Zinc Deficiency Enhances the Induction of Micronuclei and 8-Hydroxy-2′-Deoxyguanosine Via Superoxide Radical in Bone Marrow of Zinc-Deficient Rats

The aim of the present study was to examine whether zinc (Zn) deficiency augmented the frequency of micronuclei, an indicator of chromosome aberration, and the induction of 8-hydroxy-2′-deoxyguanosine (8-OHdG), a marker of cellular DNA damage derived from oxidative stress, in rat bone marrow cells or not. Both the frequency of micronuclei and the induction of 8-OHdG were significantly increased in rats fed with a Zn-deficient versus a standard diet for 6 weeks (p?<?0.005). The supplementation of Zn with a standard diet for 4 weeks to rats fed with a Zn-deficient diet for 6 weeks restored the enhanced induction of micronuclei and 8-OHdG to levels comparable to those seen in rats fed with a standard diet for 10 weeks, indicating that the shortage of Zn in the body is involved in the induction of micronuclei and 8-OHdG. Again, the membrane-permeable superoxide dismutase mimetic superoxide scavenger, 4-hydroxy-2,2,6,6-tetramethylpiperidine-1-oxyl, treatment (100 μmol/kg, twice a day) for 10 days prior to the termination of dietary treatment reduced the induction of micronuclei and 8-OHdG in rats fed with a Zn-deficient diet for 6 weeks to levels comparable to those in rats fed with a standard diet for 6 weeks, indicating that superoxide radical participates in the induction of micronuclei and 8-OHdG. In fact, the endogenous superoxide scavenger, Cu/Zn superoxide dismutase, was significantly reduced in the bone marrow cells of rats fed with a Zn-deficient diet for 6 weeks when compared to those of rats fed with a standard diet for 6 weeks (p?<?0.005). These observations demonstrate that Zn deficiency elevates the frequency of micronuclei and the induction of 8-OHdG through an increase in the biological action of the superoxide radical. This suggests an increase in carcinogenic initiation resulting from Zn deficiency-induced oxidative stress.     

Ecotypes of Holcus lanatus Tolerant to Zinc Toxicity also Tolerate Zinc Deficiency

Genotypes tolerant to zinc (Zn) toxicity, if they accumulateZn in their roots, may grow better than Zn-sensitive genotypes,even in Zn-deficient soil. In the present study, Holcus lanatusL. ecotypes differing in tolerance to Zn toxicity were grownin Zn-deficient Laffer soil which was amended with Zn to createa range of conditions from Zn deficiency to Zn toxicity. IncreasingZn additions to the soil, up to the sufficiency level, improvedgrowth of all ecotypes. At toxic levels of added Zn, the Zn-sensitiveecotype suffered a greater decrease in growth than the Zn-tolerantecotypes. All ecotypes accumulated more Zn in roots than inshoots, with root concentrations exceeding 8 g Zn kg^-1dry weightin extreme cases. When grown in Zn-deficient or Zn-sufficientsoil (up to 0.5 mg Zn kg^-1soil added), ecotypes tolerant toZn toxicity took up more Zn, grew better and had greater rootand shoot Zn concentration than the control (Zn-sensitive ecotype).Zn-tolerant ecotypes transported more Zn, copper (Cu) and iron(Fe) from roots to shoots in comparison with the Zn-sensitiveecotype. The average Zn uptake rate from Zn-deficient soil (noZn added) was greater in the Zn-tolerant ecotypes than in theZn-sensitive ecotype. In conclusion, ecotypes of H. lanatusthat are tolerant to Zn toxicity also tolerate Zn deficiencybetter than the Zn-sensitive ecotype because of their greatercapacity for taking up Zn from Zn-deficient soil. This is thefirst report of the coexistence of traits for tolerance to Zntoxicity and Zn deficiency in a single plant genotype. Copyright2000 Annals of Botany Company Copper, heavy metal, Holcus lanatus, iron, zinc deficiency, zinc toxicity     

Expression of high zinc efficiency of Aegilops tauschii and Triticum monococcum in synthetic hexaploid wheats

The effect of varied zinc (Zn) supply on shoot and root dry matter production, severity of Zn deficiency symptoms and Zn tissue concentrations was studied in two Triticum turgidum (BBAA) genotypes and three synthetic hexaploid wheat genotypes by growing plants in a Zn-deficient calcareous soil under greenhouse conditions with (+Zn=5 mg kg^-1 soil) and without (−Zn) Zn supply. Two synthetic wheats (BBAADD) were derived from two different Aegilops tauschii (DD) accessions using same Triticum turgidum (BBAA), while one synthetic wheat (BBAAAA) was derived from Triticum turgidum (BBAA) and Triticum monococcum (AA). Visible symptoms of Zn deficiency, such as occurrence of necrotic patches on leaves and reduction in shoot elongation developed more rapidly and severely in tetraploid wheats than in synthetic hexaploid wheats. Correspondingly, decreases in shoot and root dry matter production due to Zn deficiency were higher in tetraploid wheats than in synthetic hexaploid wheats. Transfer of the DD genome from Aegilops tauschii or the AA genome from Triticum monococcum to tetraploid wheat greatly improved root and particularly shoot growth under Zn-deficient, but not under Zn-sufficient conditions. Better growth and lesser Zn deficiency symptoms in synthetic hexaploid wheats than in tetraploid wheats were not accompanied by increases in Zn concentration per unit dry weight, but related more to the total amount of Zn per shoot, especially in the case of synthetic wheats derived from Aegilops tauschii. This result indicates higher Zn uptake capacity of synthetic wheats. The results demonstrated that the genes for high Zn efficiency from Aegilops tauschii (DD) and Triticum monococcum (AA) are expressed in the synthetic hexaploid wheats. These wheat relatives can be used as valuable sources of genes for improvement of Zn efficiency in wheat. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effect of zinc nutrition on salinity-induced oxidative damages in wheat genotypes differing in zinc deficiency tolerance

Zinc deficiency and salinity are well-documented soil problems and often occur simultaneously in cultivated soils. Usually, plants respond to environmental stress factors by activating their antioxidative defense mechanisms. The antioxidative response of wheat genotypes to salinity in relation to Zn nutrition is not well understood. So, we investigated the effect of Zn nutrition on the growth, membrane permeability and sulfhydryl group (–SH groups) content of root cells and antioxidative defense mechanisms of wheat plants exposed to salt stress. In a hydroponic experiment, three bread wheat genotypes (Triticum aestivum L. cvs. Rushan, Kavir, and Cross) with different Zn-deficiency tolerance were exposed to adequate (1 μM Zn) and deficient (no Zn) Zn supply and three salinity levels (0, 60, and 120 mM NaCl). The results obtained showed that adequate Zn nutrition counteracted the detrimental effect of 60 mM NaCl level on the growth of all three wheat genotypes while it had no effect on the root and shoot growth of ‘Rushan’ and ‘Kavir’ at the 120 mM NaCl treatment. At the 0 and 60 mM NaCl treatments, Zn application decreased root membrane permeability while increased –SH group content and root activity of catalase (CAT) and superoxide dismutase (SOD) in ‘Rushan’ and ‘Kavir’. In contrast, Zn had no effect on the root membrane permeability and –SH group content of ‘Rushan’ and ‘Kavir’ exposed to the 120 mM NaCl treatment. At all salinity levels, ‘Cross’ plants supplied with Zn had lower root membrane permeability and higher –SH group content compared to those grown under Zn-deficient conditions. At the 0 and 60 salinity levels, Zn-deficient roots of Kavir and Rushan genotype leaked significantly higher amounts of Fe and K than the Zn-sufficient roots. In contrast, at the 120 mM treatment, Zn application had no effect or slightly increased Fe and K concentration in the root ion leakage of these wheat genotypes. For ‘Cross’, at all salinity levels, Zn-deficient roots leaked significantly higher amounts of Fe and K compared with the Zn-sufficient roots. The differential tolerance to salt stress among wheat genotypes examined in this study was related to their tolerance to Zn-deficiency, –SH group content, and root activity of CAT and SOD. Greater tolerance to salinity of Zn-deficiency tolerant genotype ‘Cross’ is probably associated with its greater antioxidative defense capacity.     

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.     

Deoxymugineic acid increases Zn translocation in Zn-deficient rice plants

Deoxymugineic acid (DMA) is a member of the mugineic acid family phytosiderophores (MAs), which are natural metal chelators produced by graminaceous plants. Rice secretes DMA in response to Fe deficiency to take up Fe in the form of Fe(III)–MAs complex. In contrast with barley, the roots of which secrete MAs in response to Zn deficiency, the amount of DMA secreted by rice roots was slightly decreased under conditions of low Zn supply. There was a concomitant increase in endogenous DMA in rice shoots, suggesting that DMA plays a role in the translocation of Zn within Zn-deficient rice plants. The expression of OsNAS1 and OsNAS2 was not increased in Zn-deficient roots but that of OsNAS3 was increased in Zn-deficient roots and shoots. The expression of OsNAAT1 was also increased in Zn-deficient roots and dramatically increased in shoots; correspondingly, HPLC analysis was unable to detect nicotianamine in Zn-deficient shoots. The expression of OsDMAS1 was increased in Zn-deficient shoots. Analyses using the positron-emitting tracer imaging system (PETIS) showed that Zn-deficient rice roots absorbed less ⁶²Zn-DMA than ⁶²Zn²⁺. Importantly, supply of ⁶²Zn-DMA rather than ⁶²Zn²⁺ increased the translocation of ⁶²Zn into the leaves of Zn-deficient plants. This was especially evident in the discrimination center (DC). These results suggest that DMA in Zn-deficient rice plants has an important role in the distribution of Zn within the plant rather than in the absorption of Zn from the soil. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Motofumi Suzuki and Takashi Tsukamoto equally contributed to this work.     

Mitigating zinc deficiency and achieving high grain Zn in rice through integration of soil chemistry and plant physiology research

Background

Zinc deficiency has been recognized as an important factor affecting both human health and crop production. Rice (Oryza sativa) is relevant to both concerns, as it is susceptible to soil Zn deficiency and is a staple food for some of the Zn-deficient human population. Improving the processes by which Zn moves from the soil into the plant and eventually into the edible part of the grain has the potential to mitigate problems associated with Zn deficiency in crops and humans. This review article focuses on soil- and plant-related processes affecting Zn chemistry in rice-grown soils and Zn uptake and transport in a rice plant.

Scope

This review covers advances in soil chemistry regarding the reasons for inconsistent Zn deficiency in rice soils and the limitations of soil test methods for predicting Zn response for rice. We then review advances in plant physiology related to root Zn uptake and internal Zn distribution mechanisms in rice and explore interactions between specific root processes and the soil chemistry of particular environments. We aim to provide an overview of the soil science research for plant scientists and vice versa, in order to promote and facilitate future interdisciplinary collaborations.

Conclusions

Priority research areas to fill in knowledge gaps are: 1) improving our ability to predict Zn deficiency in rice soils, 2) understanding the relationship between Zn-deficiency tolerance mechanisms and grain Zn accumulation, 3) exploring the effectiveness of root Zn uptake mechanisms in contrasting soil environments.     

Effect of sulphate nutrition on arsenic translocation and photosynthesis of rice seedlings

The effect of sulphate nutrition on arsenic (As) concentration, photosynthetic and chlorophyll fluorescence parameters of rice was investigated in hydroponically grown rice seedlings (Oryza sativa L.), using three sulphate levels (1.8 μM, 0.7 mM, or 1.5 mM). The results showed that sulphate deficiency decreased As accumulation in root, but increased the translocation of As from root to shoot. Sulphate deficiency reduced maximum quantum yield (Fv/Fm), minimum fluorescence and electron transport rate (ETR) of a dark-adapted leaf. Compared with low sulphate treatments (1.8 μM), significant increases were observed in the parameters of rapid light curves, rETR_max and I _k of photosystem I (PSI) and photosystem II (PSII) of rice grown in the high sulphate treatments (1.5 mM) regardless of As additions. Therefore, an adequately high sulphate supply may result in less As translocation from root to shoot, and protecting the reaction pathways of PSI and PSII of rice seedlings grown in higher As-contaminated medium.     

A study of the role of root morphological traits in growth of barley in zinc-deficient soil

Zinc (Zn) deficiency reduces crop yields globally. This study investigated the importance of root morphological traits, especially root hairs, in plant growth and Zn uptake. Wild-type barley (Hordeum vulgare) Pallas and its root-hairless mutant brb were grown in soil and solution culture at different levels of Zn supply for 16 d. Root morphological traits (root length, diameter, and surface area) were measured using the WinRHIZOPro Image Analysis system. In soil culture, Pallas had greater shoot dry matter, shoot Zn concentration, shoot Zn content, and Zn uptake per cm(2) root surface area than brb, primarily under zinc deficiency. Both Pallas and brb developed longer roots under Zn deficiency. Development of root hairs was not affected by plant Zn status. In solution culture, there were no significant genotypic differences in any of the parameters measured, indicating that mutation in brb does not affect growth and Zn uptake. However, both Pallas and brb developed longer and thinner roots, and root hair growth was less than in soil culture, and was not affected by plant Zn status. The better growth and greater Zn uptake of Pallas compared with brb in Zn-deficient soil can be attributed primarily to greater root surface area due to root hairs in Pallas rather than other root morphological differences.     

Enrichment of fertilizers with zinc: An excellent investment for humanity and crop production in India

Micronutrient malnutrition is a growing concern in the developing world, resulting in diverse health and social problems, such as mental retardations, impairments of the immune system and overall poor health. In recent years, the zinc (Zn) deficiency problem has received increasing attention and appears to be the most serious micronutrient deficiency together with vitamin A deficiency. Zinc deficiency is particularly widespread among children and represents a major cause of child death in the world. In countries where Zn deficiency is well documented as an important public health problem, cereal-based foods are the predominant source of daily calorie and protein intake. Because the concentration of Zn in cereal crops is inherently very low, growing cereals on potentially Zn-deficient soils further decreases grain Zn concentrations. It is, therefore, not surprising that high Zn deficiency incidence in humans occurs predominantly on areas where soils are deficient in plant-available Zn, as shown in many Southeast Asian countries. India has some of the most Zn-deficient soils in the world. Nearly 50% of cultivated soils in India are low in plant-available Zn; these soils are under intensive cultivation of wheat and rice with no or little application of Zn fertilizers. Consequently, cereal crops grown on such Zn-deficient soils contribute only marginally to daily Zn intake. In the rural areas of India, rice and wheat contributes nearly 75% of the daily calorie intake. These facts clearly point to an urgent need for improved Zn concentration of cereal grains in India. Recent calculations indicate that biofortification (enrichment) of rice and wheat grain with Zn, for example by breeding, may save lives of up to 48,000 children in India annually. Breeding new cereal genotypes for high grain Zn concentration is the most realistic and cost-effective strategy to address the problem. However, this strategy is a long-term one, and the size of plant-available Zn pools in soils may greatly affect the capacity of Zn-efficient (biofortified) cultivars to take up Zn and accumulate it in grains. Therefore, application of Zn-containing fertilizers represents a quick and effective approach to biofortifying cereal grains with Zn, thus being an excellent complementary tool to the breeding strategy for successful biofortification of cereals with Zn. Increasing evidence is available from field trials showing that soil and/or foliar application of Zn fertilizers improves grain Zn concentration up to 2- or 3-fold. In the countries where Zn deficiency is both a public health issue and an important soil constraint to crop production, like in India, enrichment of widely applied fertilizers with Zn would be an excellent investment for improving grain Zn while contributing to increased crop production. Recent work by the scientists of the Indian Agricultural Research Institute indicates that the use of Zn-enriched urea in rice and wheat significantly improves both grain Zn concentration and grain yield. It is obvious that enrichment of widely applied fertilizers with Zn and/or foliar application of Zn fertilizers appear to be a high priority with the strongest potential to alleviate Zn deficiency-related problems in India. A Government action and policy plan for enrichment of selected major fertilizers with Zn is required urgently.     

Photosynthetic rate,chlorophyll fluorescence parameters,and lipid peroxidation of maize leaves as affected by zinc deficiency

Pot trial in greenhouse was conducted using cumulic cinnamon soil from North China to study the effects of zinc deficiency on CO₂ exchange, chlorophyll fluorescence, the intensity of lipid peroxidation, and the activity of superoxide dismutase (SOD) in leaves of maize seedlings. Zn deficiency resulted in a reduction of net photosynthetic rate and stomatal conductance to H₂O. The maximum quantum efficiency of photosystem 2 (PS2) and the PS2 activity were depressed, while the pool size of the plastoquinone molecules was not affected by Zn deficiency. The content of super oxygen anion radical (O₂ ^·−) and the intensity of lipid peroxidation as assessed by malonyldialdehyde content in Zn-deficient leaves were higher than those in Zn-sufficient leaves. The activity of SOD increased with Zn application. The adverse influence of Zn-deficiency on the light stage of photosynthesis is probably one of possible reasons for the limitation of photosynthetic capacity in maize leaves.     

Zinc deficiency induces enhanced depression-like behaviour and altered limbic activation reversed by antidepressant treatment in mice

A relationship between zinc (Zn)-deficiency and mood disorders has been suspected. Here we examined for the first time whether experimentally-induced Zn-deficiency in mice would alter depression- and anxiety-related behaviour assessed in established tests and whether these alterations would be sensitive to antidepressant treatment. Mice receiving a Zn-deficient diet (40% of daily requirement) had similar homecage and open field activity compared to normally fed mice, but displayed enhanced depression-like behaviour in both the forced swim and tail suspension tests which was reversed by chronic desipramine treatment. An anxiogenic effect of Zn-deficiency prevented by chronic desipramine and Hypericum perforatum treatment was observed in the novelty suppressed feeding test, but not in other anxiety tests performed. Zn-deficient mice showed exaggerated stress-evoked immediate-early gene expression in the amygdala which was normalised following DMI treatment. Taken together these data support the link between low Zn levels and depression-like behaviour and suggest experimentally-induced Zn deficiency as a putative model of depression in mice.     

The physiological mechanism of enhanced oxidizing capacity of rice (Oryza sativa L.) roots induced by phosphorus deficiency

Plants show various responses to phosphorus (P) deficiency. Root oxidizing capacity enhancement is one of adaptive mechanisms for rice (Oryza sativa L.) to P deficiency. However, it remains unclear how P deficiency enhances the root oxidizing capacity. In this study, rice seedlings were treated in P-deficient nutrient solution for different periods. Variations of reactive oxygen species (ROS), antioxidant enzyme activity, root lignin content, root porosity, root oxygen release, total oxidative substances and root structural changes in rice roots in response to P-sufficient and P-deficient treatments were investigated. Results indicated that P deficiency induced the production of H₂O₂ and O ₂ ^·? in roots significantly, which reached their maximum after 1- to 2-day P-deficient treatment. Interestingly, the endogenous total oxidative substances kept stable in rice roots. P deficiency increased the activities of peroxidase and superoxide dismutase by 89.5 and 51.8 % after 4-day P-deficient treatment, respectively. Moreover, one-day P deficiency elevated lignin accumulation. Root porosity of rice seedling under 2-day P-deficient treatment was 19.8 % higher than that under P-sufficient treatment. P deficiency also enhanced the release of both O₂ and total oxidative substances after 1- to 4-day P deficiency. In addition, results from electronic microscopy indicated that the thickness of root cell wall tended to increase after 2-day P-deficient treatment. Taken together, our results suggested that P-deficiency-induced enhancement of root oxidizing capacity in rice roots was probably associated with ROS production, antioxidant enzyme activity increment in root tissues, and the release of O₂ and oxidative substances from root inside to rhizosphere.