Data Augmentation Enhances Plant-Genomic-Enabled Predictions.

Genomic selection (GS) is revolutionizing plant breeding. However, its practical implementation is still challenging, since there are many factors that affect its accuracy. For this reason, this research explores data augmentation with the goal of improving its accuracy. Deep neural networks with data augmentation (DA) generate synthetic data from the original training set to increase the training set and to improve the prediction performance of any statistical or machine learning algorithm. There is much empirical evidence of their success in many computer vision applications. Due to this, DA was explored in the context of GS using 14 real datasets. We found empirical evidence that DA is a powerful tool to improve the prediction accuracy, since we improved the prediction accuracy of the top lines in the 14 datasets under study. On average, across datasets and traits, the gain in prediction performance of the DA approach regarding the Conventional method in the top 20% of lines in the testing set was 108.4% in terms of the NRMSE and 107.4% in terms of the MAAPE, but a worse performance was observed on the whole testing set. We encourage more empirical evaluations to support our findings.

Improving hybrid rice breeding programs via stochastic simulations: number of parents, number of hybrids, tester update, and genomic prediction of hybrid performance.

KEY MESSAGE: Schemes that use genomic prediction outperform others, updating testers increases hybrid genetic gain, and larger population sizes tend to have higher genetic gain and less depletion of genetic variance One of the most common methods to improve hybrid performance is reciprocal recurrent selection (RRS). Genomic prediction (GP) can be used to increase genetic gain in RRS by reducing cycle length, but it is also possible to use GP to predict single-cross hybrid performance. The impact of the latter method on genetic gain has yet to be previously reported. Therefore, we compared via stochastic simulations various phenotypic and genomics-assisted RRS breeding schemes which used GP to predict hybrid performance rather than reducing cycle length, which allows minimal changes to traditional breeding schemes. We also compared three breeding sizes scenarios that varied the number of genotypes crossed within heterotic pools, the number of genotypes crossed between heterotic pools, the number of hybrids evaluated, and the number of genomic predicted hybrids. Our results demonstrated that schemes that used genomic prediction of hybrid performance outperformed the others for the average interpopulation hybrid population and the best hybrid performance. Furthermore, updating the testers increased hybrid genetic gain with phenotypic RRS. As expected, the largest breeding size tested had the highest rates of genetic improvement and the lowest decrease in additive genetic variance due to the drift. Therefore, this study demonstrates the usefulness of single-cross prediction, which may be easier to implement than rapid-cycling RRS and cyclical updating of testers. We also reiterate that larger population sizes tend to have higher genetic gain and less depletion of genetic variance.

Construction and genetic characterization of an interspecific raspberry hybrids panel aiming resistance to late leaf rust and adaptation to tropical regions.

Raspberries (Rubus spp) are temperate climate fruits with profitable high returns and have the potential for diversification of fruit growing in mid to low-latitude regions. However, there are still no cultivars adapted to climatic conditions and high pressure of diseases that occurs in tropical areas. In this context, our objective was to evaluate the genetic diversity from a 116 raspberry genotypes panel obtained from interspecific crosses in a testcross scheme with four cultivars already introduced in Brazil. The panel was genotyped via genotyping-by-sequencing. 28,373 and 27,281 SNPs were obtained, using the species R. occidentalis and R. idaeus genomes as references, respectively. A third marker dataset was constructed consisting of 41,292 non-coincident markers. Overall, there were no differences in the results when using the different marker sets for the subsequent analyses. The mean heterozygosity was 0.54. The average effective population size was 174, indicating great genetic variability. The other analyses revealed that the half-sibling families were structured in three groups. It is concluded that the studied panel has great potential for breeding and further genetic studies. Moreover, only one of the three marker matrices is sufficient for diversity studies.

On the usefulness of mock genomes to define heterotic pools, testers, and hybrid predictions in orphan crops.

The advances in genomics in recent years have increased the accuracy and efficiency of breeding programs for many crops. Nevertheless, the adoption of genomic enhancement for several other crops essential in developing countries is still limited, especially for those that do not have a reference genome. These crops are more often called orphans. This is the first report to show how the results provided by different platforms, including the use of a simulated genome, called the mock genome, can generate in population structure and genetic diversity studies, especially when the intention is to use this information to support the formation of heterotic groups, choice of testers, and genomic prediction of single crosses. For that, we used a method to assemble a reference genome to perform the single-nucleotide polymorphism (SNP) calling without needing an external genome. Thus, we compared the analysis results using the mock genome with the standard approaches (array and genotyping-by-sequencing (GBS)). The results showed that the GBS-Mock presented similar results to the standard methods of genetic diversity studies, division of heterotic groups, the definition of testers, and genomic prediction. These results showed that a mock genome constructed from the population's intrinsic polymorphisms to perform the SNP calling is an effective alternative for conducting genomic studies of this nature in orphan crops, especially those that do not have a reference genome.

Genome-wide association analysis of hyperspectral reflectance data to dissect the genetic architecture of growth-related traits in maize under plant growth-promoting bacteria inoculation.

Plant growth-promoting bacteria (PGPB) may be of use for increasing crop yield and plant resilience to biotic and abiotic stressors. Using hyperspectral reflectance data to assess growth-related traits may shed light on the underlying genetics as such data can help assess biochemical and physiological traits. This study aimed to integrate hyperspectral reflectance data with genome-wide association analyses to examine maize growth-related traits under PGPB inoculation. A total of 360 inbred maize lines with 13,826 single nucleotide polymorphisms (SNPs) were evaluated with and without PGPB inoculation; 150 hyperspectral wavelength reflectances at 386-1021 nm and 131 hyperspectral indices were used in the analysis. Plant height, stalk diameter, and shoot dry mass were measured manually. Overall, hyperspectral signatures produced similar or higher genomic heritability estimates than those of manually measured phenotypes, and they were genetically correlated with manually measured phenotypes. Furthermore, several hyperspectral reflectance values and spectral indices were identified by genome-wide association analysis as potential markers for growth-related traits under PGPB inoculation. Eight SNPs were detected, which were commonly associated with manually measured and hyperspectral phenotypes. Different genomic regions were found for plant growth and hyperspectral phenotypes between with and without PGPB inoculation. Moreover, the hyperspectral phenotypes were associated with genes previously reported as candidates for nitrogen uptake efficiency, tolerance to abiotic stressors, and kernel size. In addition, a Shiny web application was developed to explore multiphenotype genome-wide association results interactively. Taken together, our results demonstrate the usefulness of hyperspectral-based phenotyping for studying maize growth-related traits in response to PGPB inoculation.

Enviromic-based kernels may optimize resource allocation with multi-trait multi-environment genomic prediction for tropical Maize.

BACKGROUND: Success in any genomic prediction platform is directly dependent on establishing a representative training set. This is a complex task, even in single-trait single-environment conditions and tends to be even more intricated wherein additional information from envirotyping and correlated traits are considered. Here, we aimed to design optimized training sets focused on genomic prediction, considering multi-trait multi-environment trials, and how those methods may increase accuracy reducing phenotyping costs. For that, we considered single-trait multi-environment trials and multi-trait multi-environment trials for three traits: grain yield, plant height, and ear height, two datasets, and two cross-validation schemes. Next, two strategies for designing optimized training sets were conceived, first considering only the genomic by environment by trait interaction (GET), while a second including large-scale environmental data (W, enviromics) as genomic by enviromic by trait interaction (GWT). The effective number of individuals (genotypes × environments × traits) was assumed as those that represent at least 98% of each kernel (GET or GWT) variation, in which those individuals were then selected by a genetic algorithm based on prediction error variance criteria to compose an optimized training set for genomic prediction purposes. RESULTS: The combined use of genomic and enviromic data efficiently designs optimized training sets for genomic prediction, improving the response to selection per dollar invested by up to 145% when compared to the model without enviromic data, and even more when compared to cross validation scheme with 70% of training set or pure phenotypic selection. Prediction models that include G × E or enviromic data + G × E yielded better prediction ability. CONCLUSIONS: Our findings indicate that a genomic by enviromic by trait interaction kernel associated with genetic algorithms is efficient and can be proposed as a promising approach to designing optimized training sets for genomic prediction when the variance-covariance matrix of traits is available. Additionally, great improvements in the genetic gains per dollar invested were observed, suggesting that a good allocation of resources can be deployed by using the proposed approach.

Envirome-wide associations enhance multi-year genome-based prediction of historical wheat breeding data.

Linking 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.

A divide-and-conquer approach for genomic prediction in rubber tree using machine learning.

Rubber tree (Hevea brasiliensis) is the main feedstock for commercial rubber; however, its long vegetative cycle has hindered the development of more productive varieties via breeding programs. With the availability of H. brasiliensis genomic data, several linkage maps with associated quantitative trait loci have been constructed and suggested as a tool for marker-assisted selection. Nonetheless, novel genomic strategies are still needed, and genomic selection (GS) may facilitate rubber tree breeding programs aimed at reducing the required cycles for performance assessment. Even though such a methodology has already been shown to be a promising tool for rubber tree breeding, increased model predictive capabilities and practical application are still needed. Here, we developed a novel machine learning-based approach for predicting rubber tree stem circumference based on molecular markers. Through a divide-and-conquer strategy, we propose a neural network prediction system with two stages: (1) subpopulation prediction and (2) phenotype estimation. This approach yielded higher accuracies than traditional statistical models in a single-environment scenario. By delivering large accuracy improvements, our methodology represents a powerful tool for use in Hevea GS strategies. Therefore, the incorporation of machine learning techniques into rubber tree GS represents an opportunity to build more robust models and optimize Hevea breeding programs.

Genotyping marker density and prediction models effects in long-term breeding schemes of cross-pollinated crops.

KEY MESSAGE: In genomic recurrent selection, the more markers, the better because they buffer the linkage disequilibrium losses caused by recombination over cycles, and consequently, provide higher responses to selection. Reductions of genotyping marker density have been extensively evaluated as potential strategies to reduce the genotyping costs of genomic selection (GS). Low-density marker panels are appealing in GS because they entail lower multicollinearity and computing time and allow more individuals to be genotyped for the same cost. However, statistical models used in GS are usually evaluated with empirical data, using "static" training sets and populations. This may be adequate for making predictions during a breeding program's initial cycles but not for the long-term. Moreover, studies that focus on long selective breeding cycles generally do not consider GS models with the effect of dominance, which is particularly important for breeding outcomes in cross-pollinated crops. Hence, dominance effects are important and unexplored in GS for long-term programs involving allogamous species. To address it, we employed two approaches: analysis of empirical maize datasets and simulations of long-term breeding applying phenotypic and genomic recurrent selection (intrapopulation and reciprocal schemes). In both schemes, we simulated twenty breeding cycles and assessed the effect of marker density reduction on the population mean, the best crosses, additive variance, selective accuracy, and response to selection with models [additive, additive-dominant, general (GCA), and this plus specific combining ability (GCA + SCA)]. Our results indicate that marker reduction based on linkage disequilibrium levels provides useful predictions only within a cycle, as accuracy significantly decreases over cycles. In the long-term, without training set updating, high-marker density provides the best responses to selection. The model to be used depends on the breeding scheme: additive for intrapopulation and additive-dominant or GCA + SCA for reciprocal.

Optimizing self-pollinated crop breeding employing genomic selection: From schemes to updating training sets.

Long-term breeding schemes using genomic selection (GS) can boost the response to selection per year. Although several studies have shown that GS delivers a higher response to selection, only a few analyze which stage GS produces better results and how to update the training population to maintain prediction accuracy. We used stochastic simulation to compare five GS breeding schemes in a self-pollinated long-term breeding program. Also, we evaluated four strategies, using distinct methods and sizes, to update the training set. Finally, regarding breeding schemes, we proposed a new approach using GS to select the best individuals in each F2 progeny, based on genomic estimated breeding values and genetic divergence, to cross them and generate a new recombination event. Our results showed that the best scenario was using GS in F2, followed by the phenotypic selection of new parents in F4. For TS updating, adding new data every cycle (over 768) to update the TS maintains the prediction accuracy at satisfactory levels for more breeding cycles. However, only the last three generations can be kept in the TS, optimizing the genetic relationship between TS and the targeted population and reducing the computing demand and risks. Hence, we believe that our results may help breeders optimize GS in their programs and improve genetic gain in long-term schemes.

Partial Least Squares Enhances Genomic Prediction of New Environments.

In plant breeding, the need to improve the prediction of future seasons or new locations and/or environments, also denoted as "leave one environment out," is of paramount importance to increase the genetic gain in breeding programs and contribute to food and nutrition security worldwide. Genomic selection (GS) has the potential to increase the accuracy of future seasons or new locations because it is a predictive methodology. However, most statistical machine learning methods used for the task of predicting a new environment or season struggle to produce moderate or high prediction accuracies. For this reason, in this study we explore the use of the partial least squares (PLS) regression methodology for this specific task, and we benchmark its performance with the Bayesian Genomic Best Linear Unbiased Predictor (GBLUP) method. The benchmarking process was done with 14 real datasets. We found that in all datasets the PLS method outperformed the popular GBLUP method by margins between 0% (in the Indica data) and 228.28% (in the Disease data) across traits, environments, and types of predictors. Our results show great empirical evidence of the power of the PLS methodology for the prediction of future seasons or new environments.

Genome and Environment Based Prediction Models and Methods of Complex Traits Incorporating Genotype × Environment Interaction.

Genomic-enabled prediction models are of paramount importance for the successful implementation of genomic selection (GS) based on breeding values. As opposed to animal breeding, plant breeding includes extensive multienvironment and multiyear field trial data. Hence, genomic-enabled prediction models should include genotype × environment (G × E) interaction, which most of the time increases the prediction performance when the response of lines are different from environment to environment. In this chapter, we describe a historical timeline since 2012 related to advances of the GS models that take into account G × E interaction. We describe theoretical and practical aspects of those GS models, including the gains in prediction performance when including G × E structures for both complex continuous and categorical scale traits. Then, we detailed and explained the main G × E genomic prediction models for complex traits measured in continuous and noncontinuous (categorical) scale. Related to G × E interaction models this review also examine the analyses of the information generated with high-throughput phenotype data (phenomic) and the joint analyses of multitrait and multienvironment field trial data that is also employed in the general assessment of multitrait G × E interaction. The inclusion of nongenomic data in increasing the accuracy and biological reliability of the G × E approach is also outlined. We show the recent advances in large-scale envirotyping (enviromics), and how the use of mechanistic computational modeling can derive the crop growth and development aspects useful for predicting phenotypes and explaining G × E.

Automated Machine Learning: A Case Study of Genomic "Image-Based" Prediction in Maize Hybrids.

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.

Genetic Trends Estimation in IRRIs Rice Drought Breeding Program and Identification of High Yielding Drought-Tolerant Lines.

Estimating genetic trends using historical data is an important parameter to check the success of the breeding programs. The estimated genetic trends can act as a guideline to target the appropriate breeding strategies and optimize the breeding program for improved genetic gains. In this study, 17 years of historical data from IRRI's rice drought breeding program was used to estimate the genetic trends and assess the breeding program's success. We also identified top-performing lines based on grain yield breeding values as an elite panel for implementing future population improvement-based breeding schemes. A two-stage approach of pedigree-based mixed model analysis was used to analyze the data and extract the breeding values and estimate the genetic trends for grain yield under non-stress, drought, and in combined data of non-stress and drought. Lower grain yield values were observed in all the drought trials. Heritability for grain yield estimates ranged between 0.20 and 0.94 under the drought trials and 0.43-0.83 under non-stress trials. Under non-stress conditions, the genetic gain of 0.21% (10.22 kg/ha/year) for genotypes and 0.17% (7.90 kg/ha/year) for checks was observed. The genetic trend under drought conditions exhibited a positive trend with the genetic gain of 0.13% (2.29 kg/ha/year) for genotypes and 0.55% (9.52 kg/ha/year) for checks. For combined analysis showed a genetic gain of 0.27% (8.32 kg/ha/year) for genotypes and 0.60% (13.69 kg/ha/year) for checks was observed. For elite panel selection, 200 promising lines were selected based on higher breeding values for grain yield and prediction accuracy of > 0.40. The breeding values of the 200 genotypes formulating the core panel ranged between 2366.17 and 4622.59 (kg/ha). A positive genetic rate was observed under all the three conditions; however, the rate of increase was lower than the required rate of 1.5% genetic gain. We propose a recurrent selection breeding strategy within the elite population with the integration of modern tools and technologies to boost the genetic gains in IRRI's drought breeding program. The elite breeding panel identified in this study forms an easily available and highly enriched genetic resource for future recurrent selection programs to boost the genetic gains.

Population-tailored mock genome enables genomic studies in species without a reference genome.

Based on molecular markers, genomic prediction enables us to speed up breeding schemes and increase the response to selection. There are several high-throughput genotyping platforms able to deliver thousands of molecular markers for genomic study purposes. However, even though its widely applied in plant breeding, species without a reference genome cannot fully benefit from genomic tools and modern breeding schemes. We used a method to assemble a population-tailored mock genome to call single-nucleotide polymorphism (SNP) markers without an available reference genome, and for the first time, we compared the results with standard genotyping platforms (array and genotyping-by-sequencing (GBS) using a reference genome) for performance in genomic prediction models. Our results indicate that using a population-tailored mock genome to call SNP delivers reliable estimates for the genomic relationship between genotypes. Furthermore, genomic prediction estimates were comparable to standard approaches, especially when considering only additive effects. However, mock genomes were slightly worse than arrays at predicting traits influenced by dominance effects, but still performed as well as standard GBS methods that use a reference genome. Nevertheless, the array-based SNP markers methods achieved the best predictive ability and reliability to estimate variance components. Overall, the mock genomes can be a worthy alternative for genomic selection studies, especially for those species where the reference genome is not available.

Steady agronomic and genetic interventions are essential for sustaining productivity in intensive rice cropping.

Intensive systems with two or three rice (Oryza sativa L.) crops per year account for about 50% of the harvested area for irrigated rice in Asia. Any reduction in productivity or sustainability of these systems has serious implications for global food security. Rice yield trends in the world's longest-running long-term continuous cropping experiment (LTCCE) were evaluated to investigate consequences of intensive cropping and to draw lessons for sustaining production in Asia. Annual production was sustained at a steady level over the 50-y period in the LTCCE through continuous adjustment of management practices and regular cultivar replacement. Within each of the three annual cropping seasons (dry, early wet, and late wet), yield decline was observed during the first phase, from 1968 to 1990. Agronomic improvements in 1991 to 1995 helped to reverse this yield decline, but yield increases did not continue thereafter from 1996 to 2017. Regular genetic and agronomic improvements were sufficient to maintain yields at steady levels in dry and early wet seasons despite a reduction in the yield potential due to changing climate. Yield declines resumed in the late wet season. Slower growth in genetic gain after the first 20 y was associated with slower breeding cycle advancement as indicated by pedigree depth. Our findings demonstrate that through adjustment of management practices and regular cultivar replacement, it is possible to sustain a high level of annual production in irrigated systems under a changing climate. However, the system was unable to achieve further increases in yield required to keep pace with the growing global rice demand.

Enviromic Assembly Increases Accuracy and Reduces Costs of the Genomic Prediction for Yield Plasticity in Maize.

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.

Introgression of Maize Diversity for Drought Tolerance: Subtropical Maize Landraces as Source of New Positive Variants.

Current climate change models predict an increased frequency and intensity of drought for much of the developing world within the next 30 years. These events will negatively affect maize yields, potentially leading to economic and social instability in many smallholder farming communities. Knowledge about the genetic resources available for traits related to drought tolerance has great importance in developing breeding program strategies. The aim of this research was to study a maize landrace introgression panel to identify chromosomal regions associated with a drought tolerance index. For that, we performed Genome-Wide Association Study (GWAS) on 1326 landrace progenies developed by the CIMMYT Genetic Resources Program, originating from 20 landraces populations collected in arid regions. Phenotypic data were obtained from early testcross trials conducted in three sites and two contrasting irrigation environments, full irrigation (well-watered) and reduced irrigation (drought). The populations were genotyped using the DArTSeq® platform, and a final set of 5,695 SNPs markers was used. The genotypic values were estimated using spatial adjustment in a two-stage analysis. First, we performed the individual analysis for each site/irrigation treatment combination. The best linear unbiased estimates (BLUEs) were used to calculate the Harmonic Mean of Relative Performance (HMRP) as a drought tolerance index for each testcross. The second stage was a joint analysis, which was performed using the HMRP to obtain the best linear unbiased predictions (BLUPs) of the index for each genotype. Then, GWAS was performed to determine the marker-index associations and the marker-Grain Yield (GY) associations for the two irrigation treatments. We detected two significant markers associated with the drought-tolerance index, four associated with GY in drought condition, and other four associated with GY in irrigated conditions each. Although each of these markers explained less than 0.1% of the phenotypic variation for the index and GY, we found two genes likely related to the plant response to drought stress. For these markers, alleles from landraces provide a slightly higher yield under drought conditions. Our results indicate that the positive diversity delivered by landraces are still present on the backcrosses and this is a potential breeding strategy for improving maize for drought tolerance and for trait introgression bringing new superior allelic diversity from landraces to breeding populations.

Optimizing Genomic-Enabled Prediction in Small-Scale Maize Hybrid Breeding Programs: A Roadmap Review.

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.