Genomic approaches for improving grain zinc and iron content in wheat

More than three billion people worldwide suffer from iron deficiency associated anemia and an equal number people suffer from zinc deficiency. These conditions are more prevalent in Sub-Saharan Africa and South Asia. In developing countries, children under the age of five with stunted growth and pregnant or lactating women were found to be at high risk of zinc and iron deficiencies. Biofortification, defined as breeding to develop varieties of staple food crops whose grain contains higher levels of micronutrients such as iron and zinc, are one of the most promising, cost-effective and sustainable ways to improve the health in resource-poor households, particularly in rural areas where families consume some part of what they grow. Biofortification through conventional breeding in wheat, particularly for grain zinc and iron, have made significant contributions, transferring important genes and quantitative trait loci (QTLs) from wild and related species into cultivated wheat. Nonetheless, the quantitative, genetically complex nature of iron and zinc levels in wheat grain limits progress through conventional breeding, making it difficult to attain genetic gain both for yield and grain mineral concentrations. Wheat biofortification can be achieved by enhancing mineral uptake, source-to-sink translocation of minerals and their deposition into grains, and the bioavailability of the minerals. A number of QTLs with major and minor effects for those traits have been detected in wheat; introducing the most effective into breeding lines will increase grain zinc and iron concentrations. New approaches to achieve this include marker assisted selection and genomic selection. Faster breeding approaches need to be combined to simultaneously increase grain mineral content and yield in wheat breeding lines

Improving zinc and iron biofortification in wheat through genomics approaches

Globally, about 20% of calories (energy) come from wheat. In some countries, it is more than 70%. More than 2 billion people are at risk for zinc deficiency and even more, people are at risk of iron deficiency, nearly a quarter of all children underage group of 5 are physically and cognitively stunted, and lack of dietary zinc is a major contributing factor. Biofortified wheat with elevated levels of zinc and iron has several potential advantages as a delivery vehicle for micronutrients in the diets of resource-poor consumers who depend on cereal-based diets. The conventional breeding strategies have been successful in the introduction of novel alleles for grain Zn and Fe that led to the release of competitive Zn enriched wheat varieties in South Asia. The major challenge over the next few decades will be to maintain the rates of genetic gains for grain yield along with increased grain Zn/Fe concentration to meet the food and nutritional security challenges. Therefore, to remain competitive, the performance of Zn-enhanced lines/varieties must be equal or superior to that of current non-biofortified elite lines/varieties. Since both yield and Zn content are invisible and quantitatively inherited traits except few intermediate effect QTL regions identified for grain Zn, increased breeding efforts and new approaches are required to combine them at high frequency, ensuring that Zn levels are steadily increased to the required levels across the breeding pipelines. The current review article provides a comprehensive list of genomic regions for enhancing grain Zn and Fe concentrations in wheat including key candidate gene families such NAS, ZIP, VLT, ZIFL, and YSL. Implementing forward breeding by taking advantage of the rapid cycling trait pipeline approaches would simultaneously introgress high Zn and Fe QTL into the high Zn and normal elite lines, further increasing Zn and Fe concentrations

Assessing Genetic Diversity to Breed Competitive Biofortified Wheat With Enhanced Grain Zn and Fe Concentrations

Breeding wheat with enhanced levels of grain zinc (Zn) and iron (Fe) is a cost-effective, sustainable solution to malnutrition problems. Modern wheat varieties have limited variation in grain Zn and Fe, but large-scale screening has identified high levels of Zn and Fe in wild relatives and progenitors of cultivated wheat. The most promising sources of high Zn and Fe are einkorn (Triticum monococcum), wild emmer (T. dicoccoides), diploid progenitors of hexaploid wheat (such as Aegilops tauschii), T. spelta, T. polonicum, and landraces of T. aestivum. This study evaluate the effects of translocations from rye and different Aegilops species in a “Pavon-76” wheat genetic background and utilized in the wheat biofortification breeding program at CIMMYT that uses diverse genetic resources, including landraces, recreated synthetic hexaploids, T. spelta and pre-breeding lines. Four translocations were identified that resulted significantly higher Zn content in “Pavon 76” genetic background than the check varieties, and they had increased levels of grain Fe as well-compared to “Pavon 76.” These lines were also included in the breeding program aimed to develop advanced high Zn breeding lines. Advanced lines derived from diverse crosses were screened under Zn-enriched soil conditions in Mexico during the 2017 and 2018 seasons. The Zn content of the grain was ranging from 35 to 69 mg/kg during 2017 and 38 to 72 mg/kg during 2018. Meanwhile grain Fe ranged from 30 to 43 mg/kg during 2017 and 32 to 52 mg/kg during 2018. A highly significant positive correlation was found between Zn and Fe (r = 0.54; P < 0.001) content of the breeding lines, therefore it was possible to breed for both properties in parallel. Yield testing of the advanced lines showed that 15% (2017) and 24% (2018) of the lines achieved 95–110% yield potential of the commercial checks and also had 12 mg/kg advantage in the Zn content suggesting that greater genetic gains and farmer-preferred wheat varieties were developed and deployed. A decade of research and breeding efforts led to the selection of “best-bet” breeding lines and the release of eight biofortified wheat varieties in target regions of South Asia and in Mexico

Identification of Genetic Loci and Candidate Genes Related to Grain Zinc and Iron Concentration Using a Zinc-Enriched Wheat ‘Zinc-Shakti’

The development of nutritionally enhanced wheat (Triticum aestivum L.) with higher levels of grain iron (Fe) and zinc (Zn) offers a sustainable solution to micronutrient deficiency among resource-poor wheat consumers. One hundred and ninety recombinant inbred lines (RILs) from ‘Kachu’ × ‘Zinc-Shakti’ cross were phenotyped for grain Fe and Zn concentrations and phenological and agronomically important traits at Ciudad Obregon, Mexico in the 2017–2018, 2018–2019, and 2019–2020 growing seasons and Diversity Arrays Technology (DArT) molecular marker data were used to determine genomic regions controlling grain micronutrients and agronomic traits. We identified seven new pleiotropic quantitative trait loci (QTL) for grain Zn and Fe on chromosomes 1B, 1D, 2B, 6A, and 7D. The stable pleiotropic QTL identified have expanded the diversity of QTL that could be used in breeding for wheat biofortification. Nine RILs with the best combination of pleiotropic QTL for Zn and Fe have been identified to be used in future crossing programs and to be screened in elite yield trials before releasing as biofortified varieties. In silico analysis revealed several candidate genes underlying QTL, including those belonging to the families of the transporters and kinases known to transport small peptides and minerals (thus assisting mineral uptake) and catalyzing phosphorylation processes, respectively

QTL Mapping of Grain Zn and Fe Concentrations in Two Hexaploid Wheat RIL Populations with Ample Transgressive Segregation

This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these termsMore than 50% of undernourished children live in Asia and more than 25% live in Africa. Coupled with an inadequate food supply, mineral deficiencies are widespread in these populations; particularly zinc (Zn) and iron (Fe) deficiencies that lead to retarded growth, adverse effects on both the immune system and an individual's cognitive abilities. Biofortification is one solution aimed at reducing the incidence of these deficiencies. To efficiently breed a biofortified wheat variety, it is important to generate knowledge of the genomic regions associated with grain Zn (GZn) and Fe (GFe) concentration. This allows for the introgression of favorable alleles into elite germplasm. In this study we evaluated two bi-parental populations of 188 recombinant inbred lines (RILs) displaying a significant range of transgressive segregation for GZn and GFe during three crop cycles in CIMMYT, Mexico. Parents of the RILs were derived from Triticum spelta L. and synthetic hexaploid wheat crosses. QTL analysis identified a number of significant QTL with a region denominated as QGZn.cimmyt-7B_1P2 on chromosome 7B explaining the largest (32.7%) proportion of phenotypic variance (PVE) for GZn and leading to an average additive effect of −1.3. The QTL with the largest average additive effect for GFe (−0.161) was found on chromosome 4A (QGFe.cimmyt-4A_P2), with 21.14% of the PVE. The region QGZn.cimmyt-7B_1P2 co-localized closest to the region QGZn.cimmyt-7B_1P1 in a consensus map built from the linkage maps of both populations. Pleiotropic or tightly linked QTL were also found on chromosome 3B, however of minor effects and PVE between 4.3 and 10.9%. Further efforts are required to utilize the QTL information in marker assisted backcrossing schemes for wheat biofortification. A strategy to follow is to intercross the transgressive individuals from both populations and then utilize them as sources in biofortification breeding pipelines

Yield and Quality in Purple-Grained Wheat Isogenic Lines

Breeding programs for purple wheat are underway in many countries but there is a lack of information on the effects of Pp (purple pericarp) genes on agronomic and quality traits in variable environments and along the product chain (grain-flour-bread). This study was based on unique material: two pairs of isogenic lines in a spring wheat cv. Saratovskaya-29 (S29) background differing only in Pp genes and grain color. In 2017, seven experiments were conducted in Kazakhstan, Russia, and Turkey with a focus on genotype and environment interaction and, in 2018, one experiment in Turkey with a focus on grain, flour, and bread quality. The eect of environment was greater compared to genotype for the productivity and quality traits studied. Nevertheless, several important traits, such as grain color and anthocyanin content, are closely controlled by genotype, offering the opportunity for selection. Phenolic content in purple-grained lines was not significantly higher in whole wheat flour than in red-colored lines. However, this trait was significantly higher in bread. For antioxidant activities, no differences between the genotypes were detected in both experiments. Comparison of two sources of Pp genes demonstrated that the lines originating from cv. Purple Feed had substantially improved productivity and quality traits compared to those from cv. Purple

Localisation of iron and zinc in grain of biofortified wheat

The dietary contributions of iron (Fe) and zinc (Zn) from cereals are determined by concentrations, locations and chemical forms. A genetically biofortified wheat line showed higher concentrations of Zn and Fe than three control lines when grown over two years. The mineral distributions determined using imaging (histochemical staining and LA-ICP-MS), sequential pearling and hand dissection showed no consistent differences between the two lines. Fe was most abundant in the aleurone layer and the scutellum and Zn in the scutellar epithelium, the endosperm transfer cells and embryonic axis. Pearling fractions showed positive correlations between the concentration of P and those of Zn and Fe in all fractions except the outermost layer. This is consistent with Fe and Zn being concentrated in phytates. Developing grains showed decreasing gradients in concentration from the proximal to the distal ends. The concentrations of Fe and Zn were therefore higher in the biofortified line than the control lines but their locations did not differ

Editorial: Wheat biofortification to alleviate global malnutrition

According to the latest FAO report on the state of food security and nutrition in the world (1), more than 720 million people faced hunger, and around 3 billion people did not have access to a healthy diet. All these problematics, exacerbated by the current COVID-19 crisis, led to an increase in the number of people affected by the so-called hidden hunger, caused by an inadequate intake of essential micronutrients (MNs) such as iron (Fe), zinc (Zn), selenium (Se) and provitamin A. Biofortification, intended as the improvement of the nutritional quality of food crops through either conventional breeding, agronomic practices ormodern biotechnologies, represents a sustainable, costeffective and long-term approach to alleviate micronutrient-deficiency. Staple crops are typically the major target of most biofortification studies, given their central role in human diet. Wheat, specifically, contributes to around 20% of the total energy and protein intake and to around 30% of the Fe and Zn intake worldwide. However, the current level of MNs present in most wheat-derived food products is not enough to meet the minimum daily intake, especially in the poorest regions of the world. For these reasons, continuing to work on wheat biofortification is fundamental to ensure the production of nutritious and sustainable food and to contribute to the reduction of MNs deficiency

Effects of glutenins (Glu-1 and Glu-3) allelic variation on dough properties and bread-making quality of CIMMYT bread wheat breeding lines

Wheat dough characteristics and end-use quality are strongly influenced by the amount and specific composition of the glutenins, the major components of gluten. Such proteins are divided into high-molecular-weight glutenins, encoded by the Glu-A1, Glu-B1 and Glu-D1 loci; and low-molecular-weight glutenins, encoded by the Glu-A3, Glu-B3 and Glu-D3 loci. Allelic variation at each of these loci has been associated with changes in wheat functionality. However, most of the studies conducted so far included a relatively limited number of genotypes. Also for this reason, it is still unclear which locus contributes more to dough characteristics and how important are the interactions between the glutenin loci. To try to answer these questions, the quality data of 4623 grain samples derived from 2550 genotypes and generated across 10 years at the CIMMYT bread wheat breeding program, was used to estimate the effect of the glutenin loci and their interactions on gluten quality and bread-making potential. Gluten strength was the trait more strongly influenced by glutenin variations, with the Glu-B1, Glu-D1 and Glu-B3 loci having the greatest effect. Among the glutenin alleles, Glu-A1a, Glu-A1b, Glu-B1al, Glu-B1i, Glu-B1f, Glu-D1d, Glu-A3b, Glu-A3d, Glu-A3f, Glu-B3c and Glu-B3d were associated in general with greater gluten strength, good extensibility and higher bread loaf volume. Differently, alleles Glu-A1c, Glu-B1a, Glu-B1d, Glu-D1a, Glu-A3e and Glu-B3j were associated with an overall poor quality. Glutenin interactions were significantly associated with most of the analyzed quality traits even if their influence was often lower compared to the effect of the single glutenin loci. This is probably the largest study ever done on the effects of the glutenins on wheat quality. The results obtained confirm the importance of such proteins on wheat quality variation and corroborate the usefulness of determining the glutenin profile to improve the selection efficiency for wheat quality in breeding programs