Yunxia Wang

The role of the root apoplast in aluminium-induced inhibition of root elongation and in aluminium resistance of plants: a review

BACKGROUND Aluminium (Al) toxicity is the most important soil constraint for plant growth and development in acid soils. The mechanism of Al-induced inhibition of root elongation is still not well understood, and it is a matter of debate whether the primary lesions of Al toxicity are apoplastic or symplastic. SCOPE The present review focuses on the role of the apoplast in Al toxicity and resistance, summarizing evidence from our own experimental work and other evidence published since 1995. CONCLUSIONS The binding of Al in the cell wall particularly to the pectic matrix and to the apoplastic face of the plasma membrane in the most Al-sensitive root zone of the root apex thus impairing apoplastic and symplastic cell functions is a major factor leading to Al-induced inhibition of root elongation. Although symplastic lesions of Al toxicity cannot be excluded, the protection of the root apoplast appears to be a prerequisite for Al resistance in both Al-tolerant and Al-accumulating plant species. In many plant species the release of organic acid anions complexing Al, thus protecting the root apoplast from Al binding, is a most important Al resistance mechanism. However, there is increasing physiological, biochemical and, most recently also, molecular evidence showing that the modification of the binding properties of the root apoplast contributes to Al resistance. A further in-depth characterization of the Al-induced apoplastic reaction in the most Al-sensitive zone of the root apex is urgently required, particularly to understand the Al resistance of the most Al-resistant plant species.

Biofortification and localization of zinc in wheat grain.

Zinc (Zn) deficiency associated with low dietary intake is a well-documented public health problem, resulting in serious health and socioeconomic problems. Field experiments were conducted with wheat to test the role of both soil and foliar application of ZnSO4 in Zn concentration of whole grain and grain fractions (e.g., bran, embryo and endosperm) in 3 locations. Foliar application of ZnSO4 was realized at different growth stages (e.g., stem elongation, boot, milk, dough stages) to study the effect of timing of foliar Zn application on grain Zn concentration. The rate of foliar Zn application at each growth stage was 4 kg of ZnSO4·7H2O ha(-1). Laser ablation (LA)-ICP-MS was used to follow the localization of Zn within grain. Soil Zn application at a rate of 50 kg of ZnSO4·7H2O ha(-1) was effective in increasing grain Zn concentration in the Zn-deficient location, but not in the locations without soil Zn deficiency. In all locations, foliar application of Zn significantly increased Zn concentration in whole grain and in each grain fraction, particularly in the case of high soil N fertilization. In Zn-deficient location, grain Zn concentration increased from 11 mg kg(-1) to 22 mg kg(-1) with foliar Zn application and to 27 mg kg(-1) with a combined application of ZnSO4 to soil and foliar. In locations without soil Zn deficiency, combination of high N application with two times foliar Zn application (e.g., at the booting and milk stages) increased grain Zn concentration, on average, from 28 mg kg(-1) to 58 mg kg(-1). Both ICP-OES and LA-ICP-MS data showed that the increase in Zn concentration of whole grain and grain fractions was pronounced when Zn was sprayed at the late growth stage (e.g., milk and dough). LA-ICP-MS data also indicated that Zn was transported into endosperm through the crease phloem. To our knowledge, this is the first study to show that the timing of foliar Zn application is of great importance in increasing grain Zn in wheat, especially in the endosperm part that is the predominant grain fraction consumed in many countries. Providing a large pool of Zn in vegetative tissues during the grain filling (e.g., via foliar Zn spray) is an important practice to increase grain Zn and contribute to human nutrition.

APOPLASTIC BINDING OF ALUMINUM IS INVOLVED IN SILICON-INDUCED AMELIORATION OF ALUMINUM TOXICITY IN MAIZE

The alleviating effect of silicon (Si) supply on aluminum (Al) toxicity was suggested to be based on ex or in planta mechanisms. In our experiments with the Al-sensitive maize (Zea mays) cultivar Lixis, Si treatment but not Si pretreatment ameliorated Al-induced root injury as revealed by less root-growth inhibition and callose formation. Si treatment did not affect monomeric Al concentrations in the nutrient solution, suggesting an in planta effect of Si on Al resistance. A fractionated analysis of Si and Al in the 1-cm root apices revealed that more than 85% of the root-tip Al was bound in the cell wall. Al contents in the apoplastic sap, the symplastic sap, and the cell wall did not differ between −Si and +Si plants. Si did not affect the Al-induced exudation of organic acid anions and phenols from the root apices. However, Al treatment greatly enhanced Si accumulation in the cell wall fraction, reducing the mobility of apoplastic Al. From our data we conclude that Si treatment leads to the formation of hydroxyaluminumsilicates in the apoplast of the root apex, thus detoxifying Al.

Stable isotope labelling and zinc distribution in grains studied by laser ablation ICP-MS in an ear culture system reveals zinc transport barriers during grain filling in wheat

Zinc (Zn) deficiency has been recognized as a potential risk for human health in many developing regions where staple food with low micronutrient density represents a major proportion of the diet. The success of strategies to increase Zn content in the edible part of crops requires better understanding of Zn transport to, and distribution within, the grains. The transfer of Zn from the growth medium to wheat (Triticum aestivum) grains in an ear culture system was investigated by using the stable Zn isotope (70) Zn, and the spatial distribution of Zn within the grains was studied by laser ablation-inductively coupled plasma-mass spectrometry (LA-ICP-MS). Zinc was readily transported in the stem up to the rachis. More Zn accumulated in the stem when higher amounts of Zn were supplied to the medium. Once Zn was transported into the grain, Zn accumulated particularly in the crease vascular tissue. The gradient of (70) Zn concentration between crease vascular tissue, aleurone layer and endosperm demonstrates that Zn is distributed within grain through the crease phloem. These results suggest that two barriers of Zn transport into wheat grains may exist: between the stem tissue rachis and the grain, and the maternal and filial tissues in the grain.

Molecular marker analysis of manganese toxicity tolerance in rice under greenhouse conditions

Two rice genotypes, an upland japonica rice Azucena and an indica rice IR1552, were found to differ in tolerance for Mn toxicity based the Mn toxicity symposium in the screening experiments in acid soil with higher available manganese level. Therefore, a recombinant inbred line (RIL) population in the F9 generation consisting of 150 lines derived from a cross between the two parents was used in soil pots experiment under greenhouse conditions using an acid soil with a high available manganese level of 25 mg Kg−1. Genotypic variation in tolerance for Mn toxicity was evaluated using Mn toxicity index (MTI) based on the degree of necrotic brown spots on the leaves and Mn concentration/content in shoots. Two hundred and seven markers covering 2419.5 cM on rice all 12 chromosomes were used by single marker loci and interval mapping. The results indicated that the two parents differed in MTI and Mn concentration and/or content in shoots, and these parameters were associated with Mn toxicity tolerance. The RILs showed a segregation for the above parameters. Molecular marker loci associated with variations in MIT values and Mn concentration/content in shoots were identified and eight QTLs associated with Mn toxicity tolerance were detected. Comparison of the marker genotype class means indicated that 6 of 8 tolerant alleles detected were contributed by the tolerant parent IR1552.

Biochemical factors conferring shoot tolerance to oxidative stress in rice grown in low zinc soil

Zinc deficiency reduces rice growth and yield, and this is, in part, due to leaf damage caused by reactive oxygen species (ROS). The aim of this study was to identify biochemical mechanisms conferring tolerance to Zn deficiency-induced oxidative stress. A field experiment and three nutrient solution experiments were conducted with the intolerant genotype IR74 and recombinant inbred lines (RILs) derived from a cross between IR74 and the tolerant landrace Jalmagna. After 2 weeks of growth in low Zn soil, stress symptoms developed in leaves of IR74, but not in the tolerant RIL46. Activity of antioxidant enzymes showed clear treatment effects, but did not explain tolerance of RIL46. On the contrary, the intolerant IR74 showed higher activities of superoxide dismutase (SOD), guaiacol peroxidase (POX), ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR) and glutathione reductase (GR) under Zn deficiency. This contrasted with a constitutively higher level of total and reduced ascorbic acid (AsA) in RIL46. Three further nutrient solution experiments focussed on enzymes and reducing substrates of the ascorbate–glutathione cycle. The first experiment included the highly sensitive RIL76 in addition to the genotypes used in the field trial, to test whether the patterns of antioxidant response observed in the field were specific to the genotypes used. This genotype had similarly low AsA level as IR74, but did not respond to Zn deficiency with an increase in enzyme activity, leading to even more pronounced leaf symptoms. In a second experiment, co-segregation of AsA concentration and Zn deficiency tolerance was confirmed in five genotypes from the IR74/Jalmagna quantitative trait loci (QTL) mapping population. A third experiment was conducted to determine whether the observed patterns of antioxidant response were specific to Zn deficiency or would also apply to oxidative stress caused by iron deficiency. Although high AsA level apparently conferred tolerance under both types of stress, the enzymatic response to iron deficiency differed from that to Zn deficiency. In particular, APX activity showed a decrease instead of an increase under low iron stress. In conclusion, we suggest that a high AsA level is a promising target for developing rice genotypes with tolerance to oxidative stress.

Significance of the Root Apoplast for Aluminium Toxicity and Resistance of Maize

The mechanism of aluminium-induced inhibition of root elongation is still not well understood. It is a matter of debate whether the primary lesions of Al toxicity are apoplastic or symplastic. The present paper summarises evidence from own experimental work and the literature which could contribute to the understanding of Al toxicity and resistance in maize focussing on the role of the apoplast.

Effects of elevated ozone, carbon dioxide, and the combination of both on the grain quality of Chinese hybrid rice

The effects of CO2 and/or O3 elevation on rice grain quality were investigated in chamber experiments with gas fumigation performed from transplanting until maturity in 2011 and 2012. Compared with the control (current CO2 and O3 concentration), elevated CO2 caused a tendency of an increase in grain chalkiness and a decrease in mineral nutrient concentrations. In contrast, elevated O3 significantly increased grain chalkiness and the concentrations of essential nutrients, while changes in starch pasting properties indicated a trend of deterioration in the cooking and eating quality. In the combination of elevated CO2 and O3 treatment, only chalkiness degree was significantly affected. It is concluded that the O3 concentration projected for the coming few decades will have more substantial effects on grain quality of Chinese hybrid rice than the projected high CO2 concentration alone, and the combination of two gases caused fewer significant changes in grain quality than individual gas treatments.

An agar nutrient solution technique as a screening tool for tolerance to zinc deficiency and iron toxicity in rice

Abstract Studies examining iron (Fe) toxicity and zinc (Zn) deficiency in rice have shown that screening experiments in nutrient solutions are of limited use because the rankings of genotypes as tolerant or intolerant can be very different from the results obtained in field-screening experiments. A possible reason for such deviation is that crucial rhizosphere processes cannot be reproduced in nutrient solutions. The objective of the present study was to evaluate the suitability of low-concentration agar nutrient solutions (ANS) as an alternative screening tool. Agar was dissolved in boiling water and mixed with nutrient solution to achieve a final agar concentration of 0.1% (w/v). Zinc deficiency was induced by supplying Zn at a low concentration (0.1 × 10−3 µmol L−1), while Fe toxicity was induced by supplying excess Fe2+ (200 mg L−1). Three-week-old seedlings were transplanted into this medium. Symptoms of Zn deficiency and Fe toxicity developed more rapidly in ANS compared with conventional nutrient solutions (CNS). For Zn deficiency this was probably because of the development of Zn depletion zones as a result of the reduced convection in the viscous agar medium. In the case of Fe toxicity we observed far less Fe precipitation in ANS compared with CNS. Genotypic comparisons showed that the tolerance rankings obtained in ANS were very similar to the field tolerance rankings, whereas this was not the case in CNS. This was particularly evident with regard to the considerable root growth inhibition detected in intolerant genotypes when stress treatments were imposed in ANS.

The impact of elevated CO2 and temperature on grain quality of rice grown under open‐air field conditions

BACKGROUND Rising atmospheric CO2 is accompanied by global warming. However, interactive effects of elevated CO2 and temperature have not been well studied on grain quality of rice. A japonica cultivar was grown in the field using a free-air CO2 enrichment facility in combination with a canopy air temperature increase system in 2014. The gas fumigation (200 µmol mol(-1) above ambient CO2 ) and temperature increase (1 °C above ambient air temperature) were performed from tillering until maturity. RESULTS Compared with the control (ambient CO2 and air temperature), elevated CO2 increased grain length and width as well as grain chalkiness but decreased protein concentrations. In contrast, the increase in canopy air temperature had less effect on these parameters except for grain chalkiness. The starch pasting properties of rice flour and taste analysis of cooked rice indicated that the palatability of rice was improved by CO2 and/or temperature elevation, with the combination of the two treatments showing the most significant changes compared with ambient rice. CONCLUSION It is concluded that projected CO2 in 2050 may have larger effects on rice grain quality than the projected temperature increase. Although deterioration in milling suitability, grain appearance and nutritional quality can be expected, the taste of cooked rice might be better in the future environment.