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Bread wheat is a major staple cereal provides more than 20% of dietary energy and protein supply to global population. However, with increasing population growth, the problem of nutritional deficiencies is increasingly affecting the health of resource people with predominantly cereal-based diet. Therefore, the development of wheat genotypes with micronutrient-dense grains along with high-yield potential is one of the major priorities of wheat biofortification program at CIMMYT. We conducted a QTL mapping study using a recombinant inbred line (RIL) population derived from a cross between a Chinese parental line with highGZnC and a Mexican commercial bread wheat cultivar Roelfs F2007 to identify QTLs that could potentially be integrated in mineral nutrient concentrations and agronomic-related traits breeding. We evaluated 200 RIL lines for mineral nutrient concentrations and agronomic-related traits over two years. A total of 60 QTLs were detected, of which 10 QTLs for GZnC, 9 for GFeC, 5 for GPC and 36 for agronomic-related traits. Moreover, a total of 55 promising candidate genes were identified from the list of associated markers for GFeC and GZnC using the recently annotated wheat genome sequence. We identified the promising genomic regions with high mineral nutrient concentrations and acceptable yield potential, which are good resource for further use in wheat biofortification breeding programs., Highlights • Rich genetic diversity for micronutrients in wheat landraces provides novel alleles. • QTLs identified for multiple traits offers breeding competitive biofortified wheat. • Genomic regions and the candidate genes identified enables marker assisted breeding.

Genomic selection (GS) models have been validated for many quantitative traits in wheat (Triticum aestivum L.) breeding. However, those models are mostly constrained within the same growing cycle and the extension of GS to the case of across cycles has been a challenge, mainly due to the low predictive accuracy resulting from two factors: reduced genetic relationships between different families and augmented environmental variances between cycles. Using the data collected from diverse field conditions at the International Wheat and Maize Improvement Center, we evaluated GS for grain yield in three elite yield trials across three wheat growing cycles. The objective of this project was to employ the secondary traits, canopy temperature, and green normalized difference vegetation index, which are closely associated with grain yield from high-throughput phenotyping platforms, to improve prediction accuracy for grain yield. The ability to predict grain yield was evaluated reciprocally across three cycles with or without secondary traits. Our results indicate that prediction accuracy increased by an average of 146% for grain yield across cycles with secondary traits. In addition, our results suggest that secondary traits phenotyped during wheat heading and early grain filling stages were optimal for enhancing the prediction accuracy for grain yield.

In this study, we defined the target population of environments (TPE) for wheat breeding in India, the largest wheat producer in South Asia, and estimated the correlated response to the selection and prediction ability of five selection environments (SEs) in Mexico. We also estimated grain yield (GY) gains in each TPE. Our analysis used meteorological, soil, and GY data from the international Elite Spring Wheat Yield Trials (ESWYT) distributed by the International Maize and Wheat Improvement Center (CIMMYT) from 2001 to 2016. We identified three TPEs: TPE 1, the optimally irrigated Northwestern Plain Zone; TPE 2, the optimally irrigated, heat-stressed North Eastern Plains Zone; and TPE 3, the drought-stressed Central-Peninsular Zone. The correlated response to selection ranged from 0.4 to 0.9 within each TPE. The highest prediction accuracies for GY per TPE were derived using models that included genotype-by-environment interaction and/or meteorological information and their interaction with the lines. The highest prediction accuracies for TPEs 1, 2, and 3 were 0.37, 0.46, and 0.51, respectively, and the respective GY gains were 118, 46, and 123 kg/ha/year. These results can help fine-tune the breeding of elite wheat germplasm with stable yields to reduce farmers’ risk from year-to-year environmental variation in India’s wheat lands, which cover 30 million ha, account for 100 million tons of grain or more each year, and provide food and livelihoods for hundreds of millions of farmers and consumers in South Asia.

Breeding for improved nutritional quality in major staples has been emerged as one of the sustainable solutions to alleviate micronutrient malnutrition problems in the developing countries. Wheat provides one-fifth of global dietary energy and protein demand worldwide. Additionally, wheat products, such as chapatti (flat bread), made of whole grain wheat flour are major sources of micronutrients like Zinc (Zn), Iron (Fe) and Manganese (Mn), Vitamin B and E. An estimated two billion people suffer from Zn and Fe deficiency mainly in South Asia and Sub-Saharan Africa. Therefore, genetic enhancement of grain Zn and Fe in an improved wheat genetic background offers cost-effective sustainable solution to the problem. Breeding for nutritional quality in wheat through enhanced concentrations of micronutrients, initiated under the HarvestPlus program by crossing high Zn and Fe sources, identified among synthetic wheats, T. spelta, and landraces from Mexico and Iran. These crosses have resulted in wheat lines with competitive yields and enhanced grain Zn in South Asia. QTL mapping and gene discovery research have identified 5–6 important QTL regions for grain Zn. The high Zn and Fe inheritance are under quantitative genetic control; further progress is possible through pyramiding large effect QTL regions in high-yielding wheats. High-throughput, non-destructive phenotyping for grain Zn and Fe using the X-ray fluorescence (XRF) analysis has facilitated the selection dramatically. Accelerated gene discovery and mapping studies, and genomic selection schemes expected to improve the breeding efficiency.

Background and aims Zinc (Zn) and iron (Fe) deficiencies are the most important forms of malnutrition globally, and caused mainly by low dietary intake. Wheat, a major staple food crop, is inherently low in these micronutrients. Identifying new QTLs for high grain Zn (GZn) and Fe (GFe) will contribute to improved micronutrient density in wheat grain.

Predictability estimated through cross-validation approach showed moderate to high level; hence, genomic selection approach holds great potential for biofortification breeding to enhance grain zinc and iron concentrations in wheat. Wheat (Triticum aestivum L.) is a major staple crop, providing 20 % of dietary energy and protein consumption worldwide. It is an important source of mineral micronutrients such as zinc (Zn) and iron (Fe) for resource poor consumers. Genomic selection (GS) approaches have great potential to accelerate development of Fe- and Zn-enriched wheat. Here, we present the results of large-scale genomic and phenotypic data from the HarvestPlus Association Mapping (HPAM) panel consisting of 330 diverse wheat lines to perform genomic predictions for grain Zn (GZnC) and Fe (GFeC) concentrations, thousand-kernel weight (TKW) and days to maturity (DTM) in wheat. The HPAM lines were phenotyped in three different locations in India and Mexico in two successive crop seasons (2011–12 and 2012–13) for GZnC, GFeC, TKW and DTM. The genomic prediction models revealed that the estimated prediction abilities ranged from 0.331 to 0.694 for Zn and from 0.324 to 0.734 for Fe according to different environments, whereas prediction abilities for TKW and DTM were as high as 0.76 and 0.64, respectively, suggesting that GS holds great potential in biofortification breeding to enhance grain Zn and Fe concentrations in bread wheat germplasm.

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.

Malnutrition because of the deficiency of minerals such as iron (Fe) and zinc (Zn) afflicts over 2 billion people worldwide. Wheat is a major staple crop, providing 20% of dietary energy and protein consumption worldwide. Breeding wheat with elevated levels of grain Zn and Fe concentrations (GZn and GFe) represents a significant opportunity to increase the intake of these micronutrients for the resource poor people who depend on it as a major source of dietary energy. Synthetic hexaploid wheats (SHWs) have large genetic variation for GZn and GFe, which can be exploited for developing wheat varieties with higher concentrations of these minerals. The objective of this study was to localise genomic regions associated with GZn and GFe, thousand kernel weight (TKW) and test weight (TW) in a mapping population derived from the cross of Seri M82 and the SHW CWI76364. Major quantitative trait loci (QTL) on chromosome 4BS were detected for GZn and GFe; the QTL explained up to 19.6% of the total phenotypic variation for GZn and showed pleiotropic effects on GFe. This indicates that simultaneous improvement of GZn and GFe is feasible. Three and five QTL for TW and TKW were detected, respectively. One of the QTL for TKW was also located on chromosome 4BS. Positive correlations between plant height and GZn/GFe were observed. The 4BS QTL is of great interest for breeding biofortified wheat by means of marker-assisted selection.

Highlights • Linear and consistent grain yield genetic gains in the HRWYT has been observed. • Several best performing lines were common in both high and low rainfall environments. • The genetic gains were explained by paralleled increases in grain weight, day to maturity and grain filling period. • These results indicate continuous genetic progress and yield stability in the HRWYT germplasm developed by CIMMYT., The effects of climate change together with the projected future demand represents a huge challenge for wheat production systems worldwide. Wheat breeding can contribute to global food security through the creation of genotypes exhibiting stress tolerance and higher yield potential. The objectives of our study were to (i) estimate the annual grain yield (GY) genetic gain of High Rainfall Wheat Yield Trials (HRWYT) grown from 2007 (15th HRWYT) to 2016 (24th HRWYT) across international environments, and (ii) determine the changes in physiological traits associated with GY genetic improvement. The GY genetic gains were estimated as genetic progress per se (GYP) and in terms of local checks (GYLC). In total, 239 international locations were classified into two groups: high- and low-rainfall environments based on climate variables and trial management practices. In the high-rainfall environment, the annual genetic gains for GYP and GYLC were 3.8 and 1.17 % (160 and 65.1 kg ha−1 yr−1), respectively. In the low-rainfall environment, the genetic gains were 0.93 and 0.73 % (40 and 33.1 kg ha−1 yr−1), for GYP and GYLC respectively. The GY of the lines included in each nursery showed a significant phenotypic correlation between high- and low-rainfall environments in all the examined years and several of the five best performing lines were common in both environments. The GY progress was mainly associated with increased grain weight (R2 = 0.35 p < 0.001), days to maturity (R2 = 0.20, p < 0.001) and grain filling period (R2 = 0.06, p < 0.05). These results indicate continuous GY genetic progress and yield stability in the HRWYT germplasm developed and distributed by CIMMYT.

The development of high-yielding wheat genotypes containing micronutrient-dense grains are the main priorities of biofortification programs. At the International Maize and Wheat Improvement Center, breeders have successfully crossed high zinc progenitors including synthetic hexaploid wheat, T. dicoccum, T. spelta and landraces to generate high-zinc varieties. In this study, we report a genome-wide association using a wheat diversity panel to dissect the genetics controlling zinc, iron, copper, manganese and phosphorus concentrations in the grain and rachis during grain development and at physiological maturity. Significant marker-trait associations (MTAs) were identified for each nutrient using multi-locus mixed model methodologies. For mature grain, markers that showed significant pleiotropic effects were found on chromosomes 1A, 3B and 5B, of which those on chromosome 5B at ∼95.5 cM were consistent over two growing seasons. Co-located MTAs were identified for the nutrient concentrations in developing grain, rachis and mature grain on multiple chromosomes. The identified genomic regions included putative candidate genes involved in metal uptake and transport and storage protein processing. These findings add to our understanding of the genetics of the five important nutrients in wheat grain and provide information on genetic markers for selecting high micronutrient genotypes.

At least two billion people around the world suffer from micronutrient deficiency, or hidden hunger, which is characterized by iron-deficiency anemia, vitamin A and zinc deficiency. As a key staple food crop, wheat provides 20% of the world’s dietary energy and protein, therefore wheat is an ideal vehicle for biofortification. Developing biofortified wheat varieties with genetically enhanced levels of grain zinc (Zn) and iron (Fe) concentrations, and protein content provides a cost-effective and sustainable solution to the resource-poor wheat consumers. Large genetic variation for Fe and Zn were found in the primitive and wild relatives of wheat, the potential high Zn and Fe containing genetic resources were used as progenitors to breed high-yielding biofortified wheat varieties with 30–40% higher Zn content. Grain protein content (GPC) determines processing and end-use quality of wheat for making diverse food products. The GPC-B1 allele from Triticum turgidum L. var. dicoccoides have been well characterized for the increase in GPC and the associated pleiotropic effect on grain Zn and Fe concentrations in wheat. In this study effect of GPC-B1 allele on grain Zn and Fe concentrations in wheat were measured in different genetic backgrounds and two different agronomic management practices (with- and without foliar Zn fertilization). Six pairs of near-isogenic lines differing for GPC-B1 gene evaluated at CIMMYT, Mexico showed that GPC-B1 influenced marginal increase for grain Zn, Fe concentrations, grain protein content and slight reduction in kernel weight and grain yield. However, the magnitude of GPC and grain Zn and Fe reductions varied depending on the genetic background. Introgression of GPC-B1 functional allele in combination with normal or delayed maturity alleles in the CIMMYT elite wheat germplasm has the potential to improve GPC and grain Zn and Fe concentrations without the negative effect on grain yield due to early senescence and accelerated maturity.

As a major cereal crop worldwide, wheat contributes on average one-fifth of the calories in the human diet and is the main source of protein and nutrients for much of the world's population. Wheat varieties with improved nutritional quality, high grain yield and desirable processing quality attributes in adapted genetic backgrounds can help alleviate nutrient deficiencies among resource poor people. This paper reports advances in targeted crosses of landraces and ancestors of common wheat ( Triticum aestivum L.), such as Aegilops tauschii , Triticum turgidum ssp. diccocoides , T. turgidum ssp. dicoccum and T. aestivum ssp. spelta species, which feature significant genetic variation for grain zinc and iron, with high-yielding bread wheat lines from the CIMMYT breeding program that have desirable processing and end-use quality. High-yielding lines that resulted from these crosses possessed preferred processing quality traits and 10–90% higher grain micronutrient concentrations than popular commercial varieties.

Zinc deficiency in humans is recognized as a widespread public health problem worldwide, but can be combated via genetic biofortification through breeding high zinc containing wheat varieties. CIMMYT (International Maize and Wheat Improvement Center, Int.) is engaged in enhancing, among others, the grain zinc concentration (GZnC) of high-yielding wheat germplasm under the HarvestPlus initiative of the Consultative Group on International Agricultural Research consortium. In the present study, we determined GZnC in a recombinant inbred line population from the cross between PBW343 and Kenya Swara in replicated trials grown in Zn-enriched field. An integrated genetic map with 1,133 loci (diversity arrays technology and simple sequence repeats markers) covering all 21 wheat chromosomes was constructed and used for quantitative trait loci (QTL) analysis. Two novel QTL of large effect were stably detected for increasing GZnC on chromosomes 2Bc (centromeric region) and 3AL (long arm). The two QTL individually explained about 10–15 % of the total phenotypic variation. The 2Bc QTL from PBW343 have pleiotropic effect and can increase thousand-kernel weight at significant level. The QTL and the closely linked markers identified will make selection for this difficult trait feasible in breeding program.

Ten QTL underlying the accumulation of Zn and Fe in the grain were mapped in a set of RILs bred from the cross Triticum spelta × T. aestivum . Five of these loci (two for Zn and three for Fe) were consistently detected across seven environments. The genetic basis of accumulation in the grain of Zn and Fe was investigated via QTL mapping in a recombinant inbred line (RIL) population bred from a cross between Triticum spelta and T. aestivum. The concentration of the two elements was measured from grain produced in three locations over two consecutive cropping seasons and from a greenhouse trial. The range in Zn and Fe concentration across the RILs was, respectively, 18.8–73.5 and 25.3–59.5 ppm, and the concentrations of the two elements were positively correlated with one another (rp =+0.79). Ten QTL (five each for Zn and Fe accumulation) were detected, mapping to seven different chromosomes. The chromosome 2B and 6A grain Zn QTL were consistently expressed across environments. The proportion of the phenotype explained (PVE) by QZn.bhu-2B was >16 %, and the locus was closely linked to the SNP marker 1101425|F|0, while QZn.bhu-6A (7.0 % PVE) was closely linked to DArT marker 3026160|F|0. Of the five Fe QTL detected, three, all mapping to chromosome 1A were detected in all seven environments. The PVE for QFe.bhu-3B was 26.0 %.

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.

Genetic biofortification to improve zinc (Zn) and iron (Fe) concentrations in bread wheat ( Triticum aestivum L.) could reduce micronutrient malnutrition-related problems in the developing world. A breeding program on wheat was started to enhance Zn and Fe concentrations and other essential traits needed in a successful commercial variety. The first set of advanced lines derived from crosses of high yielding wheats with genetic resources possessing high Zn and Fe such as Triticum spelta , landraces and synthetic wheat based on Triticum dicoccon were tested at nine locations in South Asia and Mexico for Zn and Fe concentration, grain yield and other traits. Analyses of variance across locations revealed significant genotypic, environmental and genotype × environment (G × E) effects for grain Zn and Fe concentrations and grain yield. Variances associated with environmental effects were larger than the genotypic and G × E effects for all three traits, suggesting that environmental effects have relatively greater influence. Although G × E interaction was significant, high heritabilities were observed for Zn and Fe concentrations at individual sites and across environments, reflecting non-crossover type of interaction. This trend was confirmed by the high genetic correlations between locations that showed similar ranking of entries across locations, indicating that it is possible to select the best adapted entries with high Zn and Fe concentration. Pooled data across locations showed increments of 28% and 25% over the checks for Zn and Fe. A considerable number of entries exceeded intermediate to full breeding target Zn concentrations, indicating that it is possible to develop Zn-biofortified varieties with competitive yields and other farmer preferred agronomic traits. The positive and moderately high correlation between Zn and Fe concentration suggest good prospects of simultaneous improvement for both micronutrients.

Pearl millet [Pennisetum glaucum (L.) R. Br.] is a major warm-season cereal, grown primarily for grain production in the arid and semi-arid tropical regions of Asia and Africa. Iron (Fe) and zinc (Zn) deficiencies have been reported to be a food-related primary health problem affecting nearly two billion people worldwide. Improving Fe and Zn densities of staple crops by breeding offers a cost-effective and sustainable solution to reducing micronutrient malnutrition in resource poor communities. An understanding of the genetics of these micronutrients can help to accelerate the breeding process, but little is known about the genetics and heterosis pattern of Fe and Zn densities in pearl millet. In the present study, ten inbred lines and their full diallel crosses were used to study the nature of gene action and heterosis for these micronutrients. The general combining ability (GCA) effects of parents and specific combining ability (SCA) effects of hybrids showed significant differences for both of the micronutrients. However, the predictability ratio (2σ2gca/(2σ2gca + σ2sca)) was around unity both for Fe and Zn densities, implying preponderance of additive gene action. Further, highly significant positive correlation between mid-parent values and hybrid performance, and no correlation between mid-parent values and mid-parent heterosis confirmed again the predominant role of additive gene action for these micronutrients. Barring a few exceptions with one parent, hybrids did not outperform the parents having high Fe and Zn levels. This showed that there would be little opportunity, if any, to exploit heterosis for these mineral micronutrients in pearl millet. In general, high Fe and Zn levels in both of the parental lines would be required to increase the probability of breeding high Fe and Zn hybrids.

Rust diseases continue to cause significant losses to wheat production worldwide. Although the life of effective race-specific resistance genes can be prolonged by using gene combinations, an alternative approach is to deploy varieties that posses adult plant resistance (APR) based on combinations of minor, slow rusting genes. When present alone, APR genes do not confer adequate resistance especially under high disease pressure; however, combinations of 4–5 such genes usually result in “near-immunity” or a high level of resistance. Although high diversity for APR occurs for all three rusts in improved germplasm, relatively few genes are characterized in detail. Breeding for APR to leaf rust and stripe rust in CIMMYT spring wheats was initiated in the early 1970s by crossing slow rusting parents that lacked effective race-specific resistance genes to prevalent pathogen populations and selecting plants in segregating populations under high disease pressure in field nurseries. Consequently most of the wheat germplasm distributed worldwide now possesses near-immunity or adequate levels of resistance. Some semidwarf wheats such as Kingbird, Pavon 76, Kiritati and Parula show high levels of APR to stem rust race Ug99 and its derivatives based on the Sr2-complex, or a combination of Sr2 with other uncharacterized slow rusting genes. These parents are being utilized in our crossing program and a Mexico-Kenya shuttle breeding scheme is used for selecting resistance to Ug99. High frequencies of lines with near-immunity to moderate levels of resistance are now emerging from these activities. After further yield trials and quality assessments these lines will be distributed internationally through the CIMMYT nursery system.

Approximately 25 % of global agricultural land is utilized for wheat cultivation, making wheat the largest food crop worldwide in terms of area. Wheat is the second most produced cereal crop after Maize with more than 650 million tons produced every year. Wheat productivity is increasing at less than 1 percent annually, while the annual productivity must increase at 2 % annually to meet the global demand. The potential of increasing arable land is limited; hence future increases in wheat production must be achieved by enhancing the productivity per unit area. Increasing grain yield, yield stability, resistance/tolerance to biotic and abiotic stresses, and end-use quality characteristics are among the most important wheat breeding goals.