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Abstract Common wheat (Triticum aestivum L.) is a major staple food crop, providing a fifth of food calories and proteins to the world’s human population. Despite the impressive growth in global wheat production in recent decades, further increases in grain yield are required to meet future demands. Here we estimated genetic gain and genotype stability for grain yield (GY) and determined the trait associations that contributed uniquely or in combination to increased GY, through a retrospective analysis of top-performing genotypes selected from the elite spring wheat yield trial (ESWYT) evaluated internationally during a 14-year period (2003 to 2016). Fifty-six ESWYT genotypes and four checks were sown under optimally irrigated conditions in three phenotyping trials during three consecutive growing seasons (2018–2019 to 2020–2021) at Norman E. Borlaug Research Station, Ciudad Obregon, Mexico. The mean GY rose from 6.75 (24th ESWYT) to 7.87 t ha−1 (37th ESWYT), representing a cumulative increase of 1.12 t ha−1. The annual genetic gain for GY was estimated at 0.96% (65 kg ha−1 year−1) accompanied by a positive trend in genotype stability over time. The GY progress was mainly associated with increases in biomass (BM), grain filling rate (GFR), total radiation use efficiency (RUE_total), grain weight per spike (GWS), and reduction in days to heading (DTH), which together explained 95.5% of the GY variation. Regression lines over the years showed significant increases of 0.015 kg m−2 year−1 (p

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.

We untangled key regions of the genetic architecture of grain yield (GY) in CIMMYT spring bread wheat by conducting a haplotype-based, genome-wide association study (GWAS), together with an investigation of epistatic interactions using seven large sets of elite yield trials (EYTs) consisting of a total of 6,461 advanced breeding lines. These lines were phenotyped under irrigated and stress environments in seven growing seasons (2011–2018) and genotyped with genotyping-by-sequencing markers. Genome-wide 519 haplotype blocks were constructed, using a linkage disequilibrium-based approach covering 14,036 Mb in the wheat genome. Haplotype-based GWAS identified 7, 4, 10, and 15 stable (significant in three or more EYTs) associations in irrigated (I), mild drought (MD), severe drought (SD), and heat stress (HS) testing environments, respectively. Considering all EYTs and the four testing environments together, 30 stable associations were deciphered with seven hotspots identified on chromosomes 1A, 1B, 2B, 4A, 5B, 6B, and 7B, where multiple haplotype blocks were associated with GY. Epistatic interactions contributed significantly to the genetic architecture of GY, explaining variation of 3.5–21.1%, 3.7–14.7%, 3.5–20.6%, and 4.4– 23.1% in I, MD, SD, and HS environments, respectively. Our results revealed the intricate genetic architecture of GY, controlled by both main and epistatic effects. The importance of these results for practical applications in the CIMMYT breeding program is discussed.

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.

Inherently low concentrations of zinc (Zn), iron (Fe), iodine (I), and selenium (Se) in wheat (Triticum aestivum L.) grains represent a major cause of micronutrient malnutrition (hidden hunger) in human populations. Genetic biofortification represents a highly useful solution to this problem. However, genetic biofortification alone may not achieve desirable concentrations of micronutrients for human nutrition due to several soil- and plant-related factors. This study investigated the response of genetically biofortified high-Zn wheat genotypes to soil-applied Zn and foliarly applied Zn, I, and Se in India and Pakistan. The effect of soil-applied Zn (at the rate of 50 kg ha−1 as ZnSO4·7H2O) and foliar-applied Zn (0.5% ZnSO4·7H2O), I (0.04% KIO3), Se (0.001% Na2SeO4), and a foliar cocktail (F-CT: combination of the above foliar solutions) on the grain concentrations of Zn, I, Se, and Fe of high-Zn wheat genotypes was investigated in field experiments over 2 years. The predominantly grown local wheat cultivars in both countries were also included as check cultivars. Wheat grain yield was not influenced by the micronutrient treatments at all field locations, except one location in Pakistan where F-CT resulted in increased grain yield. Foliar-applied Zn, I, and Se each significantly enhanced the grain concentration of the respective micronutrients. Combined application of these micronutrients was almost equally effective in enhancing grain Zn, I, and Se, but with a slight reduction in grain yield. Foliar-applied Zn, Zn+I, and F-CT also enhanced grain Fe. In India, high-Zn genotypes exhibited a minor grain yield penalty as compared with the local cultivar, whereas in Pakistan, high-Zn wheat genotypes could not produce grain yield higher than the local cultivar. The study demonstrates that there is a synergism between genetic and agronomic biofortification in enrichment of grains with micronutrients. Foliar Zn spray to Zn-biofortified genotypes provided additional increments in grain Zn of more than 15 mg kg−1. Thus, combining agronomic and genetic strategies will raise grain Zn over 50 mg kg−1. A combination of fertilization practice with plant breeding is strongly recommended to maximize accumulation of micronutrients in food crops and to make significant progress toward resolving the hidden hunger problem in human populations. [ABSTRACT FROM AUTHOR]

Rapid urbanization and evolving dietary preferences have heightened the demand for durum wheat and its derivatives in developing nations like Nepal. This study represents the first comprehensive exploration and evaluation of durum wheat genotypes in Nepal, addressing the escalating need for highyielding varieties. The primary objective was to identify stable and prolific durum wheat lines for release, enhancing Nepal's durum wheat breeding program. Utilizing genotypes from CIMMYT's disease screening and yield nurseries from 2011/12 to 2020/21, a total of 132 genotypes, including international checks, underwent evaluation over ten years under the Alpha Lattice design. Results revealed significant variation among genotypes for grain yield and other traits, identifying high-yielding and stable lines suitable for Nepal. Heritability analysis highlighted moderate heritability for grain number per spike, thousand-grain weight, and grain yield. Cluster analysis identified distinct clusters with high grain yield and desirable agronomic traits. Disease incidence facilitated the selection of resistant lines, with DWK38 emerging as the highest grain yielder (4416.04 kg/ha). Overall, durum wheat lines from CIMMYT exhibited robust performance in Nepal, enabling the identification of superior lines with potential benefits for farmers and consumers. The study's implications include developing and releasing superior durum lines in Nepal, providing farmers with profitable alternatives amidst evolving food habits. In conclusion, the findings from this study provide a valuable foundation for future durum wheat breeding efforts in Nepal, guiding the selection of genotypes that are well-suited to the diverse environmental challenges of the region. [ABSTRACT FROM AUTHOR]

Heat and drought are two important constraints to global wheat productivity; understanding the genotypic responses for quality parameters under harsh production conditions (drought and heat) is very important for developing nutrient‐dense wheat varieties. A set of 15 modern bread wheat (Triticum aestivum L. subsp. aestivum) and durum wheat (Triticum turgidum subsp. durum) cultivars were tested in nine environments, including three different production conditions (normal, heat and drought) during 2020–21. Genotype stability performance for yield, nutrition and quality parameters is assessed using multienvironment trials through AMMI and GGE Biplot analysis. We discovered intriguing stress dynamics in grain zinc content (Zn) and grain iron content (Fe). Under heat stress, zinc concentration increases but decreases under drought stress, while iron does the opposite. Selecting zinc, starch and kernel weight under terminal heat stress can boost yield. Protein content and yield are inversely related, making it difficult for breeders to optimise both traits. G × E interactions and stability indices across all environments have found genotypes with high‐yielding stable genotypes, G12 (MP1358) (42.09 ppm) and G5 (HI1544) (42.41 ppm) have high Fe content. G12 (MP1358) (14.98%) ranked highest in protein concentration. Meanwhile, for Zn content, G11 (MACS 4058) (45.23 ppm) and G15 (WH730) (42.44 ppm) were top performers across environments. G7 (HI 1636) and G12 (MP1358) stand out as a win‐win genotype for their high potential and stability in yield, protein, Zn and Fe content. Our study shows the complex relationships and possible suggestions for targeted breeding programmes under heat and drought stress conditions to improve wheat grain quality and micronutrient profiles without yield loss. [ABSTRACT FROM AUTHOR]

Bread wheat (T. aestivum) is one of the world's most widely consumed cereals. Since micronutrient deficiencies are becoming more common among people who primarily depend upon cereal-based diets, a need for better-quality wheat varieties has been felt. An association panel of 154 T. aestivum lines was evaluated for the following quality traits: grain appearance (GA) score, grain hardness (GH), phenol reaction (PR) score, protein percent, sodium dodecyl sulfate (SDS) sedimentation value, and test weight (TWt). In addition, the panel was also phenotyped for grain yield and related traits such as days to heading, days to maturity, plant height, and thousand kernel weight for the year 2017-18 at the Borlaug Institute for South Asia (BISA) Ludhiana and Jabalpur sites. We performed a genome-wide association analysis on this panel using 18,351 genotyping-bysequencing (GBS) markers to find marker-trait associations for quality and grain yield-related traits. We detected 55 single nucleotide polymorphism (SNP) marker trait associations (MTAs) for quality-related traits on chromosomes 7B (10), 1A (9), 2A (8), 3B (6), 2B (5), 7A (4), and 1B (3), with 3A, 4A, and 6D, having two and the rest, 4B, 5A, 5B, and 1D, having one each. Additionally, 20 SNP MTAs were detected for yield-related traits based on a field experiment conducted in Ludhiana on 7D (4) and 4D (3) chromosomes, while 44 SNP MTAs were reported for Jabalpur on chromosomes 2D (6), 7A (5), 2A (4), and 4A (4). Utilizing these loci in marker-assisted selection will benefit from further validation studies for these loci to improve hexaploid wheat for better yield and grain quality [ABSTRACT FROM AUTHOR]

The use of plant genetic resources (PGR)—wild relatives, landraces, and isolated breeding gene pools—has had substantial impacts on wheat breeding for resistance to biotic and abiotic stresses, while increasing nutritional value, end‐use quality, and grain yield. In the Global South, post‐Green Revolution genetic yield gains are generally achieved with minimal additional inputs. As a result, production has increased, and millions of hectares of natural ecosystems have been spared. Without PGR‐derived disease resistance, fungicide use would have easily doubled, massively increasing selection pressure for fungicide resistance. It is estimated that in wheat, a billion liters of fungicide application have been avoided just since 2000. This review presents examples of successful use of PGR including the relentless battle against wheat rust epidemics/pandemics, defending against diseases that jump species barriers like blast, biofortification giving nutrient‐dense varieties and the use of novel genetic variation for improving polygenic traits like climate resilience. Crop breeding genepools urgently need to be diversified to increase yields across a range of environments (>200 Mha globally), under less predictable weather and biotic stress pressure, while increasing input use efficiency. Given that the ~0.8 m PGR in wheat collections worldwide are relatively untapped and massive impacts of the tiny fraction studied, larger scale screenings and introgression promise solutions to emerging challenges, facilitated by advanced phenomic and genomic tools. The first translocations in wheat to modify rhizosphere microbiome interaction (reducing biological nitrification, reducing greenhouse gases, and increasing nitrogen use efficiency) is a landmark proof of concept. Phenomics and next‐generation sequencing have already elucidated exotic haplotypes associated with biotic and complex abiotic traits now mainstreamed in breeding. Big data from decades of global yield trials can elucidate the benefits of PGR across environments. This kind of impact cannot be achieved without widescale sharing of germplasm and other breeding technologies through networks and public–private partnerships in a pre‐competitive space. [ABSTRACT FROM AUTHOR]

In the field of plant breeding, variousmachine learning models have been developed and studied to evaluate the genomic prediction (GP) accuracy of unseen phenotypes. Deep learning has shown promise. However, most studies on deep learning in plant breeding have been limited to small datasets, and only a few have explored its application in moderate-sized datasets. In this study, we aimed to address this limitation by utilizing a moderately large dataset. We examined the performance of a deep learning (DL) model and compared it with the widely used and powerful best linear unbiased prediction (GBLUP)model. The goal was to assess the GP accuracy in the context of a five-fold cross-validation strategy and when predicting complete environments using the DL model. The results revealed the DL model outperformed the GBLUP model in terms of GP accuracy for two out of the five included traits in the five-fold cross-validation strategy, with similar results in the other traits. This indicates the superiority of the DL model in predicting these specific traits. Furthermore, when predicting complete environments using the leave-oneenvironment-out (LOEO) approach, the DL model demonstrated competitive performance. It is worth noting that the DL model employed in this study extends a previously proposed multi-modal DL model, which had been primarily applied to image data but with small datasets. By utilizing a moderately large dataset, we were able to evaluate the performance and potential of the DL model in a context with more information and challenging scenario in plant breeding. [ABSTRACT FROM AUTHOR]

Motivation Developing new crop varieties with superior performance is highly important to ensure robust and sustainable global food security. The speed of variety development is limited by long field cycles and advanced generation selections in plant breeding programs. While methods to predict yield from genotype or phenotype data have been proposed, improved performance and integrated models are needed. Results We propose a machine learning model that leverages both genotype and phenotype measurements by fusing genetic variants with multiple data sources collected by unmanned aerial systems. We use a deep multiple instance learning framework with an attention mechanism that sheds light on the importance given to each input during prediction, enhancing interpretability. Our model reaches 0.754 ± 0.024 Pearson correlation coefficient when predicting yield in similar environmental conditions; a 34.8% improvement over the genotype-only linear baseline (0.559 ± 0.050). We further predict yield on new lines in an unseen environment using only genotypes, obtaining a prediction accuracy of 0.386 ± 0.010, a 13.5% improvement over the linear baseline. Our multi-modal deep learning architecture efficiently accounts for plant health and environment, distilling the genetic contribution and providing excellent predictions. Yield prediction algorithms leveraging phenotypic observations during training therefore promise to improve breeding programs, ultimately speeding up delivery of improved varieties. Availability and implementation Available at https://github.com/BorgwardtLab/PheGeMIL (code) and https://doi.org/doi:10.5061/dryad.kprr4xh5p (data). [ABSTRACT FROM AUTHOR]

Genomic regions governing days to heading (DH), days to maturity (DM), plant height (PH), thousand-kernel weight (TKW), and test weight (TW) were investigated in a set of 190 RILs derived from a cross between a widely cultivated wheat-variety, Kachu (DPW-621-50), and a high-zinc variety, Zinc-Shakti. The RIL population was genotyped using 909 DArTseq markers and phenotyped in three environments. The constructed genetic map had a total genetic length of 4665 cM, with an average marker density of 5.13 cM. A total of thirty-seven novel quantitative trait loci (QTL), including twelve for PH, six for DH, five for DM, eight for TKW and six for TW were identified. A set of 20 stable QTLs associated with the expression of DH, DM, PH, TKW, and TW were identified in two or more environments. Three novel pleiotropic genomic-regions harboring co-localized QTLs governing two or more traits were also identified. In silico analysis revealed that the DArTseq markers were located on important putative candidate genes such as MLO-like protein, Phytochrome, Zinc finger and RING-type, Cytochrome P450 and pentatricopeptide repeat, involved in the regulation of pollen maturity, the photoperiodic modulation of flowering-time, abiotic-stress tolerance, grain-filling duration, thousand-kernel weight, seed morphology, and plant growth and development. The identified novel QTLs, particularly stable and co-localized QTLs, will be validated to estimate their effects in different genetic backgrounds for subsequent use in marker-assisted selection (MAS). [ABSTRACT FROM AUTHOR]

Undernourishment of nutrients, also known as hidden hunger, affects over 2 billion populace globally. Even though stunting among children below five years of age has decreased in India in the last ten years, India is home to roughly thirty percent of the world's population of stunted pre-schoolers. A significant improvement has been witnessed in the targeted development and deployment of biofortified crops; approximately 20 million farm households from developing counties benefit from cultivating and consuming biofortified crops. There is ample scope for including biofortified varieties in the seed chain, ensuring nutritional security. Wheat is a dietary staple in India, typically consumed as wholemeal flour in the form of flatbreads such as chapatti and roti. Wheat contributes to nearly one fifth of global energy requirements and can also provide better amounts of iron (Fe) and zinc (Zn). As a result, biofortified wheat can serve as a medium for delivery of essential micronutrients such as Fe and Zn to end users. This review discusses wheat biofortification components such as Fe and Zn dynamics, its uptake and movement in plants, the genetics of their buildup, and the inclusion of biofortified wheat varieties in the seed multiplication chain concerning India. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Wheat is one of the most widely grown and consumed food crops in the world. Spot blotch and terminal heat stress are the two significant constraints mainly in the Indo–Gangetic plains of South Asia. The study was undertaken using 185 recombinant lines (RILs) derived from the interspecific hybridization of 'Triticum aestivum (HUW234) × T. spelta (H+26)' to reveal genomic regions associated with tolerance to combined stress to spot blotch and terminal heat. Different physiological (NDVI, canopy temperature, leaf chlorophyll) and grain traits (TGW, grain size) were observed under stressed (spot blotch, terminal heat) and non-stressed environments. The mean maturity duration of RILs under combined stress was reduced by 12 days, whereas the normalized difference vegetation index (NDVI) was 46.03%. Similarly, the grain size was depleted under combined stress by 32.23% and thousand kernel weight (TKW) by 27.56% due to spot blotch and terminal heat stress, respectively. The genetic analysis using 6734 SNP markers identified 37 significant loci for the area under the disease progress curve (AUDPC) and NDVI. The genome-wide functional annotation of the SNP markers revealed gene functions such as plant chitinases, NB-ARC and NBS-LRR, and the peroxidase superfamily Cytochrome P450 have a positive role in the resistance through a hypersensitive response. Zinc finger domains, cysteine protease coding gene, F-box protein, ubiquitin, and associated proteins, play a substantial role in the combined stress of spot blotch and terminal heat in bread wheat, according to genomic domains ascribed to them. The study also highlights T. speltoides as a source of resistance to spot blotch and terminal heat tolerance. [ABSTRACT FROM AUTHOR]

Estimation of the rate of genetic gain over time allows quantification of breeding progress. Here we report on the rate of grain yield and zinc (Zn) concentration increase over 11 yr of targeted wheat (Triticum aestivum L.) biofortification breeding at the International Maize and Wheat Improvement Center (CIMMYT). Data from yield trials evaluated across multiple locations in South Asia and beyond showed that average annual increases in grain yield potential of ∼1.5% and 0.9% per year gains for grain Zn and Fe concentrations. Across locations in all countries, mean yields of the five highest‐yielding entries showed an annual gain of 109 kg ha−1 yr−1 in yield and 0.3 mg kg−1 for grain Fe and Zn as well as 0.66 g for the yield component thousand‐grain weight. There was a strong positive correlation between Fe and Zn (r =.42) across locations, whereas no genetic correlation was observed between grain yield and Zn (r =.05) across locations. Despite the slight negative relationship between yield and Zn, through targeted crossing and development of large segregating populations we were able to identify transgressive segregants combining increased yield and Zn concentrations. Significant differences between lines for grain micronutrient concentrations were detected, and significant location effects on grain Zn and Fe concentrations were observed. These results demonstrate that continuous and simultaneous genetic gain for grain yield and concentrations of Fe and Zn is possible in elite spring bread wheat lines with potential to deliver global impact through identification of superior parents for use by national breeding programs and the release of biofortified wheat cultivars. Core Ideas: Significant genetic gains for grain yield, Zn and Fe concentration, and kernel weight were measured among high‐Zn biofortified lines.Biofortified lines yield gains were at par with other breeding programs showing simultaneous gains for Zn and yield.A strong positive association between grain Zn and Fe showed similar trends in genetic gains.Competitive high‐Zn lines developed at CIMMYT are being released as biofortified cultivars in target countries.As mainstreaming of grain Zn in the CIMMYT wheat breeding program is being implemented, it is expected that a large proportion of CIMMYT elite lines distributed to partners would be Zn enriched. [ABSTRACT FROM AUTHOR]

Spring bread wheat adaptation to diverse environments is supported by various traits such as phenology and plant architecture. A large-scale genome-wide association study (GWAS) was designed to investigate and dissect the genetic architecture of phenology affecting adaptation. It used 48 datasets from 4,680 spring wheat lines. For 8 years (2014–2021), these lines were evaluated for days to heading (DH) and maturity (DM) at three sites: Jabalpur, Ludhiana, and Samastipur (Pusa), which represent the three major Indian wheat-producing zones: the Central Zone (CZ), North-Western Plain Zone (NWPZ), and North-Eastern Plain Zone (NEPZ), respectively. Ludhiana had the highest mean DH of 103.8 days and DM of 148.6 days, whereas Jabalpur had the lowest mean DH of 77.7 days and DM of 121.6 days. We identified 119 markers significantly associated with DH and DM on chromosomes 5B (76), 2B (18), 7D (10), 4D (8), 5A (1), 6B (4), 7B (1), and 3D (1). Our results clearly indicated the importance of the photoperiod-associated gene (Ppd-B1) for adaptation to the NWPZ and the Vrn-B1 gene for adaptation to the NEPZ and CZ. A maximum variation of 21.1 and 14% was explained by markers 2B_56134146 and 5B_574145576 linked to the Ppd-B1 and Vrn-B1 genes, respectively, indicating their significant role in regulating DH and DM. The results provide important insights into the genomic regions associated with the two phenological traits that influence adaptation to the major wheat-producing zones in India. [ABSTRACT FROM AUTHOR]

Accelerating breeding efforts for developing biofortified bread wheat varieties necessitates understanding the genetic control of grain zinc concentration (GZnC) and grain iron concentration (GFeC). Hence, the major objective of this study was to perform genome-wide association mapping to identify consistently significant genotyping-by-sequencing markers associated with GZnC and GFeC using a large panel of 5,585 breeding lines from the International Maize and Wheat Improvement Center. These lines were grown between 2018 and 2021 in an optimally irrigated environment at Obregon, Mexico, while some of them were also grown in a water-limiting drought-stressed environment and a space-limiting small plot environment and evaluated for GZnC and GFeC. The lines showed a large and continuous variation for GZnC ranging from 27 to 74.5 ppm and GFeC ranging from 27 to 53.4 ppm. We performed 742,113 marker-traits association tests in 73 datasets and identified 141 markers consistently associated with GZnC and GFeC in three or more datasets, which were located on all wheat chromosomes except 3A and 7D. Among them, 29 markers were associated with both GZnC and GFeC, indicating a shared genetic basis for these micronutrients and the possibility of simultaneously improving both. In addition, several significant GZnC and GFeC associated markers were common across the irrigated, water-limiting drought-stressed, and space-limiting small plots environments, thereby indicating the feasibility of indirect selection for these micronutrients in either of these environments. Moreover, the many significant markers identified had minor effects on GZnC and GFeC, suggesting a quantitative genetic control of these traits. Our findings provide important insights into the complex genetic basis of GZnC and GFeC in bread wheat while implying limited prospects for marker-assisted selection and the need for using genomic selection. [ABSTRACT FROM AUTHOR]

Wheat (Triticum aestivum L.) is one of the major staples in Nepal providing the bulk of food calories and at least 30% of Fe and Zn intake and 20% of dietary energy and protein consumption; thus, it is essential to improve its nutritional quality. To select high-yielding genotypes with elevated grain zinc and iron concentration, the sixth, seventh, eighth, and ninth HarvestPlus Yield Trials (HPYTs) were conducted across diverse locations in Nepal for four consecutive years: 2015–16, 2016–17, 2017–18, and 2018–19, using 47 biofortified and 3 non-biofortified CIMMYT-bred, bread wheat genotypes: Baj#1, Kachu#1, and WK1204 (local check). Genotypic and spatial variations were found in agro-morphological traits; grain yield and its components; and the grain zinc and iron concentration of tested genotypes. Grain zinc concentration was highest in Khumaltar and lowest in Kabre. Likewise, grain iron concentration was highest in Doti and lowest in Surkhet. Most of the biofortified genotypes were superior for grain yield and for grain zinc and iron concentration to the non-biofortified checks. Combined analyses across environments showed moderate to high heritability for both Zn (0.48–0.81) and Fe (0.46–0.79) except a low heritability for Fe observed for 7th HPYT (0.15). Grain yield was positively correlated with the number of tillers per m2, while negatively correlated with days to heading and maturity, grain iron, grain weight per spike, and thousand grain weight. The grain zinc and iron concentration were positively correlated, suggesting that the simultaneous improvement of both micronutrients is possible through wheat breeding. Extensive testing of CIMMYT derived high Zn wheat lines in Nepal led to the release of five biofortified wheat varieties in 2020 with superior yield, better disease resistance, and 30–40% increased grain Zn and adaptable to a range of wheat growing regions in the country – from the hotter lowland, or Terai, regions to the dryer mid- and high-elevation areas. [ABSTRACT FROM AUTHOR]

Spot blotch caused by the fungus Bipolaris sorokiniana poses a serious threat to bread wheat production in warm and humid wheat-growing regions of the world. Hence, the major objective of this study was to identify consistent genotyping-by-sequencing (GBS) markers associated with spot blotch resistance using genome-wide association mapping on a large set of 6,736 advanced bread wheat breeding lines from the International Maize and Wheat Improvement Center. These lines were phenotyped as seven panels at Agua Fria, Mexico between the 2013–2014 and 2019–2020 crop cycles. We identified 214 significant spot blotch associated GBS markers in all the panels, among which only 96 were significant in more than one panel, indicating a strong environmental effect on the trait and highlights the need for multiple phenotypic evaluations to identify lines with stable spot blotch resistance. The 96 consistent GBS markers were on chromosomes 1A, 1B, 1D, 2A, 3B, 4A, 5B, 5D, 6B, 7A, 7B, and 7D, including markers possibly linked to the Lr46 , Sb1 , Sb2 and Sb3 genes. We also report the association of the 2NS translocation from Aegilops ventricosa with spot blotch resistance in some environments. Moreover, the spot blotch favorable alleles at the 2NS translocation and two markers on chromosome 3BS (3B_2280114 and 3B_5601689) were associated with increased grain yield evaluated at several environments in Mexico and India, implying that selection for favorable alleles at these loci could enable simultaneous improvement for high grain yield and spot blotch resistance. Furthermore, a significant relationship between the percentage of favorable alleles in the lines and their spot blotch response was observed, which taken together with the multiple minor effect loci identified to be associated with spot blotch in this study, indicate quantitative genetic control of resistance. Overall, the results presented here have extended our knowledge on the genetic basis of spot blotch resistance in bread wheat and further efforts to improve genetic resistance to the disease are needed for reducing current and future losses under climate change. [ABSTRACT FROM AUTHOR]

Biofortification of cereal grains offers a lasting solution to combat micronutrient deficiency in developing countries where it poses developmental risks to children. Breeding efforts thus far have been directed toward increasing the grain concentrations of iron (Fe) and zinc (Zn) ions. Phytic acid (PA) chelates these metal ions, reducing their bioavailability in the digestive tract. We present a high-throughput assay for quantification of PA and its application in screening a breeding population. After extraction in 96-well megatiter plates, PA content was determined from the phosphate released after treatment with a commercially available phytase enzyme. In a set of 330 breeding lines of wheat grown in the field over 3 years as part of a HarvestPlus breeding program for high grain Fe and Zn, our assay unraveled variation for PA that ranged from 0.90 to 1.72% with a mean of 1.24%. PA content was not associated with grain yield. High yielding lines were further screened for low molar PA/Fe and PA/Zn ratios for increased metal ion bioavailability, demonstrating the utility of our assay. Genome-wide association study revealed 21 genetic associations, six of which were consistent across years. Five of these associations mapped to chromosomes 1A, 2A, 2D, 5A, and 7D. Additivity over four of these haplotypes accounted for an ∼10% reduction in PA. Our study demonstrates it is possible to scale up assays to directly select for low grain PA in forward breeding programs. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Iranian wheat landrace accessions (IWAs) were collected from country‐wide farm fields and market places in 1935 by a professor at the University of Tehran and shared with University of California at Davis, California. IWAs were further submitted to the genebank of International Maize and Wheat Improvement Center (CIMMYT), Mexico. 2,403 IWAs from CIMMYT's genebank were assayed by DArT‐seq technology to assess genetic diversity. No apparent ecogeographic patterns related to genetic diversity were detected, probably due to long‐term transport and frequent interchange of landraces among farmers. A multivariate clustering procedure combining genotypic and phenotypic information was used in selecting a core‐set, which represented 15% of the hexaploid wheat accessions included in this study. This subset captured an estimated 93% of rare (frequency <0.05) alleles. Multisite phenotypic data (India, Mexico) validated the ability of the core‐set in detecting useful variants. Potential donor accessions for multiple traits (disease resistance, zinc concentration) were identified from the core‐set for wheat‐breeding. This report illustrates a breeder friendly robust core‐set formulation strategy for harnessing the useful genetic variation stored in the genebanks. [ABSTRACT FROM AUTHOR]

Genomic breeding technologies offer new opportunities for grain yield (GY) improvement in common wheat. In this study, we have evaluated the potential of genomic selection (GS) in breeding for GY in wheat by modeling a large dataset of 48,562 GY observations from the International Maize and Wheat Improvement Center (CIMMYT), including 36 yield trials evaluated between 2012 and 2019 in Obregón, Sonora, Mexico. Our key objective was to determine the value that GS can add to the current three-stage yield testing strategy at CIMMYT, and we draw inferences from predictive modeling of GY using 420 different populations, environments, cycles, and model combinations. First, we evaluated the potential of genomic predictions for minimizing the number of replications and lines tested within a site and year and obtained mean prediction accuracies (PAs) of 0.56, 0.5, and 0.42 in Stages 1, 2, and 3 of yield testing, respectively. However, these PAs were similar to the mean pedigree-based PAs indicating that genomic relationships added no value to pedigree relationships in the yield testing stages, characterized by small family-sizes. Second, we evaluated genomic predictions for minimizing GY testing across stages/years in Obregón and observed mean PAs of 0.41, 0.31, and 0.37, respectively when GY in the full irrigation bed planting (FI BP), drought stress (DS), and late-sown heat stress environments were predicted across years using genotype × environment (G × E) interaction models. Third, we evaluated genomic predictions for minimizing the number of yield testing environments and observed that in Stage 2, the FI BP, full irrigation flat planting and early-sown heat stress environments (mean PA of 0.37 ± 0.12) and the reduced irrigation and DS environments (mean PA of 0.45 ± 0.07) had moderate predictabilities among them. However, in both predictions across years and environments, the PAs were inconsistent across years and the G × E models had no advantage over the baseline model with environment and line effects. Overall, our results provide excellent insights into the predictability of a quantitative trait like GY and will have important implications on the future design of GS for GY in wheat breeding programs globally. [ABSTRACT FROM AUTHOR]

More 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. [ABSTRACT FROM AUTHOR]

Race Ug99 of the fungus Puccinia graminis tritici that causes stem or black rust disease on wheat was first detected in Uganda in 1998. Seven races belonging to the Ug99 lineage are now known and have spread to various wheat-growing countries in the eastern African highlands, as well as Zimbabwe, South Africa, Sudan, Yemen, and Iran. Because of the susceptibility of 90%% of the wheat varieties grown worldwide, the Ug99 group of races was recognized as a major threat to wheat production and food security. Its spread, either wind-mediated or human-aided, to other countries in Africa, Asia, and beyond is evident. Screening in Kenya and Ethiopia has identified a low frequency of resistant wheat varieties and breeding materials. Identification and transfer of new sources of race-specific resistance from various wheat relatives is underway to enhance the diversity of resistance. Although new Ug99-resistant varieties that yield more than current popular varieties are being released and promoted, major efforts are required to displace current Ug99 susceptible varieties with varieties that have diverse race-specific or durable resistance and mitigate the Ug99 threat. [ABSTRACT FROM AUTHOR]

Genetic biofortification through breeding offers sustainable solution to the micronutrient malnutrition problems in the target countries. Great progress has been made in the past decade in transferring alleles for high-zinc (Zn) and iron (Fe) from diverse genetic resources into elite wheat breeding lines. However, the major challenge is to maintain simultaneous and high rates of genetic gains for grain yield and grain Zn to meet the food and nutritional security demands through the continuous delivery of biofortified varieties that are competitive to replace non-biofortified varieties successfully. Although a few intermediate effect QTL regions are identified for grain Zn, both yield and Zn content are quantitatively inherited. Increased breeding efforts and new approaches are therefore required to combine them in high frequency in CIMMYT's elite germplasm, ensuring that Zn levels are steadily increased to the required levels across the CIMMYT breeding pipelines. The addition of Zn as a core-trait will be achieved through significant acceleration in the breeding cycle, expanding population sizes, extensive Zn phenotyping, yield testing, phenotyping for biotic and abiotic stresses, molecular-assisted selection and genomic selection. While continuing to increase agronomic performance, high Zn alleles has been added as a core-trait. Eventually Zn content will be increased in the elite lines annually along with the frequency of elite lines with high yield and other agronomic traits that have potential to be released by partners. A genomics assisted "rapid cycle recurrent selection" scheme achieved through rapid generation advancement approaches are expected to enable CIMMYT wheat breeding program to mainstream grain Zn in the majority of elite lines in about 10 years. • Significant genetic variation has been observed for grain Fe and Zn among elite bread wheat lines. • Speed breeding would enable rapid cycling of high Zn and high yield parents faster. • Genomic selection combined with selection index would advance genetic gains for complex traits. • CIMMYT wheat breeding program to mainstream grain Zn in all the elite lines in about 10 years. [ABSTRACT FROM AUTHOR]

Bread wheat can be used to make different products thanks to the presence of gluten, a protein network that confers unique visco-elastic properties to wheat doughs. Gluten is composed by gliadins and glutenins. The glutenins can be further divided into high and low-molecular-weight glutenins (HMWGs and LMWGs, respectively) and are encoded by Glu-1 and Glu-3 loci. The variability of these genes is associated with differences in quality. Because of this, the identification of novel glutenin alleles is still an important target. In this study, 57 haplotypes or glutenin combinations were registered among a set of 158 Iranian landraces and five novel HMWGs alleles were identified. The landraces were also characterized for several quality traits, including gluten quality, which allowed to associate the different glutenin alleles with low or high quality. Other quality traits examined were iron, zinc, and phytate contents, which are intimately related with the nutritional quality. Important variation for these components was found as well as for the phytate:iron/zinc molar ratios (related to the potential bioavailability of these important micronutrients). The landraces identified in the present study (some of them combining high gluten quality with low phytate:zinc values) could be a useful resource for breeders who aim to improve the wheat end-use quality and especially the content of zinc and its relative bioavailability. [ABSTRACT FROM AUTHOR]

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 effect 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. [ABSTRACT FROM AUTHOR]

Durum wheat accounts for more than 50% of the total wheat-growing area in the Mediterranean region, where is used for the preparation of diverse food products, including pasta and bread. The effects of drought and heat stresses on grain morphology, grain composition (protein, iron and zinc micronutrients), processing and pasta and bread-making quality in durum wheat varieties were analyzed. The results revealed significant differences among the genotypes, as well as unique responses to the environmental stresses. Micronutrients concentration (iron and zinc), processing and pasta-making quality was favored by drought but not by heat stress. Overall, the durum wheat lines showed inferior values for bread volume compared to the bread-wheat checks. However, some durum genotypes in specific environment had almost the same performance. To develop durum wheat cultivars with similar bread-making quality to that of bread wheat, it is necessary to achieve a better balance of tenacity and extensibility. The development of durum lines with good bread-making quality could increase the commercial value of this crop. [ABSTRACT FROM AUTHOR]

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. • Wheat quality and glutenin's data of 4623 grain samples derived from 2550 genotypes was used. • Gluten strength was the trait more strongly influenced by glutenin variations. • Several alleles were associated to good or bad quality, respectively. • This is probably the largest study ever done about glutenins effects on quality. [ABSTRACT FROM AUTHOR]

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. [Display omitted] • A genetically biofortified wheat line had higher contents of zinc and iron in grain. • The locations of the minerals were similar in the biofortified and control lines. • Zinc is more highly concentrated in the embryo including the axis. • The contents of Fe and Zn were correlated with P contents. • Hence the bioavailability of the minerals is unlikely to differ between the lines. [ABSTRACT FROM AUTHOR]

Bread wheat ( T. aestivum) is one of the world's most widely consumed cereals. Since micronutrient deficiencies are becoming more common among people who primarily depend upon cereal-based diets, a need for better-quality wheat varieties has been felt. An association panel of 154 T. aestivum lines was evaluated for the following quality traits: grain appearance (GA) score, grain hardness (GH), phenol reaction (PR) score, protein percent, sodium dodecyl sulfate (SDS) sedimentation value, and test weight (TWt). In addition, the panel was also phenotyped for grain yield and related traits such as days to heading, days to maturity, plant height, and thousand kernel weight for the year 2017-18 at the Borlaug Institute for South Asia (BISA) Ludhiana and Jabalpur sites. We performed a genome-wide association analysis on this panel using 18,351 genotyping-by-sequencing (GBS) markers to find marker-trait associations for quality and grain yield-related traits. We detected 55 single nucleotide polymorphism (SNP) marker trait associations (MTAs) for quality-related traits on chromosomes 7B (10), 1A (9), 2A (8), 3B (6), 2B (5), 7A (4), and 1B (3), with 3A, 4A, and 6D, having two and the rest, 4B, 5A, 5B, and 1D, having one each. Additionally, 20 SNP MTAs were detected for yield-related traits based on a field experiment conducted in Ludhiana on 7D (4) and 4D (3) chromosomes, while 44 SNP MTAs were reported for Jabalpur on chromosomes 2D (6), 7A (5), 2A (4), and 4A (4). Utilizing these loci in marker-assisted selection will benefit from further validation studies for these loci to improve hexaploid wheat for better yield and grain quality., Competing Interests: Author UK was employed by the company Astralyan Agro OPC Pvt. Ltd. The remaining authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2024 Vishwakarma, Bhati, Kumar, Singh, Kumar, Govindan, Mavi, Thiyagarajan, Dhar and Joshi.)