Biofortification strategies to increase grain zinc and iron concentrations in wheat

Micronutrient deficiencies, especially those arising from zinc (Zn) and iron (Fe), pose serious human health problems for more than 2 billion people worldwide. Wheat is a major source of dietary energy and protein for the world's growing population, and its potential to assist in reducing micronutrient-related malnutrition can be enhanced via integration of agronomic fertilization practices and delivery of genetically-manipulated, micronutrient rich wheat varieties. Targeted breeding for these biofortified varieties was initiated by exploiting available genetic diversity for Zn and Fe from wild relatives of cultivated wheat and synthetic hexaploid progenitors. The proof-of-concept results from the performance of competitive biofortified wheat lines showed good adaptation in target environments without compromising essential core agronomic traits. Agronomic biofortification through fertilizer approaches could complement the existing breeding approach; for instance, foliar application of Zn fertilizer can increase grain Zn above the breeding target set by nutritionists. This review synthesizes the progress made in genetic and agronomic biofortification strategies for Zn and Fe enrichment of wheat.     

Improved nitrogen status enhances zinc and iron concentrations both in the whole grain and the endosperm fraction of wheat

This study was conducted to assess the role of increasing N supply in enrichment of whole grain and grain fractions, particularly the endosperm, with Zn and Fe in wheat. The endosperm is the most widely consumed part of wheat grain in many countries. Plants were grown in the greenhouse with different soil applications of N and Zn and with or without foliar Zn spray. Whole grain and grain fractions were analyzed for N, P, Zn and Fe. Increased N supply significantly enhanced the Zn and Fe concentrations in all grain fractions. In the case of high Zn supply, increasing N application enhanced the whole grain Zn concentration by up to 50% and the endosperm Zn by over 80%. Depending on foliar Zn supply, high N elevated the endosperm Fe concentration up to 100%. High N also generally decreased the P/Zn and P/Fe molar ratios in whole grain and endosperm. The results demonstrate that improved N nutrition, especially when combined with foliar Zn treatment, is effective in increasing Zn and Fe of the whole grain and particularly the endosperm fraction, at least in the greenhouse, and might be a promising strategy for tackling micronutrient deficiencies in countries where white flour is extensively consumed.     

Variation in mineral micronutrient concentrations in grain of wheat lines of diverse origin

150 lines of bread wheat representing diverse origin and 25 lines of durum, spelt, einkorn and emmer wheat species were analysed for variation in micronutrient concentrations in grain. A subset of 26 bread wheat lines was grown at six sites or seasons to identify genetically determined differences in micronutrient concentrations. Substantial variation among the 175 lines existed in grain Fe, Zn and Se concentrations. Spelt, einkorn and emmer wheats appeared to contain higher Se concentration in grain than bread and durum wheats. Significant differences between bread wheat genotypes were found for grain Fe and Zn, but not Se concentration; the latter was influenced more by the soil supply. Grain Zn, but not Fe, concentration correlated negatively with grain yield, and there was a significant decreasing trend in grain Zn concentration with the date of variety release, suggesting that genetic improvement in yield has resulted in a dilution of Zn concentration in grain. Both grain Zn and Fe concentrations also correlated positively and significantly with grain protein content and P concentration, but the correlations with kernel size, kernel weight or bran yield were weak. The results from this study are useful for developing micronutrient biofortification strategies.     

Wheat ferritins: Improving the iron content of the wheat grain

The characterization of the full complement of wheat ferritins show that the modern hexaploid wheat genome contains two ferritin genes, TaFer1 and TaFer2, each represented by three homeoalleles and placed on chromosome 5 and 4, respectively. The two genes are differentially regulated and expressed. The TaFer1 genes are, except in the endosperm, the most abundantly expressed and regulated by iron and abscisic acid status. The promoter of TaFer1, in contrast to TaFer2, has iron- and ABA-responsive elements, supporting the expression data. The TaFer1 and TaFer2 genes encode two isoforms, probably functional different and acting in heteropolymer structures of ferritin in cereals. Iron biofortification of the wheat grain is possible. Endosperm targeted intragenic overexpressing of the TaFer1-A gene results in a 50–85% higher iron content in the grain.     

Phytase activity, phytate, iron, and zinc contents in wheat pearling fractions and their variation across production locations

Pearling is an effective method for evaluating the distribution of chemical components in wheat grain. Twelve pearling fractions (P₁–P₁₂) of wheat grain were obtained using two rice polishers for 10 cultivars (six soft red wheats and four hard white wheats) grown at two locations with different environmental conditions in Jiangsu Province, China. The results show that the effects of cultivar, location, and pearling on wheat flour phytase activity, phytate, iron, and zinc contents were all significant, with pearling having the greatest effect. All the four components showed a diminishing trend as pearling progressed from the outer layers to the inner part of wheat grain. Generally, the P₂ fraction (the outer 4–8% layer of wheat grain) had the highest phytase activity and phytate and iron contents, whereas the P₁ fraction (the outer 0–4% layer) ranked the highest for zinc content. Growing location had a large influence on grain phytase, phytate, and iron, but the differences between locations decreased as pearling level increased.     

Genetic variation of bioavailable iron and zinc in grain of a maize population

More than one-third of the world's population is afflicted by iron (Fe) and zinc (Zn) deficiencies, since cereal grain as a staple food of the people contains low levels or low bioavailability of Fe and Zn because of phytate. In maize, 80% of grain phosphorus (P) is in the form of phytate, and P could be an indicator of phytate content. The objectives of this study were (1) to estimate genetic variation of Fe and Zn in a maize population including P/Fe and P/Zn molar ratios as quantitative traits; (2) to determine relations among yield, P, Fe, Zn, P/Fe and P/Zn molar ratios; and (3) to define the implications of those on biofortification (breeding) programmes. There were significant genetic variations and workable heritabilities for Fe, Zn, P/Fe and P/Zn estimated in 294 F4 lines of a maize population, but there were no associations among six traits according to both simple correlations and principal component analysis. Weak correlations between P and Fe and between P and Zn indicated feasibility of breeding non low-phytic acid maize genotypes with more appropriate phytate/Fe and phytate/Zn relations. Bioavailability of iron and zinc varied substantially in a maize population justifying utilisation of these unique parameters in biofortification programmes.     

Identification of CIMMYT spring bread wheat germplasm maintaining superior grain yield and quality under heat-stress

Unpredictable temperatures and rainfall associated with climate change are expected to affect wheat (Triticum aestivum L.) production in various countries. The development of climate-resilient spring wheat cultivars able to maintain grain yield and quality is essential to food security and economic returns. We tested 54 CIMMYT spring bread wheat genotypes, developed and/or released over a span of 50 years, in the field for two years under optimum sowing dates, as well as using two delayed sowing dates to expose crops to medium and severe heat-stress conditions. The grain yield and yield components were severely affected as the heat-stress increased. Two contrasting groups of 10 lines each were identified to determine the effect of heat-stress on bread-making quality. The first set included entries that produced high yields in optimal conditions and maintained higher yields under heat-stress (superior-yielding lines), while the second set included genotypes that did not perform well in the environment with high temperature (inferior-yielding lines). We identified genotypes exhibiting bread-making quality stability, as well as the quality traits that had higher correlation with the loaf volume in the environment without stress and under heat-stress. Of all the quality traits tested, dough extensibility (AlvL) and grain protein content had a significant influence in heat-stress adaptation. Most of the lines from the superior-yielding group were also able to maintain and even improve quality characteristics under heat-stress and therefore, could be used as parents in breeding to develop high-yielding and stable quality wheat varieties.     

Localisation of iron in wheat grain using high resolution secondary ion mass spectrometry

Insufficient iron (Fe) is one of the most prevalent micronutrient deficiencies in humans, with billions of people affected. Cereal grains are an important source of Fe for humans but the bioavailability of Fe in cereals is generally low. Information regarding the cellular and sub-cellular localisation of Fe in wheat grain will aid optimising nutrient delivery for human health. In this study high resolution secondary ion mass spectrometry (NanoSIMS) was used to map the distribution of Fe in the aleurone layer and in the endosperm of immature wheat grain. Iron was shown to be localised strongly in the phytin globoids in the aleurone cells and to a lesser extent in the cytoplasm around the starch granules in the endosperm.     

Effect of temperature variation during grain filling on wheat gluten resistance

The objective was to investigate effects of natural variation in temperature during grain filling on wheat (Triticum aestivum L) gluten quality. Seventeen field trials with four different varieties were conducted during the years 2005–2008. Temperature records were obtained from automatic weather stations located near the field trial sites. The period from heading to yellow ripeness was divided into 20 sub-phases of equal thermal time units, and a last sub-phase comprising the seven days after yellow ripeness. Partial Least Squares Regression was used to relate the temperature records of the different sub-phases to gluten quality analysed by the Kieffer Extensibility test and the SDS sedimentation test.     

Yield dilution of grain Zn in wheat grown in open-top chamber experiments with elevated CO2 and O3 exposure

Wheat (Triticum aestivum L.) grain Zn data from six open-top chamber experiments performed in south-west Sweden were combined to study the relationship between Zn accumulation and grain yield, grain protein, and yield components. Treatments included, in addition to open-top chamber controls, elevated CO₂, elevated O₃, combined CO₂ and O₃ exposure, combined elevated CO₂ and supplemental irrigation, supplemental irrigation, and ambient air comparison plots. The grain Zn concentration was strongly correlated with grain protein (R² = 0.90) over the range of the experimental treatments, representing non-soil factors. A significant yield dilution effect was found for Zn. For a 10% increase in grain yield, Zn yield was increased by 6.8% on average. Effects on Zn yield correlated strongly with effects on grain protein yield, with a slope close to unity, showing that yield dilution effects for grain Zn and grain protein were similar. Treatment effects on grain Zn concentration were related to effects on grain weight (P < 0.01) and grain number (P < 0.05), but not to harvest index. It was concluded that yield stimulation caused by rising CO₂ concentrations is likely to lead to reduced Zn concentrations of wheat grain, thus reducing the nutritional quality.     

Biochemical and morphological risk assessment of transgenic wheat with enhanced iron and zinc bioaccessibility

Decreased iron and zinc bioaccessibility, caused by the anti-nutrient phytic acid, is one of the leading reasons for micronutrient deficiency-related disorders. Biofortification of wheat with phytase gene to enhance iron and zinc bioaccessibility appears to be a fitting solution for this problem, especially in developing countries where most of the population belongs to the lower economic sector. However, societal views on crops, particularly crops that are genetically modified (GM) to express a new trait, needs to be changed. Risk assessment of GM crops can play a crucial role in fostering positive public perception, since it is imperative to ensure safety before allowing human consumption. The present study performed compositional and morphological risk assessment of T3 and T4 generations of phytase transgenic wheat by comparing their biochemical and morphological traits. Transgenic plants were analysed for their carbohydrate, protein, starch and phytic acid content along with iron bioaccessibility, zinc bioaccessibility and phytase enzyme activity. Morphological traits studied included plant height, seed number, seed weight and spike number. No significant differences were observed for carbohydrate, protein, starch content and for morphological traits; however, a significant increase was observed in phytase activity as well as iron and zinc bioaccessibility, which correlated with a significant decrease in phytic acid. These results demonstrate that phytase transgenic wheat is as native as local wheat varieties and can potentially increase iron and zinc bioaccessibility.     

小麦铁锌营养品质研究进展

"隐性饥饿",即铁、锌等微量元素缺乏症,已成为困扰我国居民的首要营养不良问题.选育高铁、锌含量、强植酸酶活性或低植酸含量的"微量营养强化型"小麦品种对于改善我国西部居民的营养状况具有重要的意义.本文主要介绍小麦铁锌营养品质的遗传改良研究进展,包括铁锌营养品质的相关化学组分(铁、锌、植酸和植酸酶)及改善小麦铁锌营养品质的遗传途径等两方面,并指出了小麦铁锌营养品质的研究方向和工作重点.     

Generation mean analysis of pearl millet [Pennisetum glaucum (L.) R. Br.] grain iron and zinc contents and agronomic traits in West Africa

Pearl millet is the most important staple food crop for millions of people across the world. Micronutrient malnutrition is the major problem for people living in the semi-arid regions of Africa. Identification of gene effects controlling the inheritance of grain Fe and Zn will be helpful in formulating suitable breeding strategies for biofortified pearl millet development. Hence, generation mean analysis was used to study epistasis and estimate gene effects for grain iron and zinc contents along with the agronomic and morphological traits. Six generations P₁, P₂, F₁, F₂, BC₁P₁ and BC₁P₂ were generated and were evaluated during the 2018–19 off season. Analysis of variance showed significant variability for all the traits in both generations. Six parameter model revealed predominance of additive gene effects for inheritance of grain iron concentration, and additive × additive type of non-allelic interactions. For grain zinc concentration additive gene effects were preponderant compared to non-additive gene effects, and only additive × dominance gene effects were significant among the three types of epistasis. Grain weight per plant was predominantly under non-additive gene effects and additive × additive and additive × dominance gene effects type of epistasis was detected in each cross. Likewise, for flowering non-additive gene effects were most important with the presence of dominance × dominance type of epistasis. For plant height, panicle circumference and length, additive × additive genes effects were the most important among the three type of non-allelic gene action.These findings can be helpful in enehancing the pearl millet breeding programs in Africa.     

Effect of water deficits on within-plot variability in growth and grain yield of spring wheat in northwest China

For adapted cultivars under normal crop densities, biological yield is largely determined by the pool of available resources, e.g. water, nutrients and photosynthetically active radiation, while the nature and intensity of intraspecific competition plays an important role in determining the magnitude of harvest index (HI). Water deficits can drastically reduce the HI from its genetic potential to zero. This study was conducted to determine the effect of drought-weighted intraspecific competition on the HI and, consequently, the grain yield in spring wheat populations along a natural moisture gradient in northwestern China. Along the natural moisture gradient (annual mean rainfall decreased 328→204→185 mm per year, supplemented with 70 mm of irrigation), culm size inequality (as measured by the Gini coefficient of above-ground biomass per culm) always increased, and Lorenz curves were more concave. HI decreased significantly in 1999 (0.364→0.345→0.307) and 2000 (0.341→0.303→0.251). There was a significant negative correlation between the Gini coefficient and the HI of spring wheat along the moisture gradient (R²=0.92, P<0.01). These results suggested that size hierarchies in spring wheat populations are closely correlated with the water regime in the field, and that under greater drought stress there are relatively more smaller plants with lower HI (size-dependent reproductive allocation). Size inequality is an index of competitive status in plant populations under stress environments. Agriculturally, greater size inequality may result in a competitive cost for energy and photosynthetic products, in other words, growth redundancy, which is detrimental to reproductive allocation and consequently, grain yield. The results support the view that stand uniformity in field crops is an important mechanism for increasing grain yield.     

Dietary strategies to improve the iron and zinc nutriture of young women following a vegetarian diet

Dietary strategies to enhance the content and bioavailability of iron and zinc in vegetarian diets were compiled. Strategies included increasing promoters and decreasing antagonists of iron and zinc absorption, adopting food preparation and processing methods which hydrolyze the phytate content of cereals and legumes, and using iron cookware. These strategies were used to devise two vegetarian menus based on food consumption patterns derived from three day weighed food records of 78 Canadian lacto-ovo-vegetarian adolescents. The iron and zinc, as well as calcium, phosphorus, thiamin, riboflavin, niacin, vitamin A, protein and energy content of the devised menus were all higher than the actual intakes and the corresponding Canadian recommended nutrient intakes. Results show the overall nutrient adequacy of the recommended vegetarian menus and indicate that young lacto-vegetarian women can potentially meet their estimated dietary requirements for absorbed iron and zinc through modest modifications to the diet. Laboratory studies designed to measure the total amount of iron and zinc absorbed from these diets by young vegetarian women are needed to verify the efficacy of the devised menus.     

Effects of elevated atmospheric CO2 on grain quality of wheat

Wheat (Triticum aestivum L.) is one of the most important agricultural crops worldwide. Due to its high content of starch and unique gluten proteins, wheat grain is used for many food and non-food applications. Although grain quality is an important topic for food and feed as well as industrial processing, the consequences of future increases in atmospheric carbon dioxide (CO₂) concentrations on quality parameters such as nutritional and bread-making rheological properties are still unclear. Wheat productivity increases under CO₂ enrichment. Concomitantly, the chemical composition of vegetative plant parts is often changed and grain quality is altered. In particular, the decrease in grain protein concentration and changes in protein composition may have serious economic and health implications. Additionally, CO₂ enrichment affects amino acid composition and the concentrations of macro- and micro-elements. However, experimental results are often inconsistent. The present review summarises the results from numerous CO₂ enrichment experiments using different exposure techniques in order to quantify the potential impacts of projected atmospheric CO₂ levels on wheat grain yield and on aspects of grain composition relevant to processing and human nutrition.     

Influence of vegetative cycle of asparagus (Asparagus officinalis L.) on copper,iron, zinc and manganese content

The essential elements copper (Cu), iron (Fe), zinc (Zn) and manganese (Mn) were analyzed in fresh asparagus to determine the effects of the vegetative cycle of the plant on the micronutrient content. Asparagus samples were classified in two groups by diameter (<11 mm and >14 mm). Asparagus from a sample group with the same diameter were divided into two portions (apical and basal) according to distance from the tip. The concentrations of copper, iron, zinc and maganese increased during the vegetative cycle of the asparagus, mainly in the apical portion which showed significantly greater concentrations with respect to the basal portion. The >14 mm diameter asparagus presented higher levels of copper, zinc and manganese, whereas the concentration of iron was greater in the <11 mm diameter asparagus. The mean element levels were (mg/kg dry weight): Cu, 18.9±3.9; Fe, 91.7±33.7; Zn, 69.5±24.6 and Mn, 20.9±5.0).