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Maize stalk rot reduces grain yield and quality. Information about the genetics of resistance to maize stalk rot could help breeders design effective breeding strategies for the trait. Genomic prediction may be a more effective breeding strategy for stalk-rot resistance than marker-assisted selection. We performed a genome-wide association study (GWAS) and genomic prediction of resistance in testcross hybrids of 677 inbred lines from the Tuxpeño and non-Tuxpeño heterotic pools grown in three environments and genotyped with 200,681 single-nucleotide polymorphisms (SNPs). Eighteen SNPs associated with stalk rot shared genomic regions with gene families previously associated with plant biotic and abiotic responses. More favorable SNP haplotypes traced to tropical than to temperate progenitors of the inbred lines. Incorporating genotype-by-environment (G × E) interaction increased genomic prediction accuracy.

Abstract Fostering a culture of continuous improvement through regular monitoring of genetic trends in breeding pipelines is essential to improve efficiency and increase accountability. This is the first global study to estimate genetic trends across the International Maize and Wheat Improvement Center (CIMMYT) tropical maize breeding pipelines in eastern and southern Africa (ESA), South Asia, and Latin America over the past decade. Data from a total of 4152 advanced breeding trials and 34,813 entries, conducted at 1331 locations in 28 countries globally, were used for this study. Genetic trends for grain yield reached up to 138 kg ha−1 yr−1 in ESA, 118 kg ha−1 yr−1 South Asia and 143 kg ha−1 yr−1 in Latin America. Genetic trend was, in part, related to the extent of deployment of new breeding tools in each pipeline, strength of an extensive phenotyping network, and funding stability. Over the past decade, CIMMYT’s breeding pipelines have significantly evolved, incorporating new tools/technologies to increase selection accuracy and intensity, while reducing cycle time. The first pipeline, Eastern Africa Product Profile 1a (EA-PP1a), to implement marker-assisted forward-breeding for resistance to key diseases, coupled with rapid-cycle genomic selection for drought, recorded a genetic trend of 2.46% per year highlighting the potential for deploying new tools/technologies to increase genetic gain.

The two most important activities in maize breeding are the development of inbred lines with high values of general combining ability (GCA) and specific combining ability (SCA), and the identification of hybrids with high yield potentials. Genomic selection (GS) is a promising genomic tool to perform selection on the untested breeding material based on the genomic estimated breeding values estimated from the genomic prediction (GP). In this study, GP analyses were carried out to estimate the performance of hybrids, GCA, and SCA for grain yield (GY) in three maize line-by-tester trials, where all the material was phenotyped in 10 to 11 multiple-location trials and genotyped with a mid-density molecular marker platform. Results showed that the prediction abilities for the performance of hybrids ranged from 0.59 to 0.81 across all trials in the model including the additive effect of lines and testers. In the model including both additive and non-additive effects, the prediction abilities for the performance of hybrids were improved and ranged from 0.64 to 0.86 across all trials. The prediction abilities of the GCA for GY were low, ranging between − 0.14 and 0.13 across all trials in the model including only inbred lines; the prediction abilities of the GCA for GY were improved and ranged from 0.49 to 0.55 across all trials in the model including both inbred lines and testers, while the prediction abilities of the SCA for GY were negative across all trials. The prediction abilities for GY between testers varied from − 0.66 to 0.82; the performance of hybrids between testers is difficult to predict. GS offers the opportunity to predict the performance of new hybrids and the GCA of new inbred lines based on the molecular marker information, the total breeding cost could be reduced dramatically by phenotyping fewer multiple-location trials.

Zinc (Zn) deficiency is a major risk factor for human health, affecting about 30% of the world’s population. To study the potential of genomic selection (GS) for maize with increased Zn concentration, an association panel and two doubled haploid (DH) populations were evaluated in three environments. Three genomic prediction models, M (M1: Environment + Line, M2: Environment + Line + Genomic, and M3: Environment + Line + Genomic + Genomic x Environment) incorporating main effects (lines and genomic) and the interaction between genomic and environment (G x E) were assessed to estimate the prediction ability (rMP) for each model. Two distinct cross-validation (CV) schemes simulating two genomic prediction breeding scenarios were used. CV1 predicts the performance of newly developed lines, whereas CV2 predicts the performance of lines tested in sparse multi-location trials. Predictions for Zn in CV1 ranged from -0.01 to 0.56 for DH1, 0.04 to 0.50 for DH2 and -0.001 to 0.47 for the association panel. For CV2, rMP values ranged from 0.67 to 0.71 for DH1, 0.40 to 0.56 for DH2 and 0.64 to 0.72 for the association panel. The genomic prediction model which included G x E had the highest average rMP for both CV1 (0.39 and 0.44) and CV2 (0.71 and 0.51) for the association panel and DH2 population, respectively. These results suggest that GS has potential to accelerate breeding for enhanced kernel Zn concentration by facilitating selection of superior genotypes.

CIMMYT maize lines (CMLs), which represent the tropical maize germplasm, are freely available worldwide. All currently released 615 CMLs and fourteen temperate maize inbred lines were genotyped with 180 kompetitive allele-specific PCR single nucleotide polymorphisms to develop a reference fingerprinting SNP dataset that can be used to perform quality control (QC) and genetic diversity analyses. The QC analysis identified 25 CMLs with purity, identity, or mislabeling issues. Further field observation, purification, and re-genotyping of these CMLs are required. The reference fingerprinting SNP dataset was developed for all of the currently released CMLs with 152 high-quality SNPs. The results of principal component analysis and average genetic distances between subgroups showed a clear genetic divergence between temperate and tropical maize, whereas the three tropical subgroups partially overlapped with one another. More than 99% of the pairs of CMLs had genetic distances greater than 0.30, showing their high genetic diversity, and most CMLs are distantly related. The heterotic patterns, estimated with the molecular markers, are consistent with those estimated using pedigree information in two major maize breeding programs at CIMMYT. These research findings are helpful for ensuring the regeneration and distribution of the true CMLs, via QC analysis, and for facilitating the effective utilization of the CMLs, globally.

Common rust is one of the major foliar diseases in maize, leading to significant grain yield losses and poor grain quality. To dissect the genetic architecture of common rust resistance, a genome-wide association study (GWAS) panel and a bi-parental doubled haploid (DH) population, DH1, were used to perform GWAS and linkage mapping analyses. The GWAS results revealed six single-nucleotide polymorphisms (SNPs) significantly associated with quantitative resistance of common rust at a very stringent threshold of P-value 3.70 × 10–6 at bins 1.05, 1.10, 3.04, 3.05, 4.08, and 10.04. Linkage mapping identified five quantitative trait loci (QTL) at bins 1.03, 2.06, 4.08, 7.03, and 9.00. The phenotypic variation explained (PVE) value of each QTL ranged from 5.40 to 12.45%, accounting for the total PVE value of 40.67%. Joint GWAS and linkage mapping analyses identified a stable genomic region located at bin 4.08. Five significant SNPs were only identified by GWAS, and four QTL were only detected by linkage mapping. The significantly associated SNP of S10_95231291 detected in the GWAS analysis was first reported. The linkage mapping analysis detected two new QTL on chromosomes 7 and 10. The major QTL on chromosome 7 in the region between 144,567,253 and 149,717,562 bp had the largest PVE value of 12.45%. Four candidate genes of GRMZM2G328500, GRMZM2G162250, GRMZM2G114893, and GRMZM2G138949 were identified, which played important roles in the response of stress resilience and the regulation of plant growth and development. Genomic prediction (GP) accuracies observed in the GWAS panel and DH1 population were 0.61 and 0.51, respectively. This study provided new insight into the genetic architecture of quantitative resistance of common rust. In tropical maize, common rust could be improved by pyramiding the new sources of quantitative resistance through marker-assisted selection (MAS) or genomic selection (GS), rather than the implementation of MAS for the single dominant race-specific resistance gene.

Enriching of kernel zinc (Zn) concentration in maize is one of the most effective ways to solve the problem of Zn deficiency in low and middle income countries where maize is the major staple food, and 17% of the global population is affected with Zn deficiency. Genomic selection (GS) has shown to be an effective approach to accelerate genetic gains in plant breeding. In the present study, an association-mapping panel and two maize double-haploid (DH) populations, both genotyped with genotyping-by-sequencing (GBS) and repeat amplification sequencing (rAmpSeq) markers, were used to estimate the genomic prediction accuracy of kernel Zn concentration in maize. Results showed that the prediction accuracy of two DH populations was higher than that of the association mapping population using the same set of markers. The prediction accuracy estimated with the GBS markers was significantly higher than that estimated with the rAmpSeq markers in the same population. The maximum prediction accuracy with minimum standard error was observed when half of the genotypes were included in the training set and 3,000 and 500 markers were used for prediction in the association mapping panel and the DH populations, respectively. Appropriate levels of minor allele frequency and missing rate should be considered and selected to achieve good prediction accuracy and reduce the computation burden by balancing the number of markers and marker quality. Training set development with broad phenotypic variation is possible to improve prediction accuracy. The transferability of the GS models across populations was assessed, the prediction accuracies in a few pairwise populations were above or close to 0.20, which indicates the prediction accuracies across years and populations have to be assessed in a larger breeding dataset with closer relationship between the training and prediction sets in further studies. GS outperformed MAS (marker-assisted-selection) on predicting the kernel Zn concentration in maize, the decision of a breeding strategy to implement GS individually or to implement MAS and GS stepwise for improving kernel Zn concentration in maize requires further research. Results of this study provide valuable information for understanding how to implement GS for improving kernel Zn concentration in maize.

Maize is a major source of food security and economic development in sub-Saharan Africa (SSA), Latin America, and the Caribbean, and is among the top three cereal crops in Asia. Yet, maize is deficient in certain essential amino acids, vitamins, and minerals. Biofortified maize cultivars enriched with essential minerals and vitamins could be particularly impactful in rural areas with limited access to diversified diet, dietary supplements, and fortified foods. Significant progress has been made in developing, testing, and deploying maize cultivars biofortified with quality protein maize (QPM), provitamin A, and kernel zinc. In this review, we outline the status and prospects of developing nutritionally enriched maize by successfully harnessing conventional and molecular marker-assisted breeding, highlighting the need for intensification of efforts to create greater impacts on malnutrition in maize-consuming populations, especially in the low- and middle-income countries. Molecular marker-assisted selection methods are particularly useful for improving nutritional traits since conventional breeding methods are relatively constrained by the cost and throughput of nutritional trait phenotyping.

Vitamin A deficiency (VAD) is a public health issue worldwide. Provitamin A (PVA) biofortified maize serves as an alternative to help combat VAD. Breeding efforts to develop maize varieties with high PVA carotenoid content combine molecular and phenotypic selection strategies. The phenotypic assessment of carotenoids is currently done using liquid chromatography, a precise but time- and resource-consuming methodology. Using near-infrared spectroscopy (NIRS) could increase the breeding efficiency. This study used ultra-performance liquid chromatography (UPLC) data from 1857 tropical maize genotypes as a training set and NIRS data to do an independent test of a set of 650 genotypes to predict PVA carotenoids using Bayesian and modified partial least square (MPLS) regression models. Both regression methods produced similar prediction accuracies for the total carotenoids (r2 = 0.75), lutein (r2 = 0.55), zeaxanthin (r2= 0.61), β-carotene (r2 = 0.22) and β-cryptoxanthin (BCX) (r2 = 0.57). These results demonstrate that Bayesian and MPLS regression of BCX on NIRS data can be used to predict BCX content, the current focus on PVA enhancement, and thus offers opportunities for high-throughput phenotyping at a low cost, especially in the early stages of PVA maize breeding pipeline when many genotypes must be screened.

Genotype, environmental temperature, and agronomic management of parents influence seed yield in three-way cross hybrid maize seed production. The objective of this research was to generate information on the seed production of six three-way cross hybrids and their progenitors, adapted to tropical lowlands. Data on days to—and duration of—flowering, distance to spike and stigmas, and seed yield of five female single crosses and five male inbred lines were recorded for different combinations of four planting densities and four sowing dates in Mexico. The effect of planting density was not significant. The male inbred line T10 was the earliest and highest seed yield and T31 the latest, occupying second place in yield. The single crosses T32/T10 and T13/T14 were the earliest and had the highest effective seed yield. At the earliest sowing date, the females were later in their flowering, accumulated fewer growing degree days (GDD), and obtained higher yields since the grain-filling period coincided with hot days and cool nights. To achieve greater floral synchronization and therefore greater production of hybrid seed, differential planting dates for parents are recommended based on information from the accumulated GDD of each parent. The three-way cross hybrids were classified according to the expected seed yield of the females and the complexity in the synchronization of flowering of their parents.

Medium and small seed companies require information on maize parental seed production to make rational choices on what germplasm to introduce into their seed and breeding pipelines. The objective of this study was to generate public data on the appropriate hybrid seed production information of five female lines (T21, T26, T28, T33 and T38) and one line (T10) as a male hybrid parent. The effect of female and male sowing date and sowing density on hybrid seed production characteristics including flowering time, flowering duration, distance between tassel and stigmas and effective seed yield was determined. Recommendations to stagger male and female sowing to achieve flowering synchrony for the hybrid parents were complemented with data of seedling growth stages. The results were similar for T21, T26, T28, and T38; sowing the female and T10 on a row proportion of 4 females:1 male, and on a second date, when the coleoptiles of the female plants are halfway to emerge, sowing the missing male row to complete the 4:2 ratio. T33 is a late flowering line, therefore it is desirable to sow T33 first and T10 five days later, or when the T33 coleoptile begins to emerge. Plant densities did not cause differences in most plant characteristics. Line T21 showed good female parental traits. While this study provides a knowledge framework for seed production technology for these single cross hybrids, data specific to seed production regions need to be generated by seed companies to define the best regimes for hybrid production.

Genetic improvement of maize with elevated levels of zinc (Zn) can reduce Zn deficiency among populations who rely on maize as a staple. Inbred lines of quality protein maize (QPM) and non-QPM with elevated Zn levels in the kernel have been identified. However, information about the optimal strategy to utilize the germplasm in breeding for high-Zn concentration is lacking. As a preliminary step, this study was conducted to ascertain the potential of QPM, non-QPM, or a combination of QPM and non-QPM hybrids for attaining desirable Zn concentration. Twenty elite inbreds, 10 QPM and 10 non-QPM, were crossed according to a modified mating design to generate hybrids, which were evaluated in four environments in Mexico during 2015 and 2016 in order to evaluate their merits as parents of hybrids. The highest mean values of Zn were observed when high-Zn QPM lines were crossed with high-Zn non-QPM lines. Hybrids with high Zn and grain yield were identified. General combining ability (GCA) effects for Zn concentration were more preponderant than specific combining ability (SCA) effects, suggesting the importance of additive gene action for the inheritance of Zn.

High throughput phenotyping technologies are lagging behind modern marker technology impairing the use of secondary traits to increase genetic gains in plant breeding. We aimed to assess whether the combined use of hyperspectral data with modern marker technology could be used to improve across location pre-harvest yield predictions using different statistical models. A maize bi-parental doubled haploid (DH) population derived from F1, which consisted of 97 lines was evaluated in testcross combination under heat stress as well as combined heat and drought stress during the 2014 and 2016 summer season in Ciudad Obregon, Sonora, Mexico (27°20" N, 109°54" W, 38 m asl). Full hyperspectral data, indicative of crop physiological processes at the canopy level, was repeatedly measured throughout the grain filling period and related to grain yield. Partial least squares regression (PLSR), random forest (RF), ridge regression (RR) and Bayesian ridge regression (BayesB) were used to assess prediction accuracies on grain yield within (two-fold cross-validation) and across environments (leave-one-environment-out-cross-validation) using molecular markers (M), hyperspectral data (H) and the combination of both (HM). Highest prediction accuracy for grain yield averaged across within and across location predictions (rGP) were obtained for BayesB followed by RR, RF and PLSR. The combined use of hyperspectral and molecular marker data as input factor on average had higher predictions for grain yield than hyperspectral data or molecular marker data alone. The highest prediction accuracy for grain yield across environments was measured for BayesB when molecular marker data and hyperspectral data were used as input factors, while the highest within environment prediction was obtained when BayesB was used in combination with hyperspectral data. It is discussed how the combined use of hyperspectral data with molecular marker technology could be used to introduce physiological genomic estimated breeding values (PGEBV) as a pre-harvest decision support tool to select genetically superior lines.