RESUMEN

Nitrate isotopes (δ15N-NO3- and δ18O-NO3-) are a potentially powerful tool for tracking the biological removal of reactive nitrogen (N) as it is transported from land to sea. However, uncertainties about, 1) the variability of the strength of biological isotopic fractionation during anaerobic benthic NO3- reduction (the kinetic enrichment factor: εdenit), and, 2) how accurately these εdenit values are expressed in overlying aerobic surface waters (the effective enrichment factor: εeff), currently limit their use in freshwater systems. Here we used a combination of incubation experiments and numerical modelling to construct a simple framework for defining freshwater εdenit based on interactions between benthic denitrification and diffusive transport to surface waters. Under non-limited, anaerobic conditions the εdenit values produced in submerged soils (n = 3) and sediments (n = 4) with denitrification rates between 10 and 600 mg N m-2 d-1 ranged from -3‰ to -28‰. Critically, model results indicated that diffusive transport would homogenise this to an effective fractionation range of -6 ±â€¯4‰. Evidence for biological and hydrological variability of NO3- isotope fractionation means that values measured in aerobic surface water environments are most appropriately evaluated by a range of fractionation values, rather than commonly used single 'site specific' εdenit values.

RESUMEN

We sought to explain rice (Oryza sativa) genotype differences in tolerance of zinc (Zn) deficiency in flooded paddy soils and the counter-intuitive observation, made in earlier field experiments, that Zn uptake per plant increases with increasing planting density. We grew tolerant and intolerant genotypes in a Zn-deficient flooded soil at high and low planting densities and found (a) plant Zn concentrations and growth increased with planting density and more so in the tolerant genotype, whereas the concentrations of other nutrients decreased, indicating a specific effect on Zn uptake; (b) the effects of planting density and genotype on Zn uptake could only be explained if the plants induced changes in the soil to make Zn more soluble; and (c) the genotype and planting density effects were both associated with decreases in dissolved CO2 in the rhizosphere soil solution and resulting increases in pH. We suggest that the increases in pH caused solubilization of soil Zn by dissolution of alkali-soluble, Zn-complexing organic ligands from soil organic matter. We conclude that differences in venting of soil CO2 through root aerenchyma were responsible for the genotype and planting density effects.

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Understanding Zn uptake dynamics is critical to rice grain Zn biofortification. Here we examined soil Zn availability and Zn uptake pathways as affected by genotype (high-grain Zn varieties IR69428 and IR68144), Zn fertilization and water management in two pot experiments. Results showed significant interactions (P < 0.05) between genotypes and Zn fertilization on DTPA (diethylenetriaminepentaacetic acid)-extractable soil Zn from early tillering to flowering. DTPA-extractable Zn in soils grown with IR69428 was positively correlated with stem (r = 0.78, P < 0.01), flagleaf (r = 0.60, P < 0.01) and grain (r = 0.67, P < 0.01) Zn concentrations, suggesting improved soil Zn availability and continued soil Zn uptake by IR69428 even at maturity. Conversely for IR68144, DTPA-extractable Zn was positively correlated only with leaf Zn uptake (r = 0.60, P < 0.01) at active tillering, indicating dependence on remobilization for grain Zn loading. Furthermore, the highest grain Zn concentration (P < 0.05) was produced by a combination of IR69428 and Zn fertilization applied at panicle initiation (38.5 µg g-1) compared with other treatments (P < 0.05). The results highlight that Zn uptake behavior of a rice genotype determines the fate of Zn from the soil to the grain. This has implications on overcoming Zn translocation barriers between vegetative parts and grains, and achieving grain Zn biofortification targets (30.0 µg g-1).

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RESUMEN

Enriching iron (Fe) and zinc (Zn) content in rice grains, while minimizing cadmium (Cd) levels, is important for human health and nutrition. Natural genetic variation in rice grain Zn enables Zn-biofortification through conventional breeding, but limited natural Fe variation has led to a need for genetic modification approaches, including over-expressing genes responsible for Fe storage, chelators, and transporters. Generally, Cd uptake and allocation is associated with divalent metal cations (including Fe and Zn) transporters, but the details of this process are still unknown in rice. In addition to genetic variation, metal uptake is sometimes limited by its bioavailability in the soil. The availability of Fe, Zn, and Cd for plant uptake varies widely depending on soil redox potential. The typical practice of flooding rice increases Fe while decreasing Zn and Cd availability. On the other hand, moderate soil drying improves Zn uptake but also increases Cd and decreases Fe uptake. Use of Zn- or Fe-containing fertilizers complements breeding efforts by providing sufficient metals for plant uptake. In addition, the timing of nitrogen fertilization has also been shown to affect metal accumulation in grains. The purpose of this mini-review is to identify knowledge gaps and prioritize strategies for improving the nutritional value and safety of rice.

RESUMEN

Zinc (Zn) deficiency is a major constraint to rice production and Zn is also often deficient in humans with rice-based diets. Efforts to breed more Zn-efficient rice are constrained by poor understanding of the mechanisms of tolerance to deficiency. Here we assess the contributions of root growth and root Zn uptake efficiency, and we seek to explain the results in terms of specific mechanisms. We made a field experiment in a highly Zn-deficient rice soil in the Philippines with deficiency-tolerant and -sensitive genotypes, and measured growth, Zn uptake and root development. We also measured the effect of planting density. Tolerant genotypes produced more crown roots per plant and had greater uptake rates per unit root surface area; the latter was at least as important as root number to overall tolerance. Tolerant and sensitive genotypes took up more Zn per plant at greater planting densities. The greater uptake per unit root surface area, and the planting density effect can only be explained by root-induced changes in the rhizosphere, either solubilizing Zn, or neutralizing a toxin that impedes Zn uptake (possibly [Formula: see text] or Fe(2+)), or both. Traits for these and crown root number are potential breeding targets.

RESUMEN

The rice ingestion rate in Madagascar is among the highest globally; however studies concerning metal(loid) concentrations in Madagascar rice are lacking. For Madagascar unpolished rice (n = 51 landraces), levels of toxic elements (e.g., total mercury, methylmercury, arsenic and cadmium) as well as essential micronutrients (e.g., zinc and selenium) were uniformly low, indicating potentially both positive and negative health effects. Aside from manganese (Wilcoxon rank sum, p < 0.01), no significant differences in concentrations for all trace elements were observed between rice with red bran (n = 20)and brown bran (n = 31) (Wilcoxon rank sum, p = 0.06-0.91). Compared to all elements in rice,rubidium (i.e., tracer for phloem transport) was most positively correlated with methylmercury (Pearson'sr = 0.33, p < 0.05) and total mercury (r = 0.44, p < 0.05), while strontium (i.e., tracer for xylemtransport) was least correlated with total mercury and methylmercury (r < 0.01 for both), suggesting inorganic mercury and methylmercury were possibly more mobile in phloem compared to xylem.

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Zn deficiency is a widespread problem in rice (Oryza sativa L.) grown under flooded conditions, limiting growth and grain Zn accumulation. Genotypes with Zn deficiency tolerance or high grain Zn have been identified in breeding programmes, but little is known about the physiological mechanisms conferring these traits. A protocol was developed for growing rice to maturity in agar nutrient solution (ANS), with optimum Zn-sufficient growth achieved at 1.5 µM ZnSO4.7H2O. The redox potential in ANS showed a decrease from +350 mV to -200 mV, mimicking the reduced conditions of flooded paddy soils. In subsequent experiments, rice genotypes contrasting for Zn deficiency tolerance and grain Zn were grown in ANS with sufficient and deficient Zn to assess differences in root uptake of Zn, root-to-shoot Zn translocation, and in the predominant sources of Zn accumulation in the grain. Zn efficiency of a genotype was highly influenced by root-to-shoot translocation of Zn and total Zn uptake. Translocation of Zn from root to shoot was more limiting at later growth stages than at the vegetative stage. Under Zn-sufficient conditions, continued root uptake during the grain-filling stage was the predominant source of grain Zn loading in rice, whereas, under Zn-deficient conditions, some genotypes demonstrated remobilization of Zn from shoot and root to grain in addition to root uptake. Understanding the mechanisms of grain Zn loading in rice is crucial in selecting high grain Zn donors for target-specific breeding and also to establish fertilizer and water management strategies for achieving high grain Zn.

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RESUMEN

One of the important factors that influences Zn deficiency tolerance and grain Zn loading in crops is the within-plant allocation of Zn. Three independent experiments were carried out to understand the internal Zn distribution patterns in rice genotypes grown in Zn-sufficient and Zn-deficient agar nutrient solution (ANS). In one of the experiments, two rice genotypes (IR55179 and KP) contrasting in Zn deficiency tolerance were leaf-labeled with (65)Zn. In the other two experiments, two Zn biofortification breeding lines (IR69428 and SWHOO) were either root- or leaf-labeled with (65)Zn. Rice genotype IR55179 showed significantly higher Zn deficiency tolerance than KP at 21 and 42 days after planting. When KP was Zn-deficient, it failed to translocate (65)Zn from the labeled leaf to newly emerging leaves. Similarly, the root-to-shoot translocation of unlabeled Zn was lower in KP than in IR55179. These results suggest that some Zn-efficient rice genotypes have greater ability to translocate Zn from older to actively growing tissues than genotypes sensitive to Zn deficiency. Among the two Zn biofortication breeding lines that were leaf-labeled with (65)Zn at 10 days before panicle initiation stage, (65)Zn distribution in the grains at maturity was similar between both genotypes in Zn-sufficient conditions. However, under Zn-deficient conditions, SWHOO accumulated significantly higher (65)Zn in grains than IR69428, indicating that SWHOO is a better remobilizer than IR69428. When the roots of these two Zn biofortication breeding lines were exposed to (65)Zn solution at 10 days after flowering, IR69428 showed higher root uptake of (65)Zn than SWHOO in Zn-sufficient conditions, but (65)Zn allocation in the aerial parts of the plant was similar between both genotypes.