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.     

小麦籽粒微量元素含量的研究进展

以禾谷类作物为主食引起的人体摄取必需微量元素(特别是Fe和Zn)不足,已对现代社会和谐发展造成沉重的经济负担和安全隐患。小麦(Triticum aestivumL.)是中国和全球大多数人主要的食物和矿质元素来源。小麦籽粒中Zn、Fe含量普遍较低,已引起国内外学者们的高度关注。本文综述了小麦籽粒微量元素含量器官间、基因型间的差异及其影响因素和相关性状研究进展,介绍了小麦籽粒吸收和富集微量元素的生理与遗传基础,展望了提高小麦籽粒微量元素的研究内容和方向。     

Enhancing the mineral and vitamin content of wheat and maize through plant breeding

More than half of the world's population suffers micronutrient undernourishment. The main sources of vitamins and minerals (iron, zinc, and vitamin A) for low-income rural and urban populations are staple foods of plant origin that often contain low levels or low bioavailability of these micronutrients. Biofortification aims to develop micronutrient-enhanced crop varieties through conventional plant breeding. HarvestPlus, the CGIAR's biofortification initiative, seeks to breed and disseminate crop varieties with enhanced micronutrient content that can improve the nutrition of the “hard to reach” (by fortification or supplementation programmes) rural and urban poor in targeted countries/regions. In attempting to enhance micronutrient levels in maize and wheat through conventional plant breeding, it is important to identify genetic resources with high levels of the targeted micronutrients, to consider the heritability of the targeted traits, to explore the availability of high throughput screening tools and to gain a better understanding of genotype by environment interactions. Biofortified maize and wheat varieties must have the trait combinations which encourage adoption such as high yield potential, disease resistance, and consumer acceptability. When defining breeding strategies and targeting micronutrient levels, researchers need to consider the desired micronutrient increases, food intake and retention and bioavailability as they relate to food processing, anti-nutritional factors and promoters. Finally, ex ante studies are required to quantify the burden of micronutrient deficiency and the potential of biofortification to achieve a significant improvement in human micronutrient status in the deficient target population in order to determine whether a biofortification program is cost-effective.     

Enhancing the mineral and vitamin content of wheat and maize through plant breeding

More than half of the world's population suffers micronutrient undernourishment. The main sources of vitamins and minerals (iron, zinc, and vitamin A) for low-income rural and urban populations are staple foods of plant origin that often contain low levels or low bioavailability of these micronutrients. Biofortification aims to develop micronutrient-enhanced crop varieties through conventional plant breeding. HarvestPlus, the CGIAR's biofortification initiative, seeks to breed and disseminate crop varieties with enhanced micronutrient content that can improve the nutrition of the “hard to reach” (by fortification or supplementation programmes) rural and urban poor in targeted countries/regions. In attempting to enhance micronutrient levels in maize and wheat through conventional plant breeding, it is important to identify genetic resources with high levels of the targeted micronutrients, to consider the heritability of the targeted traits, to explore the availability of high throughput screening tools and to gain a better understanding of genotype by environment interactions. Biofortified maize and wheat varieties must have the trait combinations which encourage adoption such as high yield potential, disease resistance, and consumer acceptability. When defining breeding strategies and targeting micronutrient levels, researchers need to consider the desired micronutrient increases, food intake and retention and bioavailability as they relate to food processing, anti-nutritional factors and promoters. Finally, ex ante studies are required to quantify the burden of micronutrient deficiency and the potential of biofortification to achieve a significant improvement in human micronutrient status in the deficient target population in order to determine whether a biofortification program is cost-effective.     

Genetic diversity for grain nutrients contents in a core collection of finger millet (Eleusine coracana (L.) Gaertn.) germplasm

Finger millet is a promising source of micronutrients and protein besides energy and can contribute to the alleviation of iron (Fe), zinc (Zn) and protein malnutrition affecting women and preschool children in African and south-east Asian countries. The most cost effective approach for mitigating micronutrient and protein malnutrition is to introduce staple crop cultivars selected and/or bred for Fe, Zn and protein dense grain. Breeding finger millet for enhanced grain nutrients is still in its infancy. Analysis, detection and exploitation of the existing variability among the germplasm accessions are the initial steps in breeding micronutrient and protein-dense finger millet cultivars. Evaluation of finger millet core collection for grain nutrients and agronomic traits revealed a substantial genetic variability for grain Fe, Zn, calcium (Ca) and protein contents. The accessions rich in nutrient contents were identified and their agronomic diversity assessed. The accessions rich in Zn content have significantly higher grain yield potential than those rich in Fe and protein content. Grain nutrient-specific accessions and those contrasting for nutrient contents were identified for use in the strategic research and cultivar development in finger millet.     

氮肥对不同玉米品种子粒微量元素含量和生物有效性的影响

研究田间施用氮肥对吉林省4个玉米品种子粒中Zn、Fe、Mn、Cu含量以及Zn与Fe生物有效性的影响。结果表明,施用氮肥可增加玉米子粒中Zn、Fe含量,过量施氮玉米子粒中Zn和Fe含量不再增加甚至呈下降趋势;子粒中Mn含量随氮肥增加而提高,Cu含量随氮肥增加而降低,4个品种中只有吉四单19在过量施氮时子粒Mn含量呈下降趋势。随氮肥水平提高,玉米子粒中Zn、Fe和Mn累积量呈递增趋势,Cu累积量变化不显著。全磷与这些矿物质元素的比值(P/Zn、P/Fe)表明,氮肥施用将显著降低玉米子粒中Zn和Fe的生物有效性。     

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.     

小麦籽粒矿质元素的基因型差异及对锌强化的响应

为了分析锌强化对不同小麦品种籽粒矿质元素含量的影响,以40个不同小麦品种(系)为材料,在扬花期叶面喷施0.5%的Zn肥,用原子吸收光谱法测定Zn处理与不处理(对照)下不同小麦品种籽粒中6种矿质元素(Zn、Fe、Ca、Mg、Cu、Mn)的含量,并分析这些元素对Zn强化的响应。结果表明,Zn处理后各品种籽粒中Zn含量均比对照有极显著上升,但上升幅度因品种而异。籽粒中Fe、Ca、Mg、Cu、Mn含量对Zn强化的响应因元素种类和品种而异,就品种平均值而言,Mn含量在Zn处理和对照间有极显著差异,Fe、Mg和Cu含量在处理与对照间无显著差异。大部分品种Zn处理后籽粒Ca含量有所下降,Mn则有所增加。对施Zn和对照间被测矿质元素的变幅进行主成分分析,前三个主成分(PC1、PC2、PC3)累积贡献率为0.76,基于前三个主成分对40个品种(系)进行类平均法的聚类分析,将40个品种分为3类:第1类包含有8个品种,Zn处理后籽粒中Ca、Mg、Cu、Fe含量总体呈现降低趋势,而Mn、Zn含量呈增高趋势;第2类包含23个品种,Zn处理后籽粒Ca、Cu含量总体有一定的降低,其余4种元素含量则有一定幅度的提高;第3类共有9个品种,施Zn后Fe含量总体有一定幅度的降低,其余5种元素含量则有一定幅度的增加。来自第2类和第3类中的12个品种在Zn处理后所测定的全部6种元素或者其中大部分元素的含量提高,适宜进行Zn强化,而其余品种Zn强化后部分或全部元素(Zn除外)与对照相比有所下降,不宜作为Zn营养强化的载体品种。     

Hydrocyanate,oxalate, phytate,calcium and zinc in selected brands of Nigerian cocoa beverage

Hydrocyanate, oxalate, phytate, calcium and zinc were determined in five brands of cocoa beverage which were coded NC, BT, PN, CT and CA. Hydrocyanate ranged from 5.40 to 9.64 mg/100 g dry matter (DM), oxalate 68 to 146 mg/100 g DM, phytate 590 to 750 mg/100 g DM, calcium 28.7 to 116.4 mg/100 g DM and zinc 0.516 to 0.675 mg/100 g DM. The computed phytate:zinc, calcium:phytate and [calcium] [phytate]/[zinc] molar ratios ranged from 89 to 132, 0.80 to 3.01 and 0.64 to 3.03 respectively. The discussion is focused on toxic levels of hydrocyanate and oxalate, and the significance of the molar ratios in predicting the bioavailability of dietary zinc.     

Phosphorus application affects the nutritional quality of millet grain in the Sahel

The application of phosphorus (P) and crop residues (CR) to acid sandy soils of the Sahel has been shown to increase yields of pearl millet (Pennisetum glaucum L.) several-fold. Information is lacking, however, about possible detrimental effects of such yield-enhancing amendments on grain quality, in particular the bioavailability of zinc (Zn) as defined by the phytate:zinc molar ratio (PZMR) and the concentrations of calcium, micronutrients, and protein. To determine the effects of CR and P on grain quality, millet seeds taken from the grain stores of 14 farmers and from a 2-yr on-station fertilizer experiment were analyzed for macronutrients, Zn, copper, iron, and phytate-P. The on-station experiment comprised four millet lines, P applied at 0 and 13 kg ha⁻¹, and CR applied at 500 and 2000 kg ha⁻¹ as surface mulch or ash. Grain from farmers' unfertilized millet had PZMRs ranging from 15 to 30. Application of P increased the concentrations of phytic acid in the grain between 25 and 29% and decreased Zn concentrations between 6 and 11%. The reasons for this were greater P uptake and a dilution of Zn by the large yield increases after P application. Phosphorus application decreased protein concentrations in both years, and increased the PZMRs from 20 to 28 in 1992 and from 21 to 29 in 1993. Although CR markedly increased millet yield, their application had little effect on PZMRs. While PZMRs above 15 are generally considered critical for Zn nutrition of humans, meat consumption and traditional practices of millet processing may increase Zn bioavailability in local dishes. Further studies of full diets are therefore needed, particularly among rural groups at particularly high risk of Zn deficiency such as nursing women and small children before definitive conclusions can be drawn about the effects of P fertilizer application to millet on the nutritional status of farmers in the Sahel.     

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.     

Breeding for increased grain protein and micronutrient content in wheat: Ten years of the GPC-B1 gene

To provide food and nutrition security for a growing world population, continued improvements in the yield and nutritional quality of agricultural crops will be required. Wheat is an important source of calories, protein and micronutrients and is thus a priority to breed for improvements in these traits. The GRAIN PROTEIN CONTENT-B1 (GPC-B1) gene is a positive regulator of nutrient translocation which increases protein, iron and zinc concentration in the wheat grain. In the ten years since it was cloned, the impacts of GPC-B1 allelic variation on quality and yield traits have been extensively analyzed in diverse genetic backgrounds in field studies spanning forty environments and seven countries. In this review, we compile data from twenty-five studies to summarize the impact of GPC-B1 allelic variation on fifty different traits. Taken together, the results demonstrate that the functional copy of the GPC-B1 gene is associated with consistent positive effects on grain protein, Fe and Zn content with only marginally negative impacts on yield. We conclude that the GPC-B1 gene has the potential to increase nutritional and end use quality in a wide range of modern cultivars and environments and discuss the possibilities for its application in wheat breeding.     

Correlation of Plant Morphological and Grain Quality Traits with Mineral Element Contents in Yunnan Rice

Correlations between four grain quality, 24 plant morphological traits and eight element contents of 653 accessions from Yunnan rice were analyzed. P, K, Ca and Mg contents of core collection were closely correlated to the most plant morphological and grain quality traits, and there were highly significant correlations （P 〈0.01） among some traits including P content to number of stems and tillers, K content and amylose content, Ca content and plant height, Mg content and protein content. Mn, Zn, Cu and Fe contents of core collection were closely related to a few traits, such as Fe content and gel consistency （-0.1121^＊＊）, Zn content and seed setting rate （-0.1411^＊＊）, Cu content and number of grains per panicle （-0.1398^＊＊）, Mn content and plant height （-0.2492^＊＊）.     

Molecular genetic approaches to increasing mineral availability and vitamin content of cereals

The present paper summarizes the current state of knowledge on molecular genetic approaches to increasing iron and zinc availability and vitamin content in cereals. We have also attempted to integrate the scientific issues into the wider context of human nutrition. In the cereal grain, iron and zinc are preferentially stored together with phytate in membrane-enclosed globoids in the protein storage vacuole (PSV) found in the aleurone and the embryo scutellum. The PSV is accordingly central for understanding mineral deposition during grain filling and mobilization of minerals during germination. Recent studies in Arabidopsis have led to the first identification of iron and zinc transporters of the PSV and further illustrate some of the dynamics associated with mineral and phytate transport and deposition into the vacuole. This provides new opportunities for modulating iron and zinc deposition in the cereal grain. Current strategies towards increasing the iron content of the endosperm are largely based on the expression of legume ferritin genes in an endosperm-specific manner. However, it is apparent that this approach, at least in rice, only allows a two- to three-fold increase in the iron content of the grain due to exhaustion of the iron stores in leaves. Further increases thus have to rely on additional uptake and transport of iron from the root. Phytate is generally considered to be the single most important anti-nutritional factor for iron and zinc availability. In the current paper we summarize attempts to increase phytase activity in the grain by transformation and evaluate the potential of this approach as well as the reduction of phytate biosynthesis for improving the bioavailability of iron and zinc. Vitamins constitute the second important group of micronutrients in grain and we discuss current efforts to increase the amounts of provitamin A, vitamin C and vitamin E.     

Effects of Different Nitrogen Fertilizer Levels and Native Soil Properties on Rice Grain Fe, Zn and Protein Contents

Deposition of protein and metal ions (Fe, Zn) in rice grains is a complex polygenic trait showing considerable environmental effect. To analyze the effect of nitrogen application levels and native soil properties on rice grain protein, iron (Fe) and zinc (Zn) contents, 32 rice genotypes were grown at three different locations each under 80 and 120 kg/hm2 nitrogen fertilizer applications. In treatments with nitrogen fertilizer application, the brown rice grain protein content (GPC) increased significantly (1.1% to 7.0%) under higher nitrogen fertilizer application (120 kg/hm2) whereas grain Fe/Zn contents showed non-significant effect of nitrogen application level, thus suggesting that the rate of uptake and translocation of macro-elements does not influence the uptake and translocation of micro-elements. The pH, organic matter content and inherent Fe/Zn levels of native soil showed significant effects on grain Fe and Zn contents of all the rice genotypes. Grain Zn content of almost all the tested rice genotypes was found to increase at Location III having loamy soil texture, neutral pH value (pH 6.83) and higher organic matter content than the other two locations (Locations I and II), indicating significant influence of native soil properties on brown rice grain Zn content while grain Fe content showed significant genotype × environment interaction effect. Genotypic difference was found to be the most significant factor to affect grain Fe/Zn contents in all the tested rice genotypes, indicating that although native soil properties influence phyto-availability of micronutrients and consequently influencing absorption, translocation and grain deposition of Fe/Zn ions, yet genetic makeup of a plant determines its response to varied soil conditions and other external factors. Two indica rice genotypes R-RF-31 (27.62 μg/g grain Zn content and 7.80% GPC) and R1033-968-2-1 (30.05 μg/g grain Zn content and 8.47% GPC) were identified as high grain Zn and moderate GPC rice genotypes. These results indicate that soil property and organic matter content increase the availability of Fe and Zn in rhizosphere, which in turn enhances the uptake, translocation and redistribution of Fe/Zn into rice grains.     

In vitro availability of iron and zinc in white and coloured ragi (Eleusine coracana): role of tannin and phytate

White and brown ragi (Eleusine coracana) varieties were analysed for tannin, phytate phosphorus, total phosphorus, iron, ionisable iron, zinc and soluble zinc content. White ragi had no detectable tannin while in brown varieties it ranged from 351 to 2392 mg per 100 g. Germination brought about a progressive decrease in tannin and phytate phosphorus and an increase in ionisable ion and soluble zinc content of grain ragi. Both in raw and germinated grain, ionizable iron was significantly higher in white than in brown varieties. While ionisable iron was inversely correlated with the level of tannin and phytate phosphorus, soluble zinc was negatively correlated with phytate phosphorus. After extraction of tannin, ionisable iron of brown ragi rose by 85%. On the other hand, in white varieties, addition of tannin extracted from brown ragi, resulted in a 52–65% decrease in ionisable iron content. These studies indicated that poor iron availability in ragi as judged by its low ionisable iron content was due to the presence of tannin in the grain.     

The influence of milling on the nutritive value of flour from cereal grains. 5. Maize

Different grades of maize flour were produced by dry milling of yellow maize. The chemical composition of wholemeal, three semi-sifted flours and degermed maize was determined, and the nutritional value of the flours were investigated in balance experiments with growing rats. Distinct differences in nutrient and fibre content were found. Amino acid composition was significantly different in the milled fractions, and the content of minerals and phytate in the flours was determined by the extent to which the germ was retained. Phytate: zinc molar ratios ranged from 43 to 2. Weight gain of rats was closely correlated to the concentration of lysine (r=0.99) and tryptophan (r=0.97). Weight gain of rats fed degermed maize was reduced to half of that of rats fed wholemeal. True protein digestibility was high, but protein quality varied widely between the different flour fractions. Energy density of the flours was quite similar. Rats fed wholemeal had the highest intake of zinc and the highest apparent zinc absorption and retention but the lowest femur zinc concentration. Factors present in whole maize may interfere with zinc utilization. The results might indicate an interaction between phytate and fibre. Supplementation of wholemeal with lysine and tryptophan increased the biological value of the protein and the weight gain considerably, but apparent absorption and retention of zinc as well as femur zinc concentration were similar in unsupplemented and supplemented rats. The percentage of zinc absorbed from degermed maize flour was high, but degermed maize has a very low content of minerals and is highly deficient in lysine and tryptophan.     

Effect of domestic processing on the cooking time,nutrients, antinutrients andin vitro Protein digestibility of the African yambean (Sphenostylis stenocarpa)

The effects of processing (soaking, dehulling, fermentation and heat treatment) on the cooking time, protien, mineral, tannin, phytate and in vitro protein digestibility (IVPD) of the african yambean (AYB) were examined. The cooking time ranged from 90–155 minutes. Soaking reduced cooking time by about 50 percent. Soaking for 12 hours was the most appropriate to reduce cooking time, tannin and phytate levels. It improved in vitro protein digestibility (IVPD). Prolonged soaking (24 hours) decreased calcium (Ca) and iron (Fe) values by 19 percent and 35 percent, respectively. Dehulling showed that Ca, Fe, magnesium (Mg) and zinc (Zn) were concentrated in the seed coat of the AYB. The seeds soaked and dehulled retained Mg and Zn. Dehulling reduced tannin but had no significant effect on phytate and the IVPD of the AYB except for seeds soaked for 12 hours before dehulling. Soaking for 24 hours before dehulling significantly increased crude protein content by 16 percent (p<0.05). Blanching and roasting increased the IVPD by 8–11 percent. Fermentation had no effect on the crude protein, Ca, Fe, Mg and Zn but significantly reduced phytate content of the AYB. Fermentation had no advantage over heat treatment with respect to improving the in vitro protein digestibility of the AYB.     

Effect of drought and elevated temperature on grain zinc and iron concentrations in CIMMYT spring wheat

Abiotic stress caused by increasing temperature and drought is a major limiting factor for wheat productivity around the world. Wheat plays an important role in feeding the world, but climate change threatens its future harvest and nutritional quality. In this study, grain iron (Fe) and zinc (Zn) concentrations of 54 wheat varieties, including CIMMYT derived historic and modern wheat varieties grown in six different environmental conditions, were analyzed. The objective of the study was to evaluate the effect of water and heat stress on the nutritional value of wheat grains with a main emphasis on grain protein content, Zn and Fe concentrations. Significant effects of environment on protein content and grain micronutrients concentration were observed. The protein and Zn concentrations increased in the water and heat stressed environments, whereas Zn and Fe yield per unit area was higher in non-stress conditions. The results suggest that genetic gains in the yield potential of CIMMYT derived wheat varieties have tended to reduce grain Zn, in some instances; however, environmental variability might influence the extent to which this effect manifests itself.     

玉米抗倒性研究进展

玉米倒伏严重影响产量、品质及机械化收获。通过育种和栽培方式提高抗倒性,对玉米丰产稳产尤为重要。本文综述玉米茎秆力学特性、化学成分、植株形态结构等对个体抗倒性的影响,栽培措施和病虫害对群体抗倒性的调控及玉米茎秆性状的遗传特性,展望未来提高玉米抗倒性的技术途径。