AM colonisation, plant growth, and plant nutrition.     

Uptake of zinc and phosphorus by plants is affected by zinc fertiliser material and arbuscular mycorrhizas

Background and Aims

Water solubility of zinc (Zn) fertilisers affects their plant availability. Further, simultaneous application of Zn and phosphorus (P) fertiliser can have antagonistic effects on plant Zn uptake. Arbuscular mycorrhizas (AM) can improve plant Zn and P uptake. We conducted a glasshouse experiment to test the effect of different Zn fertiliser materials, in conjunction with P fertiliser application, and colonisation by AM, on plant nutrition and biomass.

Methods

We grew a mycorrhiza-defective tomato genotype (rmc) and its mycorrhizal wild-type progenitor (76R) in soil with six different Zn fertilisers ranging in water solubility (Zn sulphate, Zn oxide, Zn oxide (nano), Zn phosphate, Zn carbonate, Zn phosphate carbonate), and supplemental P. We measured plant biomass, Zn and P contents, mycorrhizal colonisation and water use efficiency.

Results

Whereas water solubility of the Zn fertilisers was not correlated with plant biomass or Zn uptake, plant Zn and P contents differed among Zn fertiliser treatments. Plant Zn and P uptake was enhanced when supplied as Zn phosphate carbonate. Mycorrhizal plants took up more P than non-mycorrhizal plants; the reverse was true for Zn.

Conclusions

Zinc fertiliser composition and AM have a profound effect on plant Zn and P uptake.     

Positive feedbacks between decomposition and soil nitrogen availability along fertility gradients

Background and aims

We determined the relationship between site N supply and decomposition rates with respect to controls exerted by environment, litter chemistry, and fungal colonization.

Methods

Two reciprocal transplant decomposition experiments were established, one in each of two long-term experiments in oak woodlands in Minnesota, USA: a fire frequency/vegetation gradient, along which soil N availability varies markedly, and a long-term N fertilization experiment. Both experiments used native Quercus ellipsoidalis E.J. Hill and Andropogon gerardii Vitman leaf litter and either root litter or wooden dowels.

Results

Leaf litter decay rates generally increased with soil N availability in both experiments while belowground litter decayed more slowly with increasing soil N. Litter chemistry differed among litter types, and these differences had significant effects on belowground (but not aboveground) decay rates and on aboveground litter N dynamics during decomposition. Fungal colonization of detritus was positively correlated with soil fertility and decay rates.

Conclusions

Higher soil fertility associated with low fire frequency was associated with greater leaf litter production, higher rates of fungal colonization of detritus, more rapid leaf litter decomposition rates, and greater N release in the root litter, all of which likely enhance soil fertility. During decomposition, both greater mass loss and litter N release provide mechanisms through which the plant and decomposer communities provide positive feedbacks to soil fertility as ultimately driven by decreasing fire frequency in N-limited soils and vice versa.     

Acquisition of phosphorus and other poorly mobile nutrients by roots. Where do plant nutrition models fail?

Background

In the context of increasing global food demand, ecological intensification of agroecosystems is required to increase nutrient use efficiency in plants while decreasing fertilizer inputs. Better exploration and exploitation of soil resources is a major issue for phosphorus, given that rock phosphate ores are finite resources, which are going to be exhausted in decades from now on.

Scope

We review the processes governing the acquisition by plants of poorly mobile nutrients in soils, with a particular focus on processes at the root?Csoil interface. Rhizosphere processes are poorly accounted for in most plant nutrition models. This lack largely explains why present-day models fail at predicting the actual uptake of poorly mobile nutrients such as phosphorus under low input conditions. A first section is dedicated to biophysical processes and the spatial/temporal development of the rhizosphere. A second section concentrates on biochemical/biogeochemical processes, while a third section addresses biological/ecological processes operating in the rhizosphere.

Conclusions

New routes for improving soil nutrient efficiency are addressed, with a particular focus on breeding and ecological engineering options. Better mimicking natural ecosystems and exploiting plant diversity appears as an appealing way forward, on this long and winding road towards ecological intensification of agroecosystems.     

Predictability of the Zn and Cd phytoextraction efficiency of a Salix smithiana clone by DGT and conventional bioavailability assays

Aims

Phytomanagement of metal-polluted soils requires information on plant responses to metal availability in soil, but the predictability of metal accumulation in plant shoots and/or roots may be limited by metal toxicity and inherent shortfalls of the bioavailability assays.

Methods

We measured the uptake of Cd and Zn in a Salix smithiana clone grown in a pot experiment on soils with different characteristics and metal availabilities, determined by conventional soil single extractions (0.05 M Na₂-EDTA and 1 M NH₄NO₃), soil solution obtained by centrifugation, and diffusive gradients in thin films (DGT). The Cd and Zn phytoavailability after a 2-year phytoextraction by willow was assessed by metal accumulation in the straw of the following barley culture.

Results

The phytoextraction efficiency was largest on a moderately polluted acid soil. Biomass and shoot Zn concentrations of S. smithiana were better predicted by DGT-measured Zn concentrations in soil solution (C _DGT) than by Zn concentrations in the soil solution and extractable soil fractions. The weaker correlation for Cd in shoots may be related to relative Cd enrichment in the plant tissues. The metal accumulation in barley straw was unaffected or increased after a 2-year phytoextraction.

Conclusions

The shoot Zn and Cd removal of the tested Salix clone can be predicted by C _DGT concentrations and is highest on either calcareous or moderately polluted acid soils. Single extraction with NH₄NO₃ and the C _DGT value of Cd were not able to predict shoot Cd removal on the tested soils. Only shoot removal of Zn was predicted fairly well by the C _DGT value.