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Northern corn leaf blight (NCLB), caused by Setosphaeria turcica, is a major fungal disease affecting maize production in sub-Saharan Africa. Utilizing host plant resistance to mitigate yield losses associated with NCLB can serve as a cost-effective strategy. In this study, we conducted a high-resolution genome-wide association study (GWAS) in an association mapping panel and linkage mapping with three doubled haploid (DH) and three F3 populations of tropical maize. These populations were phenotyped for NCLB resistance across six hotspot environments in Kenya. Across environments and genotypes, NCLB scores ranged from 2.12 to 5.17 (on a scale of 1–9). NCLB disease severity scores exhibited significant genotypic variance and moderate-to-high heritability. From the six biparental populations, 23 quantitative trait loci (QTLs) were identified, each explaining between 2.7% and 15.8% of the observed phenotypic variance. Collectively, the detected QTLs explained 34.28%, 51.37%, 41.12%, 12.46%, 12.11%, and 14.66% of the total phenotypic variance in DH populations 1, 2, and 3 and F3 populations 4, 5, and 6, respectively. GWAS, using 337,110 high-quality single nucleotide polymorphisms (SNPs), identified 15 marker–trait associations and several putative candidate genes linked to NCLB resistance in maize. Joint linkage association mapping (JLAM) identified 37 QTLs for NCLB resistance. Using linkage mapping, JLAM, and GWAS, several QTLs were identified within the genomic region spanning 4 to 15 Mbp on chromosome 2. This genomic region represents a promising target for enhancing NCLB resistance via marker-assisted breeding. Genome-wide predictions revealed moderate correlations with mean values of 0.45, 0.44, 0.55, and 0.42 for within GWAS panel, DH pop1, DH pop2, and DH pop3, respectively. Prediction by incorporating marker-by-environment interactions did not show much improvement. Overall, our findings indicate that NCLB resistance is quantitative in nature and is controlled by few major-effect and many minor-effect QTLs. We conclude that genomic regions consistently detected across mapping approaches and populations should be prioritized for improving NCLB resistance, while genome-wide prediction results can help incorporate both major- and minor-effect genes. This study contributes to a deeper understanding of the genetic and molecular mechanisms driving maize resistance to NCLB. more...

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 more...

Genomic selection (GS) is a predictive methodology that is revolutionizing plant and animal breeding. However, the practical application of the GS methodology is challenging since a successful implementation requires a good identification of the best lines. For this reason, some approaches have been proposed to be able to select the top (or bottom) lines with more Precision. Despite the varying popularity of methods, with some being notably more efficient than others, this paper delves into the fundamentals of these techniques. We used five models/methods: (1) RC, known as the Bayesian Best Linear Unbiased Predictor (GBLUP); (2) R, which is like RC but uses a threshold; (3) RO, Regression Optimum, that leverages the RC model in its training process to fine-tune the threshold; (4) B, Threshold Bayesian Probit Binary model (TGBLUP) with a threshold of 0.5 to classify the cultivars as top or non-top; (5) BO is the TGBLUP but the threshold used is an optimal probability threshold that guarantees similar Sensitivity and Specificity. We also present a benchmark comparison of existing approaches for selecting the top (or bottom) performers, utilizing five real datasets for comprehensive analysis. For methods that necessitate a rigorous tuning process, we suggest a streamlined tuning approach that significantly decreases implementation time without notably compromising performance. Our analysis revealed that the regression optimal (RO) method outperformed other models across the five real datasets, achieving superior results in terms of the F1 score. Specifically, RO was more effective than models R, B, RC, and BO by 60.87, 42.37, 17.63, and 9.62%, respectively. When looking at the Kappa coefficient, the RO model was better than models B, BO, R, and RC by 37.46, 36.21, 52.18, and 3.95%, respectively. In terms of Sensitivity, the RO model outperformed models B, R, and RC by 145.74, 250.41, and 86.20, respectively. The second-best model was the model BO. It is important to point out that in the first stage, the BO and RO approaches train a classification and regression model, respectively, to classify the lines as the top (bottom) or not the top (not the bottom). However, both the BO and RO approaches optimize a threshold in the second stage to perform the classification of the lines that minimize the difference between the Sensitivity and Specificity. The BO and RO methods are superior for the selection of the top (or bottom) lines. For this reason, we encourage breeders to adopt these approaches to increase genetic gain in plant breeding programs. more...

Three F2-derived biparental doubled haploid (DH) maize populations were generated for genetic mapping of resistance to common rust. Each of the three populations has the same susceptible parent, but a different resistance donor parent. Population 1 and 3 consist of 320 lines each, population 2 consists of 260 lines. The DH lines were evaluated for their susceptibility to common rust in two years and with two replications in each year. For phenotyping, a visual score (VS) for susceptibility was assigned. Additionally, unmanned aerial vehicle (UAV) derived multispectral and thermal infrared data was recorded and combined in different vegetation indices (“remote sensing”, RS). The DH lines were genotyped with the DarTseq method, to obtain data on single nucleotide polymorphisms (SNPs). After quality control, 9051 markers remained. Missing values were “imputed” by the empirical mean of the marker scores of the respective locus. We used the data for comparison of genome-wide association studies and genomic prediction when based on different phenotyping methods, that is either VS or RS data. The data may be interesting for reuse for instance for benchmarking genomic prediction models, for phytopathological studies addressing common rust, or for specifications of vegetation indices. more...

Abstract Bread wheat (Triticum aestivum L.) is a globally important food crop, which was domesticated about 8–10,000 years ago. Bread wheat is an allopolyploid, and it evolved from two hybridization events of three species. To widen the genetic base in breeding, bread wheat has been re‐synthesized by crossing durum wheat (Triticum turgidum ssp. durum) and goat grass (Aegilops tauschii Coss), leading to so‐called synthetic hexaploid wheat (SHW). We applied the quantitative genetics tools of “hybrid prediction”—originally developed for the prediction of wheat hybrids generated from different heterotic groups — to a situation of allopolyploidization. Our use‐case predicts the phenotypes of SHW for three quantitatively inherited global wheat diseases, namely tan spot (TS), septoria nodorum blotch (SNB), and spot blotch (SB). Our results revealed prediction abilities comparable to studies in ‘traditional’ elite or hybrid wheat. Prediction abilities were highest using a marker model and performing random cross‐validation, predicting the performance of untested SHW (0.483 for SB to 0.730 for TS). When testing parents not necessarily used in SHW, combination prediction abilities were slightly lower (0.378 for SB to 0.718 for TS), yet still promising. Despite the limited phenotypic data, our results provide a general example for predictive models targeting an allopolyploidization event and a method that can guide the use of genetic resources available in gene banks. more...

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. more...

AbstractWhile several statistical machine learning methods have been developed and studied for assessing the genomic prediction (GP) accuracy of unobserved phenotypes in plant breeding research, few methods have linked genomics and phenomics (imaging). Deep learning (DL) neural networks have been developed to increase the GP accuracy of unobserved phenotypes while simultaneously accounting for the complexity of genotype–environment interaction (GE); however, unlike conventional GP models, DL has not been investigated for when genomics is linked with phenomics. In this study we used 2 wheat data sets (DS1 and DS2) to compare a novel DL method with conventional GP models. Models fitted for DS1 were GBLUP, gradient boosting machine (GBM), support vector regression (SVR) and the DL method. Results indicated that for 1 year, DL provided better GP accuracy than results obtained by the other models. However, GP accuracy obtained for other years indicated that the GBLUP model was slightly superior to the DL. DS2 is comprised only of genomic data from wheat lines tested for 3 years, 2 environments (drought and irrigated) and 2–4 traits. DS2 results showed that when predicting the irrigated environment with the drought environment, DL had higher accuracy than the GBLUP model in all analyzed traits and years. When predicting drought environment with information on the irrigated environment, the DL model and GBLUP model had similar accuracy. The DL method used in this study is novel and presents a strong degree of generalization as several modules can potentially be incorporated and concatenated to produce an output for a multi-input data structure. more...

Development and deployment of high-yielding maize varieties with native resistance to Fall armyworm (FAW), turcicum leaf blight (TLB), and gray leaf spot (GLS) infestation is critical for addressing the food insecurity in sub-Saharan Africa. The objectives of this study were to determine the inheritance of resistance for FAW, identity hybrids which in addition to FAW resistance, also show resistance to TLB and GLS, and investigate the usefulness of models based on general combining ability (GCA) and SNP markers in predicting the performance of new untested hybrids. Half-diallel mating scheme was used to generate 105 F1 hybrids from 15 parents and another 55 F1 hybrids from 11 parents. These were evaluated in two experiments, each with commercial checks in multiple locations under FAW artificial infestation and optimum management in Kenya. Under artificial FAW infestation, significant mean squares among hybrids and hybrids x environment were observed for most traits in both experiments, including at least one of the three assessments carried out for foliar damage caused by FAW. Interaction of GCA x environment and specific combining ability (SCA) x environment interactions were significant for all traits under FAW infestation and optimal conditions. Moderate to high heritability estimates were observed for GY under both management conditions. Correlation between GY and two of the three scorings (one and three weeks after infestation) for foliar damage caused by FAW were negative (-0.27 and -0.38) and significant. Positive and significant correlation (0.84) was observed between FAW-inflicted ear damage and the percentage of rotten ears. We identified many superior-performing hybrids compared to the best commercial checks for both GY and FAW resistance associated traits. Inbred lines CML312, CML567, CML488, DTPYC9-F46-1-2-1-2, CKDHL164288, CKDHL166062, and CLRCY039 had significant and positive GCA for GY (positive) and FAW resistance-associated traits (negative). CML567 was a parent in four of the top ten hybrids under optimum and FAW conditions. Both additive and non-additive gene action were important in the inheritance of FAW resistance. Both GCA and marker-based models showed high correlation with field performance, but marker-based models exhibited considerably higher correlation. The best performing hybrids identified in this study could be used as potential single cross testers in the development of three-way FAW resistance hybrids. Overall, our results provide insights that help breeders to design effective breeding strategies to develop FAW resistant hybrids that are high yielding under FAW and optimum conditions. more...

Abstract The success of genomic selection (GS) in breeding schemes relies on its ability to provide accurate predictions of unobserved lines at early stages. Multigeneration data provides opportunities to increase the training data size and thus, the likelihood of extracting useful information from ancestors to improve prediction accuracy. The genomic best linear unbiased predictions (GBLUPs) are performed by borrowing information through kinship relationships between individuals. Multigeneration data usually becomes heterogeneous with complex family relationship patterns that are increasingly entangled with each generation. Under these conditions, historical data may not be optimal for model training as the accuracy could be compromised. The sparse selection index (SSI) is a method for training set (TRN) optimization, in which training individuals provide predictions to some but not all predicted subjects. We added an additional trimming process to the original SSI (trimmed SSI) to remove less important training individuals for prediction. Using a large multigeneration (8 yr) wheat (Triticum aestivum L.) grain yield dataset (n = 68,836), we found increases in accuracy as more years are included in the TRN, with improvements of ∼0.05 in the GBLUP accuracy when using 5 yr of historical data relative to when using only 1 yr. The SSI method showed a small gain over the GBLUP accuracy but with an important reduction on the TRN size. These reduced TRNs were formed with a similar number of subjects from each training generation. Our results suggest that the SSI provides a more stable ranking of genotypes than the GBLUP as the TRN becomes larger. more...

AbstractLinking high-throughput environmental data (enviromics) to genomic prediction (GP) is a cost-effective strategy for increasing selection intensity under genotype-by-environment interactions (G × E). This study developed a data-driven approach based on Environment–Phenotype Association (EPA) aimed at recycling important G × E information from historical breeding data. EPA was developed in two applications: (1) scanning a secondary source of genetic variation, weighted from the shared reaction-norms of past-evaluated genotypes and (2) pinpointing weights of the similarity among trial-sites (locations), given the historical impact of each envirotyping data variable for a given site. These results were then used as a dimensionality reduction strategy, integrating historical data to feed multi-environment GP models, which led to the development of four new G × E kernels considering genomics, enviromics, and EPA outcomes. The wheat trial data used included 36 locations, 8 years, and three target populations of environments (TPEs) in India. Four prediction scenarios and six kernel models within/across TPEs were tested. Our results suggest that the conventional GBLUP, without enviromic data or when omitting EPA, is inefficient in predicting the performance of wheat lines in future years. Nevertheless, when EPA was introduced as an intermediary learning step to reduce the dimensionality of the G × E kernels while connecting phenotypic and environmental-wide variation, a significant enhancement of G × E prediction accuracy was evident. EPA revealed that the effect of seasonality makes strategies such as “covariable selection” unfeasible because G × E is year-germplasm specific. We propose that the EPA effectively serves as a “reinforcement learner” algorithm capable of uncovering the effect of seasonality over the reaction-norms, with the benefits of better forecasting the similarities between past and future trialing sites. EPA combines the benefits of dimensionality reduction while reducing the uncertainty of genotype-by-year predictions and increasing the resolution of GP for the genotype-specific level. more...

AbstractIn this study, we extend research on genomic prediction (GP) to polysomic polyploid plant species with the main objective to investigate single-trait (ST) and multitrait (MT) multienvironment (ME) models using field trial data from 3 locations in Sweden [Helgegården (HEL), Mosslunda (MOS), Umeå (UM)] over 2 years (2020, 2021) of 253 potato cultivars and breeding clones for 5 tuber weight traits and 2 tuber flesh quality characteristics. This research investigated the GP of 4 genome-based prediction models with genotype × environment interactions (GEs): (1) ST reaction norm model (M1M2M2M3M4M4M4M4,M3M2 more...

Fusarium ear rot (FER) is a destructive maize fungal disease worldwide. In this study, three tropical maize populations consisting of 874 inbred lines were used to perform genome-wide association study (GWAS) and genomic prediction (GP) analyses of FER resistance. Broad phenotypic variation and high heritability for FER were observed, although it was highly influenced by large genotype-by-environment interactions. In the 874 inbred lines, GWAS with general linear model (GLM) identified 3034 single-nucleotide polymorphisms (SNPs) significantly associated with FER resistance at the P-value threshold of 1 × 10−5, the average phenotypic variation explained (PVE) by these associations was 3% with a range from 2.33% to 6.92%, and 49 of these associations had PVE values greater than 5%. The GWAS analysis with mixed linear model (MLM) identified 19 significantly associated SNPs at the P-value threshold of 1 × 10−4, the average PVE of these associations was 1.60% with a range from 1.39% to 2.04%. Within each of the three populations, the number of significantly associated SNPs identified by GLM and MLM ranged from 25 to 41, and from 5 to 22, respectively. Overlapping SNP associations across populations were rare. A few stable genomic regions conferring FER resistance were identified, which located in bins 3.04/05, 7.02/04, 9.00/01, 9.04, 9.06/07, and 10.03/04. The genomic regions in bins 9.00/01 and 9.04 are new. GP produced moderate accuracies with genome-wide markers, and relatively high accuracies with SNP associations detected from GWAS. Moderate prediction accuracies were observed when the training and validation sets were closely related. These results implied that FER resistance in maize is controlled by minor QTL with small effects, and highly influenced by the genetic background of the populations studied. Genomic selection (GS) by incorporating SNP associations detected from GWAS is a promising tool for improving FER resistance in maize. more...

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. more...

“Sparse testing” refers to reduced multi-environment breeding trials in which not all genotypes of interest are grown in each environment. Using genomic-enabled prediction and a model embracing genotype × environment interaction (GE), the non-observed genotype-in-environment combinations can be predicted. Consequently, the overall costs can be reduced and the testing capacities can be increased. The accuracy of predicting the unobserved data depends on different factors including (1) how many genotypes overlap between environments, (2) in how many environments each genotype is grown, and (3) which prediction method is used. In this research, we studied the predictive ability obtained when using a fixed number of plots and different sparse testing designs. The considered designs included the extreme cases of (1) no overlap of genotypes between environments, and (2) complete overlap of the genotypes between environments. In the latter case, the prediction set fully consists of genotypes that have not been tested at all. Moreover, we gradually go from one extreme to the other considering (3) intermediates between the two previous cases with varying numbers of different or non-overlapping (NO)/overlapping (O) genotypes. The empirical study is built upon two different maize hybrid data sets consisting of different genotypes crossed to two different testers (T1 and T2) and each data set was analyzed separately. For each set, phenotypic records on yield from three different environments are available. Three different prediction models were implemented, two main effects models (M1 and M2), and a model (M3) including GE. The results showed that the genome-based model including GE (M3) captured more phenotypic variation than the models that did not include this component. Also, M3 provided higher prediction accuracy than models M1 and M2 for the different allocation scenarios. Reducing the size of the calibration sets decreased the prediction accuracy under all allocation designs with M3 being the less affected model; however, using the genome-enabled models (i.e., M2 and M3) the predictive ability is recovered when more genotypes are tested across environments. Our results indicate that a substantial part of the testing resources can be saved when using genome-based models including GE for optimizing sparse testing designs. more...

A combined multistage linear genomic selection index (CMLGSI) is a linear combination of phenotypic and genomic estimated breeding values useful for predicting the individual net genetic merit, which in turn is a linear combination of the true unobservable breeding values of the traits weighted by their respective economic values. The CMLGSI is a cost-saving strategy for improving multiple traits because the breeder does not need to measure all traits at each stage. The optimum (OCMLGSI) and decorrelated (DCMLGSI) indices are the main CMLGSIs. Whereas the OCMLGSI takes into consideration the index correlation values among stages, the DCMLGSI imposes the restriction that the index correlation values among stages be zero. Using real and simulated datasets, we compared the efficiency of both indices in a two-stage context. The criteria we applied to compare the efficiency of both indices were that the total selection response of each index must be lower than or equal to the single-stage combined linear genomic selection index (CLGSI) response and that the correlation of each index with the net genetic merit should be maximum. Using four different total proportions for the real dataset, the estimated total OCMLGSI and DCMLGSI responses explained 97.5% and 90%, respectively, of the estimated single-stage CLGSI selection response. In addition, at stage two, the estimated correlations of the OCMLGSI and the DCMLGSI with the net genetic merit were 0.84 and 0.63, respectively. We found similar results for the simulated datasets. Thus, we recommend using the OCMLGSI when performing multistage selection. more...

Machine learning methods such as multilayer perceptrons (MLP) and Convolutional Neural Networks (CNN) have emerged as promising methods for genomic prediction (GP). In this context, we assess the performance of MLP and CNN on regression and classification tasks in a case study with maize hybrids. The genomic information was provided to the MLP as a relationship matrix and to the CNN as “genomic images.” In the regression task, the machine learning models were compared along with GBLUP. Under the classification task, MLP and CNN were compared. In this case, the traits (plant height and grain yield) were discretized in such a way to create balanced (moderate selection intensity) and unbalanced (extreme selection intensity) datasets for further evaluations. An automatic hyperparameter search for MLP and CNN was performed, and the best models were reported. For both task types, several metrics were calculated under a validation scheme to assess the effect of the prediction method and other variables. Overall, MLP and CNN presented competitive results to GBLUP. Also, we bring new insights on automated machine learning for genomic prediction and its implications to plant breeding. more...

Abstract Genomic selection (GS) is a predictive methodology that trains statistical machine‐learning models with a reference population that is used to perform genome‐enabled predictions of new lines. In plant breeding, it has the potential to increase the speed and reduce the cost of selection. However, to optimize resources, sparse testing methods have been proposed. A common approach is to guarantee a proportion of nonoverlapping and overlapping lines allocated randomly in locations, that is, lines appearing in some locations but not in all. In this study we propose using incomplete block designs (IBD), principally, for the allocation of lines to locations in such a way that not all lines are observed in all locations. We compare this allocation with a random allocation of lines to locations guaranteeing that the lines are allocated to the same number of locations as under the IBD design. We implemented this benchmarking on several crop data sets under the Bayesian genomic best linear unbiased predictor (GBLUP) model, finding that allocation under the principle of IBD outperformed random allocation by between 1.4% and 26.5% across locations, traits, and data sets in terms of mean square error. Although a wide range of performance improvements were observed, our results provide evidence that using IBD for the allocation of lines to locations can help improve predictive performance compared with random allocation. This has the potential to be applied to large‐scale plant breeding programs. more...

We investigated increasing genetic gain for grain yield using early generation genomic selection (GS). A training set of 1,334 elite wheat breeding lines tested over three field seasons was used to generate Genomic Estimated Breeding Values (GEBVs) for grain yield under irrigated conditions applying markers and three different prediction methods: (1) Genomic Best Linear Unbiased Predictor (GBLUP), (2) GBLUP with the imputation of missing genotypic data by Ridge Regression BLUP (rrGBLUP_imp), and (3) Reproducing Kernel Hilbert Space (RKHS) a.k.a. Gaussian Kernel (GK). F2 GEBVs were generated for 1,924 individuals from 38 biparental cross populations between 21 parents selected from the training set. Results showed that F2 GEBVs from the different methods were not correlated. Experiment 1 consisted of selecting F2s with the highest average GEBVs and advancing them to form genomically selected bulks and make intercross populations aiming to combine favorable alleles for yield. F4:6 lines were derived from genomically selected bulks, intercrosses, and conventional breeding methods with similar numbers from each. Results of field-testing for Experiment 1 did not find any difference in yield with genomic compared to conventional selection. Experiment 2 compared the predictive ability of the different GEBV calculation methods in F2 using a set of single plant-derived F2:4 lines from randomly selected F2 plants. Grain yield results from Experiment 2 showed a significant positive correlation between observed yields of F2:4 lines and predicted yield GEBVs of F2 single plants from GK (the predictive ability of 0.248, P < 0.001) and GBLUP (0.195, P < 0.01) but no correlation with rrGBLUP_imp. Results demonstrate the potential for the application of GS in early generations of wheat breeding and the importance of using the appropriate statistical model for GEBV calculation, which may not be the same as the best model for inbreds. more...

Abstract Sparse testing in genome‐enabled prediction in plant breeding can be emulated throughout different line allocations where some lines are observed in all environments (overlap) and others are observed in only one environment (nonoverlap). We studied three general cases of the composition of the sparse testing allocation design for genome‐enabled prediction of wheat (Triticum aestivum L.) breeding: (a) completely nonoverlapping wheat lines in environments, (b) completely overlapping wheat lines in all environments, and (c) a proportion of nonoverlapping/overlapping wheat lines allocated in the environments. We also studied several cases in which the size of the testing population was systematically decreased. The study used three extensive wheat data sets (W1, W2, and W3). Three different genome‐enabled prediction models (M1–M3) were used to study the effect of the sparse testing in terms of the genomic prediction accuracy. Model M1 included only main effects of environments and lines; M2 included main effects of environments, lines, and genomic effects; whereas the remaining model (M3) also incorporated the genomic × environment interaction (GE). The results show that the GE component of the genome‐based model M3 captures a larger genetic variability than the main genomic effects term from models M1 and M2. In addition, model M3 provides higher prediction accuracy than models M1 and M2 for the same allocation designs (different combinations of nonoverlapping/overlapping lines in environments and training set sizes). Overlapped sets of 30–50 lines in all the environments provided stable genomic‐enabled prediction accuracy. Reducing the size of the testing populations under all allocation designs decreases the prediction accuracy, which recovers when more lines are tested in all environments. Model M3 offers the possibility of maintaining the prediction accuracy throughout both extreme situations of all nonoverlapping lines and all overlapping lines. more...

Abstract Genomic selection in maize (Zea mays L.) has been one factor that has increased the rate of genetic gain when compared with other cereals. However, the technological foundations in maize also exist in other cereal crops that would allow prediction of hybrid performance based on general (GCA) and specific (SCA) combining abilities applied through genomic‐enabled prediction models. Further, the incorporation of genotype × environment (G × E) interaction effects present an opportunity to deploy hybrids to targeted environments. To test these concepts, a factorial mating design of elite yet divergent grain sorghum lines generated hybrids for evaluation. Inbred parents were genotyped, and markers were used to assess population structure and develop the genomic relationship matrix (GRM). Grain yield, height, and days to anthesis were collected for hybrids in replicated trials, and best linear unbiased estimates were used to train classical GCA‐SCA–based and genomic (GB) models under a hierarchical Bayesian framework. To incorporate population structure, GB was fitted using the GRM of both parents and hybrids. For GB models, G × E interaction effects were included by the Hadamard product between GRM and environments. A leave‐one‐out cross‐validation scheme was used to study the prediction capacity of models. Classical and genomic models effectively predicted hybrid performance and prediction accuracy increased by including genomic data. Genomic models effectively partitioned the variation due to GCA, SCA, and their interaction with the environment. A strategy to implement genomic selection for hybrid sorghum [Sorghum bicolor (L.) Moench] breeding is presented herein. more...

Abstract Genomic selection (GS) is revolutionizing conventional ways of developing new plants and animals. However, because it is a predictive methodology, GS strongly depends on statistical and machine learning to perform these predictions. For continuous outcomes, more models are available for GS. Unfortunately, for count data outcomes, there are few efficient statistical machine learning models for large datasets or for datasets with fewer observations than independent variables. For this reason, in this paper, we applied the univariate version of the Poisson deep neural network (PDNN) proposed earlier for genomic predictions of count data. The model was implemented with (a) the negative log‐likelihood of Poisson distribution as the loss function, (b) the rectified linear activation unit as the activation function in hidden layers, and (c) the exponential activation function in the output layer. The advantage of the PDNN model is that it captures complex patterns in the data by implementing many nonlinear transformations in the hidden layers. Moreover, since it was implemented in Tensorflow as the back‐end, and in Keras as the front‐end, the model can be applied to moderate and large datasets, which is a significant advantage over previous GS models for count data. The PDNN model was compared with deep learning models with continuous outcomes, conventional generalized Poisson regression models, and conventional Bayesian regression methods. We found that the PDNN model outperformed the Bayesian regression and generalized Poisson regression methods in terms of prediction accuracy, although it was not better than the conventional deep neural network with continuous outcomes. more...

Quantitative genetics states that phenotypic variation is a consequence of the interaction between genetic and environmental factors. Predictive breeding is based on this statement, and because of this, ways of modeling genetic effects are still evolving. At the same time, the same refinement must be used for processing environmental information. Here, we present an “enviromic assembly approach,” which includes using ecophysiology knowledge in shaping environmental relatedness into whole-genome predictions (GP) for plant breeding (referred to as enviromic-aided genomic prediction, E-GP). We propose that the quality of an environment is defined by the core of environmental typologies and their frequencies, which describe different zones of plant adaptation. From this, we derived markers of environmental similarity cost-effectively. Combined with the traditional additive and non-additive effects, this approach may better represent the putative phenotypic variation observed across diverse growing conditions (i.e., phenotypic plasticity). Then, we designed optimized multi-environment trials coupling genetic algorithms, enviromic assembly, and genomic kinships capable of providing in-silico realization of the genotype-environment combinations that must be phenotyped in the field. As proof of concept, we highlighted two E-GP applications: (1) managing the lack of phenotypic information in training accurate GP models across diverse environments and (2) guiding an early screening for yield plasticity exerting optimized phenotyping efforts. Our approach was tested using two tropical maize sets, two types of enviromics assembly, six experimental network sizes, and two types of optimized training set across environments. We observed that E-GP outperforms benchmark GP in all scenarios, especially when considering smaller training sets. The representativeness of genotype-environment combinations is more critical than the size of multi-environment trials (METs). The conventional genomic best-unbiased prediction (GBLUP) is inefficient in predicting the quality of a yet-to-be-seen environment, while enviromic assembly enabled it by increasing the accuracy of yield plasticity predictions. Furthermore, we discussed theoretical backgrounds underlying how intrinsic envirotype-phenotype covariances within the phenotypic records can impact the accuracy of GP. The E-GP is an efficient approach to better use environmental databases to deliver climate-smart solutions, reduce field costs, and anticipate future scenarios. more...

The constrained linear genomic selection index (CLGSI) is a linear combination of genomic estimated breeding values useful for predicting the net genetic merit, which in turn is a linear combination of true unobservable breeding values of the traits weighted by their respective economic values. The CLGSI is the most general genomic index and allows imposing constraints on the expected genetic gain per trait to make some traits change their mean values based on a predetermined level, while the rest of them remain without restrictions. In addition, it includes the unconstrained linear genomic index as a particular case. Using two real datasets and simulated data for seven selection cycles, we compared the theoretical results of the CLGSI with the theoretical results of the constrained linear phenotypic selection index (CLPSI). The criteria used to compare CLGSI vs. CLPSI efficiency were the estimated expected genetic gain per trait values, the selection response, and the interval between selection cycles. The results indicated that because the interval between selection cycles is shorter for the CLGSI than for the CLPSI, CLGSI is more efficient than CLPSI per unit of time, but its efficiency could be lower per selection cycle. Thus, CLGSI is a good option for performing genomic selection when there are genotyped candidates for selection. more...

The usefulness of genomic prediction (GP) for many animal and plant breeding programs has been highlighted for many studies in the last 20 years. In maize breeding programs, mostly dedicated to delivering more highly adapted and productive hybrids, this approach has been proved successful for both large- and small-scale breeding programs worldwide. Here, we present some of the strategies developed to improve the accuracy of GP in tropical maize, focusing on its use under low budget and small-scale conditions achieved for most of the hybrid breeding programs in developing countries. We highlight the most important outcomes obtained by the University of São Paulo (USP, Brazil) and how they can improve the accuracy of prediction in tropical maize hybrids. Our roadmap starts with the efforts for germplasm characterization, moving on to the practices for mating design, and the selection of the genotypes that are used to compose the training population in field phenotyping trials. Factors including population structure and the importance of non-additive effects (dominance and epistasis) controlling the desired trait are also outlined. Finally, we explain how the source of the molecular markers, environmental, and the modeling of genotype–environment interaction can affect the accuracy of GP. Results of 7 years of research in a public maize hybrid breeding program under tropical conditions are discussed, and with the great advances that have been made, we find that what is yet to come is exciting. The use of open-source software for the quality control of molecular markers, implementing GP, and envirotyping pipelines may reduce costs in an efficient computational manner. We conclude that exploring new models/tools using high-throughput phenotyping data along with large-scale envirotyping may bring more resolution and realism when predicting genotype performances. Despite the initial costs, mostly for genotyping, the GP platforms in combination with these other data sources can be a cost-effective approach for predicting the performance of maize hybrids for a large set of growing conditions. more...

To enable a scalable sparse testing genomic selection (GS) strategy at preliminary yield trials in the CIMMYT maize breeding program, optimal approaches to incorporate genotype by environment interaction (GEI) in genomic prediction models are explored. Two cross-validation schemes were evaluated: CV1, predicting the genetic merit of new bi-parental populations that have been evaluated in some environments and not others, and CV2, predicting the genetic merit of half of a bi-parental population that has been phenotyped in some environments and not others using the coefficient of determination (CDmean) to determine optimized subsets of a full-sib family to be evaluated in each environment. We report similar prediction accuracies in CV1 and CV2, however, CV2 has an intuitive appeal in that all bi-parental populations have representation across environments, allowing efficient use of information across environments. It is also ideal for building robust historical data because all individuals of a full-sib family have phenotypic data, albeit in different environments. Results show that grouping of environments according to similar growing/management conditions improved prediction accuracy and reduced computational requirements, providing a scalable, parsimonious approach to multi-environmental trials and GS in early testing stages. We further demonstrate that complementing the full-sib calibration set with optimized historical data results in improved prediction accuracy for the cross-validation schemes. more...

In maize, doubled haploid (DH) line production capacity of large-sized maize breeding programs often exceeds the capacity to phenotypically evaluate the complete set of testcross candidates in multi-location trials. The ability to partially select DH lines based on genotypic data while maintaining or improving genetic gains for key traits using phenotypic selection can result in significant resource savings. The present study aimed to evaluate genomic selection (GS) prediction scenarios for grain yield and agronomic traits of one of the tropical maize breeding pipelines of CIMMYT in eastern Africa, based on multi-year empirical data for designing a GS-based strategy at the early stages of the pipeline. We used field data from 3,068 tropical maize DH lines genotyped using rAmpSeq markers and evaluated as test crosses in well-watered (WW) and water-stress (WS) environments in Kenya from 2017 to 2019. Three prediction schemes were compared: (1) 1 year of performance data to predict a second year; (2) 2 years of pooled data to predict performance in the third year, and (3) using individual or pooled data plus converting a certain proportion of individuals from the testing set (TST) to the training set (TRN) to predict the next year's data. Employing five-fold cross-validation, the mean prediction accuracies for grain yield (GY) varied from 0.19 to 0.29 under WW and 0.22 to 0.31 under WS, when the 1-year datasets were used training set to predict a second year's data as a testing set. The mean prediction accuracies increased to 0.32 under WW and 0.31 under WS when the 2-year datasets were used as a training set to predict the third-year data set. In a forward prediction scenario, good predictive abilities (0.53 to 0.71) were found when the training set consisted of the previous year's breeding data and converting 30% of the next year's data from the testing set to the training set. The prediction accuracy for anthesis date and plant height across WW and WS environments obtained using 1-year data and integrating 10, 30, 50, 70, and 90% of the TST set to TRN set was much higher than those trained in individual years. We demonstrate that by increasing the TRN set to include genotypic and phenotypic data from the previous year and combining only 10–30% of the lines from the year of testing, the predicting accuracy can be increased, which in turn could be used to replace the first stage of field-based screening partially, thus saving significant costs associated with the testcross formation and multi-location testcross evaluation. more...

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. more...

Anther extrusion (AE) is the most important male floral trait for hybrid wheat seed production. AE is a complex quantitative trait that is difficult to phenotype reliably in field experiments not only due to high genotype-by-environment effects but also due to the short expression window in the field condition. In this study, we conducted a genome-wide association scan (GWAS) and explored the possibility of applying genomic prediction (GP) for AE in the CIMMYT hybrid wheat breeding program. An elite set of male lines (n = 603) were phenotype for anther count (AC) and anther visual score (VS) across three field experiments in 2017–2019 and genotyped with the 20K Infinitum is elect SNP array. GWAS produced five marker trait associations with small effects. For GP, the main effects of lines (L), environment (E), genomic (G) and pedigree relationships (A), and their interaction effects with environments were used to develop seven statistical models of incremental complexity. The base model used only L and E, whereas the most complex model included L, E, G, A, and G × E and A × E. These models were evaluated in three cross-validation scenarios (CV0, CV1, and CV2). In cross-validation CV0, data from two environments were used to predict an untested environment; in random cross-validation CV1, the test set was never evaluated in any environment; and in CV2, the genotypes in the test set were evaluated in only a subset of environments. The prediction accuracies ranged from −0.03 to 0.74 for AC and −0.01 to 0.54 for VS across different models and CV schemes. For both traits, the highest prediction accuracies with low variance were observed in CV2, and inclusion of the interaction effects increased prediction accuracy for AC only. In CV0, the prediction accuracy was 0.73 and 0.45 for AC and VS, respectively, indicating the high reliability of across environment prediction. Genomic prediction appears to be a very reliable tool for AE in hybrid wheat breeding. Moreover, high prediction accuracy in CV0 demonstrates the possibility of implementing genomic selection across breeding cycles in related germplasm, aiding the rapid breeding cycle. more...

Abstract When including genotype × environment interactions (G × E) in genomic prediction models, Hadamard or Kronecker products have been used to model the covariance structure of interactions. The relation between these two types of modeling has not been made clear in genomic prediction literature. Here, we demonstrate that a certain model based on a Hadamard formulation and another using the Kronecker product lead to exactly the same statistical model. Moreover, we illustrate how a multiplication of entries of covariance matrices is related to modeling locus × environmental‐variable interactions explicitly. Finally, we use a wheat and a maize data set to illustrate that the environmental covariance E can be specified easily, also if no information on environmental variables – such as temperature or precipitation – is available. Given that lines have been tested in different environments, the corresponding environmental covariance can simply be estimated from the training set as phenotypic covariance between environments. To achieve a high level of increase in predictive ability, the environmental covariance has to be defined appropriately and records on the performance of the lines of the test set under different environmental conditions have to be included in the training set. more...

Abstract Wheat quality improvement is an important objective in all wheat breeding programs. However, due to the cost, time and quantity of seed required, wheat quality is typically analyzed only in the last stages of the breeding cycle on a limited number of samples. The use of genomic prediction could greatly help to select for wheat quality more efficiently by reducing the cost and time required for this analysis. Here were evaluated the prediction performances of 13 wheat quality traits under two multi‐trait models (Bayesian multi‐trait multi‐environment [BMTME] and multi‐trait ridge regression [MTR]) using five data sets of wheat lines evaluated in the field during two consecutive years. Lines in the second year (testing) were predicted using the quality information obtained in the first year (training). For most quality traits were found moderate to high prediction accuracies, suggesting that the use of genomic selection could be feasible. The best predictions were obtained with the BMTME model in all traits and the worst with the MTR model. The best predictions with the BMTME model under the mean arctangent absolute percentage error (MAAPE) were for test weight across the five data sets, whereas the worst predictions were for the alveograph trait ALVPL. In contrast, under Pearson's correlation, the best predictions depended on the data set. The results obtained suggest that the BMTME model should be preferred for multi‐trait prediction analyses. This model allows to obtain not only the correlation among traits, but also the correlation among environments, helping to increase the prediction accuracy. more...

The rapid development of molecular markers and sequencing technologies has made it possible to use genomic prediction (GP) and selection (GS) in animal and plant breeding. However, when the number of observations (n) is large (thousands or millions), computational difficulties when handling these large genomic kernel relationship matrices (inverting and decomposing) increase exponentially. This problem increases when genomic × environment interaction and multi-trait kernels are included in the model. In this research we propose selecting a small number of lines m(m < n) for constructing an approximate kernel of lower rank than the original and thus exponentially decreasing the required computing time. First, we describe the full genomic method for single environment (FGSE) with a covariance matrix (kernel) including all n lines. Second, we select m lines and approximate the original kernel for the single environment model (APSE). Similarly, but including main effects and G × E, we explain a full genomic method with genotype × environment model (FGGE), and including m lines, we approximated the kernel method with G × E (APGE). We applied the proposed method to two different wheat data sets of different sizes (n) using the standard linear kernel Genomic Best Linear Unbiased Predictor (GBLUP) and also using eigen value decomposition. In both data sets, we compared the prediction performance and computing time for FGSE versus APSE; we also compared FGGE versus APGE. Results showed a competitive prediction performance of the approximated methods with a significant reduction in computing time. Genomic prediction accuracy depends on the decay of the eigenvalues (amount of variance information loss) of the original kernel as well as on the size of the selected lines m. more...

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. more...

One of the major issues in plant breeding is the occurrence of genotype × environment (GE) interaction. Several models have been created to understand this phenomenon and explore it. In the genomic era, several models were employed to improve selection by using markers and account for GE interaction simultaneously. Some of these models use special genetic covariance matrices. In addition, the scale of multi-environment trials is getting larger, and this increases the computational challenges. In this context, we propose an R package that, in general, allows building GE genomic covariance matrices and fitting linear mixed models, in particular, to a few genomic GE models. Here we propose two functions: one to prepare the genomic kernels accounting for the genomic GE and another to perform genomic prediction using a Bayesian linear mixed model. A specific treatment is given for sparse covariance matrices, in particular, to block diagonal matrices that are present in some GE models in order to decrease the computational demand. In empirical comparisons with Bayesian Genomic Linear Regression (BGLR), accuracies and the mean squared error were similar; however, the computational time was up to five times lower than when using the classic approach. Bayesian Genomic Genotype × Environment Interaction (BGGE) is a fast, efficient option for creating genomic GE kernels and making genomic predictions. more...

Multi-environment trials are routinely conducted in plant breeding to select candidates for the next selection cycle. In this study, we compare the prediction accuracy of four developed genomic-enabled prediction models: (1) single-environment, main genotypic effect model (SM); (2) multi-environment, main genotypic effects model (MM); (3) multi-environment, single variance G×E deviation model (MDs); and (4) multi-environment, environment-specific variance G×E deviation model (MDe). Each of these four models were fitted using two kernel methods: a linear kernel Genomic Best Linear Unbiased Predictor, GBLUP (GB), and a nonlinear kernel Gaussian kernel (GK). The eight model-method combinations were applied to two extensive Brazilian maize data sets (HEL and USP data sets), having different numbers of maize hybrids evaluated in different environments for grain yield (GY), plant height (PH), and ear height (EH). Results show that the MDe and the MDs models fitted with the Gaussian kernel (MDe-GK, and MDs-GK) had the highest prediction accuracy. For GY in the HEL data set, the increase in prediction accuracy of SM-GK over SM-GB ranged from 9 to 32%. For the MM, MDs, and MDe models, the increase in prediction accuracy of GK over GB ranged from 9 to 49%. For GY in the USP data set, the increase in prediction accuracy of SM-GK over SM-GB ranged from 0 to 7%. For the MM, MDs, and MDe models, the increase in prediction accuracy of GK over GB ranged from 34 to 70%. For traits PH and EH, gains in prediction accuracy of models with GK compared to models with GB were smaller than those achieved in GY. Also, these gains in prediction accuracy decreased when a more difficult prediction problem was studied. more...

Much of the world’s population growth will occur in regions where food insecurity is prevalent, with large increases in food demand projected in regions of Africa and South Asia. While improving food security in these regions will require a multi-faceted approach, improved performance of crop varieties in these regions will play a critical role. Current rates of genetic gain in breeding programs serving Africa and South Asia fall below rates achieved in other regions of the world. Given resource constraints, increased genetic gain in these regions cannot be achieved by simply expanding the size of breeding programs. New approaches to breeding are required. The Genomic Open-source Breeding informatics initiative (GOBii) and Excellence in Breeding Platform (EiB) are working with public sector breeding programs to build capacity, develop breeding strategies, and build breeding informatics capabilities to enable routine use of new technologies that can improve the efficiency of breeding programs and increase genetic gains. Simulations evaluating breeding strategies indicate cost-effective implementations of genomic selection (GS) are feasible using relatively small training sets, and proof-of-concept implementations have been validated in the International Maize and Wheat Improvement Center (CIMMYT) maize breeding program. Progress on GOBii, EiB, and implementation of GS in CIMMYT and International Crops Research Institute for the Semi-Arid Tropics (ICRISAT) breeding programs are discussed, as well as strategies for routine implementation of GS in breeding programs serving Africa and South Asia. more...

Genomic selection (GS) is an emerging methodology that helps select superior lines among experimental cultivars in plant breeding programs. It offers the opportunity to increase the productivity of cultivars by delivering increased genetic gains and reducing the breeding cycles. This methodology requires inexpensive and sufficiently dense marker information to be successful, and with whole genome sequencing, it has become an important tool in many crops. The recent assembly of the pearl millet genome has made it possible to employ GS models to improve the selection procedure in pearl millet breeding programs. Here, three GS models were implemented and compared using grain yield and dense molecular marker information of pearl millet obtained from two different genotyping platforms (C [conventional GBS RAD-seq] and T [tunable GBS tGBS]). The models were evaluated using three different cross-validation (CV) schemes mimicking real situations that breeders face in breeding programs: CV2 resembles an incomplete field trial, CV1 predicts the performance of untested hybrids, and CV0 predicts the performance of hybrids in unobserved environments. We found that (i) adding phenotypic information of parental inbreds to the calibration sets improved predictive ability, (ii) accounting for genotype-by-environment interaction also increased the performance of the models, and (iii) superior strategies should consider the use of the molecular markers derived from the T platform (tGBS). more...

Genomic selection predicts the genomic estimated breeding values (GEBVs) of individuals not previously phenotyped. Several studies have investigated the accuracy of genomic predictions in maize but there is little empirical evidence on the practical performance of lines selected based on phenotype in comparison with those selected solely on GEBVs in advanced testcross yield trials. The main objectives of this study were to (1) empirically compare the performance of tropical maize hybrids selected through phenotypic selection (PS) and genomic selection (GS) under well-watered (WW) and managed drought stress (WS) conditions in Kenya, and (2) compare the cost–benefit analysis of GS and PS. For this study, we used two experimental maize data sets (stage I and stage II yield trials). The stage I data set consisted of 1492 doubled haploid (DH) lines genotyped with rAmpSeq SNPs. A subset of these lines (855) representing various DH populations within the stage I cohort was crossed with an individual single-cross tester chosen to complement each population. These testcross hybrids were evaluated in replicated trials under WW and WS conditions for grain yield and other agronomic traits, while the remaining 637 DH lines were predicted using the 855 lines as a training set. The second data set (stage II) consists of 348 DH lines from the first data set. Among these 348 best DH lines, 172 lines selected were solely based on GEBVs, and 176 lines were selected based on phenotypic performance. Each of the 348 DH lines were crossed with three common testers from complementary heterotic groups, and the resulting 1042 testcross hybrids and six commercial checks were evaluated in four to five WW locations and one WS condition in Kenya. For stage I trials, the cross-validated prediction accuracy for grain yield was 0.67 and 0.65 under WW and WS conditions, respectively. We found similar responses to selection using PS and GS for grain yield other agronomic traits under WW and WS conditions. The top 15% of hybrids advanced through GS and PS gave 21%–23% higher grain yield under WW and 51%–52% more grain yield under WS than the mean of the checks. The GS reduced the cost by 32% over the PS with similar selection gains. We concluded that the use of GS for yield under WW and WS conditions in maize can produce selection candidates with similar performance as those generated from conventional PS, but at a lower cost, and therefore, should be incorporated into maize breeding pipelines to increase breeding program efficiency. more...

Droughts and high temperatures are the main abiotic constraints hampering durum wheat production. This study investigated the accumulation of phenolic acids (PAs) in the wholemeal flour of six durum wheat cultivars under drought and heat stress. Phenolic acids were extracted from wholemeals and analysed through HPLC-DAD analysis. Ferulic acid was the most represented PA, varying from 390.1 to 785.6 µg/g dry matter across all cultivars and growth conditions, followed by sinapic acids, p-coumaric, vanillic, syringic, and p-hydroxybenzoic acids. Among the cultivars, Cirno had the highest PAs content, especially under severe drought conditions. Heat stress enhanced the accumulation of minor individual PAs, whereas severe drought increased ferulic acid and total PAs. Broad-sense heritability was low (0.23) for p-coumaric acid but ≥0.69 for all other components. Positive correlations occurred between PA content and grain morphology and between test weight and grain yield. Durum wheat genotypes with good yields and high accumulation of PAs across different growing conditions could be significant for durum wheat resilience and health-promoting value. more...

Developing genomic selection (GS) models is an important step in applying GS to accelerate the rate of genetic gain in grain yield in plant breeding. In this study, seven genomic prediction models under two cross-validation (CV) scenarios were tested on 287 advanced elite spring wheat lines phenotyped for grain yield (GY), thousand-grain weight (GW), grain number (GN), and thermal time for flowering (TTF) in 18 international environments (year-location combinations) in major wheat-producing countries in 2010 and 2011. Prediction models with genomic and pedigree information included main effects and interaction with environments. Two random CV schemes were applied to predict a subset of lines that were not observed in any of the 18 environments (CV1), and a subset of lines that were not observed in a set of the environments, but were observed in other environments (CV2). Genomic prediction models, including genotype × environment (G×E) interaction, had the highest average prediction ability under the CV1 scenario for GY (0.31), GN (0.32), GW (0.45), and TTF (0.27). For CV2, the average prediction ability of the model including the interaction terms was generally high for GY (0.38), GN (0.43), GW (0.63), and TTF (0.53). Wheat lines in site-year combinations in Mexico and India had relatively high prediction ability for GY and GW. Results indicated that prediction ability of lines not observed in certain environments could be relatively high for genomic selection when predicting G×E interaction in multi-environment trials. more...

In this study, we used genotype × environment interactions (G×E) models for hybrid prediction, where similarity between lines was assessed by pedigree and molecular markers, and similarity between environments was accounted for by environmental covariables. We use five genomic and pedigree models (M1–M5) under four cross-validation (CV) schemes: prediction of hybrids when the training set (i) includes hybrids of all males and females evaluated only in some environments (T2FM), (ii) excludes all progenies from a randomly selected male (T1M), (iii) includes all progenies from 20% randomly selected females in combination with all males (T1F), and (iv) includes one randomly selected male plus 40% randomly selected females that were crossed with it (T0FM). Models were tested on a total of 1888 wheat ( L.) hybrids including 18 males and 667 females in three consecutive years. For grain yield, the most complex model (M5) under T2FM had slightly higher prediction accuracy than the less complex model. For T1F, the prediction accuracy of hybrids for grain yield and other traits of the most complete model was 0.50 to 0.55. For T1M, Model M3 exhibited high prediction accuracies for flowering traits (0.71), whereas the more complex model (M5) demonstrated high accuracy for grain yield (0.5). For T0FM, the prediction accuracy for grain yield of Model M5 was 0.61. Including genomic and pedigree gave relatively high prediction accuracy even when both parents were untested. Results show that it is possible to predict unobserved hybrids when modeling genomic general combining ability (GCA) and specific combining ability (SCA) and their interactions with environments. more...

Genomic prediction studies incorporating genotype × environment (G×E) interaction effects are limited in durum wheat. We tested the genomic-enabled prediction accuracy (PA) of Genomic Best Linear Unbiased Predictor (GBLUP) models—six non-G × E and three G × E models—on three basic cross-validation (CV) schemes— in predicting incomplete field trials (CV2), new lines (CV1), and lines in untested environments (CV0)— in a durum wheat panel grown under yield potential, drought stress, and heat stress conditions. For CV0, three scenarios were considered: (i) leave-one environment out (CV0-Env); (ii) leave one site out (CV0-Site); and (iii) leave 1 yr out (CV0-Year). The reaction norm models with G × E effects showed higher PA than the non-G × E models. Among the CV schemes, CV2 and CV0-Env had higher PA (0.58 each) than the CV1 scheme (0.35). When the average of all the models and CV schemes were considered, among the eight traits— grain yield, thousand grain weight, grain number, days to anthesis, days to maturity, plant height, and normalized difference vegetation index at vegetative (NDVIvg) and grain filling (NDVIllg)—, plant height had the highest PA (0.68) and moderate values were observed for grain yield (0.34). The results indicated that genomic selection models incorporating G × E interaction show great promise for forward prediction and application in durum wheat breeding to increase genetic gains. more...

Wheat ( L.) breeding programs test experimental lines in multiple locations over multiple years to get an accurate assessment of grain yield and yield stability. Selections in early generations of the breeding pipeline are based on information from only one or few locations and thus materials are advanced with little knowledge of the genotype × environment interaction (G × E) effects. Later, large trials are conducted in several locations to assess the performance of more advanced lines across environments. Genomic selection (GS) models that include G × E covariates allow us to borrow information not only from related materials, but also from historical and correlated environments to better predict performance within and across specific environments. We used reaction norm models with several cross-validation schemes to demonstrate the increased breeding efficiency of Kansas State University’s hard red winter wheat breeding program. The GS reaction norm models line effect (L) + environment effect (E), L + E + genotype environment (G), and L + E + G + (G × E) effects) showed high accuracy values (>0.4) when predicting the yield performance in untested environments, sites or both. The GS model L + E + G + (G × E) presented the highest prediction ability ( = 0.54) when predicting yield in incomplete field trials for locations with a moderate number of lines. The difficulty of predicting future years (forward prediction) is indicated by the relatively low accuracy ( = 0.171) seen even when environments with 300+ lines were included. more...

The leaf spotting diseases in wheat that include Septoria tritici blotch (STB) caused by , Stagonospora nodorum blotch (SNB) caused by , and tan spot (TS) caused by pose challenges to breeding programs in selecting for resistance. A promising approach that could enable selection prior to phenotyping is genomic selection that uses genome-wide markers to estimate breeding values (BVs) for quantitative traits. To evaluate this approach for seedling and/or adult plant resistance (APR) to STB, SNB, and TS, we compared the predictive ability of least-squares (LS) approach with genomic-enabled prediction models including genomic best linear unbiased predictor (GBLUP), Bayesian ridge regression (BRR), Bayes A (BA), Bayes B (BB), Bayes Cπ (BC), Bayesian least absolute shrinkage and selection operator (BL), and reproducing kernel Hilbert spaces markers (RKHS-M), a pedigree-based model (RKHS-P) and RKHS markers and pedigree (RKHS-MP). We observed that LS gave the lowest prediction accuracies and RKHS-MP, the highest. The genomic-enabled prediction models and RKHS-P gave similar accuracies. The increase in accuracy using genomic prediction models over LS was 48%. The mean genomic prediction accuracies were 0.45 for STB (APR), 0.55 for SNB (seedling), 0.66 for TS (seedling) and 0.48 for TS (APR). We also compared markers from two whole-genome profiling approaches: genotyping by sequencing (GBS) and diversity arrays technology sequencing (DArTseq) for prediction. While, GBS markers performed slightly better than DArTseq, combining markers from the two approaches did not improve accuracies. We conclude that implementing GS in breeding for these diseases would help to achieve higher accuracies and rapid gains from selection. more...

Marker-assisted recurrent selection (MARS) is a breeding method used to accumulate favorable alleles that for example confer tolerance to drought in inbred lines from several genomic regions within a single population. A bi-parental cross formed from two parents that combine resistance to Striga hermonthica with drought tolerance, which was improved through MARS, was used to assess changes in the frequency of favorable alleles and its impact on inbred line improvement. A total of 200 testcrosses of randomly selected S1 lines derived from the original (C0) and advanced selection cycles of this bi-parental population, were evaluated under drought stress (DS) and well-watered (WW) conditions at Ikenne and under artificial Striga infestation at Abuja and Mokwa in Nigeria in 2014 and 2015. Also, 60 randomly selected S1 lines each derived from the four cycles (C0, C1, C2, C3) were genotyped with 233 SNP markers using KASP assay. The results showed that the frequency of favorable alleles increased with MARS in the bi-parental population with none of the markers showing fixation. The gain in grain yield was not significant under DS condition due to the combined effect of DS and armyworm infestation in 2015. Because the parents used for developing the bi-parental cross combined tolerance to drought with resistance to Striga, improvement in grain yield under DS did not result in undesirable changes in resistance to the parasite in the bi-parental maize population improved through MARS. MARS increased the mean number of combinations of favorable alleles in S1 lines from 114 in C0 to 124 in C3. The level of heterozygosity decreased by 15%, while homozygosity increased by 13% due to the loss of some genotypes in the population. This study demonstrated the effectiveness of MARS in increasing the frequency of favorable alleles for tolerance to drought without disrupting the level of resistance to Striga in a bi-parental population targeted as a source of improved maize inbred lines. more...

The objective of marker assisted recurrent selection (MARS) is to increase the frequency of favorable marker alleles in a population before inbred line extraction. This approach was used to improve drought tolerance and grain yield (GY) in a biparental cross of two elite drought tolerant lines. The testcrosses of randomly selected 50 S1 lines from each of the three selection cycles (C0, C1, C2) of the MARS population, parental testcrosses and the cross between the two parents (F1) were evaluated under drought stress (DS) and well watered (WW) well as under rainfed conditions to determine genetic gains in GY and other agronomic traits. Also, the S1 lines derived from each selection types were genotyped with single nucleotide polymorphism (SNP) markers. Testcrosses derived from C2 produced significantly higher grain field under DS than those derived from C0 with a relative genetic gain of 7% per cycle. Also, the testcrosses of S1 lines from C2 showed an average genetic gain of 1% per cycle under WW condition and 3% per cycle under rainfed condition. Molecular analysis revealed that the frequency of favorable marker alleles increased from 0.510 at C0 to 0.515 at C2, while the effective number of alleles (Ne) per locus decreased from C0 (1.93) to C2 (1.87). Our results underscore the effectiveness of MARS for improvement of GY under DS condition. more...

Genomic selection (GS) unlike marker-assisted backcrossing (MABC) predicts breeding values of lines using genome-wide marker profiling and allows selection of lines prior to field-phenotyping, thereby shortening the breeding cycle. A collection of 320 elite breeding lines was selected and phenotyped extensively for yield and yield related traits at two different locations (Delhi and Patancheru, India) during the crop seasons 2011-12 and 2012-13 under rainfed and irrigated conditions. In parallel, these lines were also genotyped using DArTseq platform to generate data on 3,000 polymorphic markers. Phenotypic and genotypic data were used with six statistical GS models to estimate the prediction accuracies. GS models were tested for four yield related traits viz. seed yield, 100 seed weight, days to 50% flowering and days to maturity. Prediction accuracy for the models tested varied from 0.138 (seed yield) to 0.912 (100 seed weight), whereas performance of models did not show any significant difference for estimating prediction accuracy within traits. Kinship matrix calculated using genotyping data reaffirmed existence of two different groups within selected lines. There was not much effect of population structure on prediction accuracy. In brief, present study establishes the necessary resources for deployment of GS in chickpea breeding. more...

Durable resistance to the rust diseases of wheat ( L.) can be achieved by developing lines that have race-nonspecific adult plant resistance conferred by multiple minor slow-rusting genes. Genomic selection (GS) is a promising tool for accumulating favorable alleles of slow-rusting genes. In this study, five CIMMYT wheat populations evaluated for resistance were used to predict resistance to stem rust () and yellow rust () using Bayesian least absolute shrinkage and selection operator (LASSO) (BL), ridge regression (RR), and support vector regression with linear or radial basis function kernel models. All parents and populations were genotyped using 1400 Diversity Arrays Technology markers and different prediction problems were assessed. Results show that prediction ability for yellow rust was lower than for stem rust, probably due to differences in the conditions of infection of both diseases. For within population and environment, the correlation between predicted and observed values (Pearson’s correlation [ρ]) was greater than 0.50 in 90% of the evaluations whereas for yellow rust, ρ ranged from 0.0637 to 0.6253. The BL and RR models have similar prediction ability, with a slight superiority of the BL confirming reports about the additive nature of rust resistance. When making predictions between environments and/or between populations, including information from another environment or environments or another population or populations improved prediction. more...

Crop modelling has the potential to contribute to global food and nutrition security. This paper briefly examines the history of crop modelling by international crop research centres of the CGIAR (formerly Consultative Group on International Agricultural Research but now known simply as CGIAR), whose primary focus is on less developed countries. Basic principles of crop modelling building up to a Genotype × Environment × Management × Socioeconomic (G × E × M × S) paradigm, are explained. Modelling has contributed to better understanding of crop performance and yield gaps, better prediction of pest and insect outbreaks, and improving the efficiency of crop management including irrigation systems and optimization of planting dates. New developments include, for example, use of remote sensed data and mobile phone technology linked to crop management decision support models, data sharing in the new era of big data, and the use of genomic selection and crop simulation models linked to environmental data to help make crop breeding decisions. Socio-economic applications include foresight analysis of agricultural systems under global change scenarios, and the consequences of potential food system shocks are also described. These approaches are discussed in this paper which also calls for closer collaboration among disciplines in order to better serve the crop research and development communities by providing model based recommendations ranging from policy development at the level of governmental agencies to direct crop management support for resource poor farmers. more...