Enhanced radiation use efficiency and grain filling rate as the main drivers of grain yield genetic gains in the CIMMYT elite spring wheat yield trial.

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 < 0.01), 0.074 g m-2 year-1 (p < 0.05), and 0.017 g MJ-1 year-1 (p < 0.001) for BM, GFR, and RUE_total, respectively. Grain weight per spike exhibited a positive but no significant trend (0.014 g year-1, p = 0.07), whereas a negative tendency for DTH was observed (- 0.43 days year-1, p < 0.001). Analysis of the top ten highest-yielding genotypes revealed differential GY-associated trait contributions, demonstrating that improved GY can be attained through different mechanisms and indicating that no single trait criterion is adopted by CIMMYT breeders for developing new superior lines. We conclude that CIMMYT's Bread Wheat Breeding Program has continued to deliver adapted and more productive wheat genotypes to National partners worldwide, mainly driven by enhancing RUE_total and GFR and that future yield increases could be achieved by intercrossing genetically diverse top performer genotypes.

Multi-modal deep learning improves grain yield prediction in wheat breeding by fusing genomics and phenomics.

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

Sparse kernel models provide optimization of training set design for genomic prediction in multiyear wheat breeding data.

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.

Dissecting the Genetic Architecture of Phenology Affecting Adaptation of Spring Bread Wheat Genotypes to the Major Wheat-Producing Zones in India.

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

Genome-Wide Association Mapping Identifies Key Genomic Regions for Grain Zinc and Iron Biofortification in Bread Wheat.

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

Genome-Wide Association Mapping Indicates Quantitative Genetic Control of Spot Blotch Resistance in Bread Wheat and the Favorable Effects of Some Spot Blotch Loci on Grain Yield.

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

Juvenile Heat Tolerance in Wheat for Attaining Higher Grain Yield by Shifting to Early Sowing in October in South Asia.

Farmers in northwestern and central India have been exploring to sow their wheat much earlier (October) than normal (November) to sustain productivity by escaping terminal heat stress and to utilize the available soil moisture after the harvesting of rice crop. However, current popular varieties are poorly adapted to early sowing due to the exposure of juvenile plants to the warmer temperatures in the month of October and early November. Therefore, a study was undertaken to identify wheat genotypes suited to October sowing under warmer temperatures in India. A diverse collection of 3322 bread wheat varieties and elite lines was prepared in CIMMYT, Mexico, and planted in the 3rd week of October during the crop season 2012-2013 in six locations (Ludhiana, Karnal, New Delhi, Indore, Pune and Dharwad) spread over northwestern plains zone (NWPZ) and central and Peninsular zone (CZ and PZ; designated as CPZ) of India. Agronomic traits data from the seedling stage to maturity were recorded. Results indicated substantial diversity for yield and yield-associated traits, with some lines showing indications of higher yields under October sowing. Based on agronomic performance and disease resistance, the top 48 lines (and two local checks) were identified and planted in the next crop season (2013-2014) in a replicated trial in all six locations under October sowing (third week). High yielding lines that could tolerate higher temperature in October sowing were identified for both zones; however, performance for grain yield was more promising in the NWPZ. Hence, a new trial of 30 lines was planted only in NWPZ under October sowing. Lines showing significantly superior yield over the best check and the most popular cultivars in the zone were identified. The study suggested that agronomically superior wheat varieties with early heat tolerance can be obtained that can provide yield up to 8 t/ha by planting in the third to fourth week of October.

Genome-enabled prediction for sparse testing in multi-environmental wheat trials.

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.

Harnessing translational research in wheat for climate resilience.

Despite being the world's most widely grown crop, research investments in wheat (Triticum aestivum and Triticum durum) fall behind those in other staple crops. Current yield gains will not meet 2050 needs, and climate stresses compound this challenge. However, there is good evidence that heat and drought resilience can be boosted through translating promising ideas into novel breeding technologies using powerful new tools in genetics and remote sensing, for example. Such technologies can also be applied to identify climate resilience traits from among the vast and largely untapped reserve of wheat genetic resources in collections worldwide. This review describes multi-pronged research opportunities at the focus of the Heat and Drought Wheat Improvement Consortium (coordinated by CIMMYT), which together create a pipeline to boost heat and drought resilience, specifically: improving crop design targets using big data approaches; developing phenomic tools for field-based screening and research; applying genomic technologies to elucidate the bases of climate resilience traits; and applying these outputs in developing next-generation breeding methods. The global impact of these outputs will be validated through the International Wheat Improvement Network, a global germplasm development and testing system that contributes key productivity traits to approximately half of the global wheat-growing area.

High-resolution spectral information enables phenotyping of leaf epicuticular wax in wheat.

BACKGROUND: Epicuticular wax (EW) is the first line of defense in plants for protection against biotic and abiotic factors in the environment. In wheat, EW is associated with resilience to heat and drought stress, however, the current limitations on phenotyping EW restrict the integration of this secondary trait into wheat breeding pipelines. In this study we evaluated the use of light reflectance as a proxy for EW load and developed an efficient indirect method for the selection of genotypes with high EW density. RESULTS: Cuticular waxes affect the light that is reflected, absorbed and transmitted by plants. The narrow spectral regions statistically associated with EW overlap with bands linked to photosynthetic radiation (500 nm), carotenoid absorbance (400 nm) and water content (~ 900 nm) in plants. The narrow spectral indices developed predicted 65% (EWI-13) and 44% (EWI-1) of the variation in this trait utilizing single-leaf reflectance. However, the normalized difference indices EWI-4 and EWI-9 improved the phenotyping efficiency with canopy reflectance across all field experimental trials. Indirect selection for EW with EWI-4 and EWI-9 led to a selection efficiency of 70% compared to phenotyping with the chemical method. The regression model EWM-7 integrated eight narrow wavelengths and accurately predicted 71% of the variation in the EW load (mg·dm-2) with leaf reflectance, but under field conditions, a single-wavelength model consistently estimated EW with an average RMSE of 1.24 mg·dm-2 utilizing ground and aerial canopy reflectance. CONCLUSIONS: Overall, the indices EWI-1, EWI-13 and the model EWM-7 are reliable tools for indirect selection for EW based on leaf reflectance, and the indices EWI-4, EWI-9 and the model EWM-1 are reliable for selection based on canopy reflectance. However, further research is needed to define how the background effects and geometry of the canopy impact the accuracy of these phenotyping methods.

Target Population of Environments for Wheat Breeding in India: Definition, Prediction and Genetic Gains.

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.

Elucidating the genetics of grain yield and stress-resilience in bread wheat using a large-scale genome-wide association mapping study with 55,568 lines.

Wheat grain yield (GY) improvement using genomic tools is important for achieving yield breakthroughs. To dissect the genetic architecture of wheat GY potential and stress-resilience, we have designed this large-scale genome-wide association study using 100 datasets, comprising 105,000 GY observations from 55,568 wheat lines evaluated between 2003 and 2019 by the International Maize and Wheat Improvement Center and national partners. We report 801 GY-associated genotyping-by-sequencing markers significant in more than one dataset and the highest number of them were on chromosomes 2A, 6B, 6A, 5B, 1B and 7B. We then used the linkage disequilibrium (LD) between the consistently significant markers to designate 214 GY-associated LD-blocks and observed that 84.5% of the 58 GY-associated LD-blocks in severe-drought, 100% of the 48 GY-associated LD-blocks in early-heat and 85.9% of the 71 GY-associated LD-blocks in late-heat, overlapped with the GY-associated LD-blocks in the irrigated-bed planting environment, substantiating that simultaneous improvement for GY potential and stress-resilience is feasible. Furthermore, we generated the GY-associated marker profiles and analyzed the GY favorable allele frequencies for a large panel of 73,142 wheat lines, resulting in 44.5 million datapoints. Overall, the extensive resources presented in this study provide great opportunities to accelerate breeding for high-yielding and stress-resilient wheat varieties.

Haplotype-Based, Genome-Wide Association Study Reveals Stable Genomic Regions for Grain Yield in CIMMYT Spring Bread Wheat.

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

Improved Accuracy of High-Throughput Phenotyping From Unmanned Aerial Systems by Extracting Traits Directly From Orthorectified Images.

The development of high-throughput genotyping and phenotyping has provided access to many tools to accelerate plant breeding programs. Unmanned Aerial Systems (UAS)-based remote sensing is being broadly implemented for field-based high-throughput phenotyping due to its low cost and the capacity to rapidly cover large breeding populations. The Structure-from-Motion photogrammetry processes aerial images taken from multiple perspectives over a field to an orthomosaic photo of a complete field experiment, allowing spectral or morphological trait extraction from the canopy surface for each individual field plot. However, some phenotypic information observable in each raw aerial image seems to be lost to the orthomosaic photo, probably due to photogrammetry processes such as pixel merging and blending. To formally assess this, we introduced a set of image processing methods to extract phenotypes from orthorectified raw aerial images and compared them to the negative control of extracting the same traits from processed orthomosaic images. We predict that standard measures of accuracy in terms of the broad-sense heritability of the remote sensing spectral traits will be higher using the orthorectified photos than with the orthomosaic image. Using three case studies, we therefore compared the broad-sense heritability of phenotypes in wheat breeding nurseries including, (1) canopy temperature from thermal imaging, (2) canopy normalized difference vegetation index (NDVI), and (3) early-stage ground cover from multispectral imaging. We evaluated heritability estimates of these phenotypes extracted from multiple orthorectified aerial images via four statistical models and compared the results with heritability estimates of these phenotypes extracted from a single orthomosaic image. Our results indicate that extracting traits directly from multiple orthorectified aerial images yielded increased estimates of heritability for all three phenotypes through proper modeling, compared to estimation using traits extracted from the orthomosaic image. In summary, the image processing methods demonstrated in this study have the potential to improve the quality of the plant trait extracted from high-throughput imaging. This, in turn, can enable breeders to utilize phenomics technologies more effectively for improved selection.

Genomic Selection for Grain Yield in the CIMMYT Wheat Breeding Program-Status and Perspectives.

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

Breeder friendly phenotyping.

The word phenotyping can nowadays invoke visions of a drone or phenocart moving swiftly across research plots collecting high-resolution data sets on a wide array of traits. This has been made possible by recent advances in sensor technology and data processing. Nonetheless, more comprehensive often destructive phenotyping still has much to offer in breeding as well as research. This review considers the 'breeder friendliness' of phenotyping within three main domains: (i) the 'minimum data set', where being 'handy' or accessible and easy to collect and use is paramount, visual assessment often being preferred; (ii) the high throughput phenotyping (HTP), relatively new for most breeders, and requiring significantly greater investment with technical hurdles for implementation and a steeper learning curve than the minimum data set; (iii) detailed characterization or 'precision' phenotyping, typically customized for a set of traits associated with a target environment and requiring significant time and resources. While having been the subject of debate in the past, extra investment for phenotyping is becoming more accepted to capitalize on recent developments in crop genomics and prediction models, that can be built from the high-throughput and detailed precision phenotypes. This review considers different contexts for phenotyping, including breeding, exploration of genetic resources, parent building and translational research to deliver other new breeding resources, and how the different categories of phenotyping listed above apply to each. Some of the same tools and rules of thumb apply equally well to phenotyping for genetic analysis of complex traits and gene discovery.

Regularized selection indices for breeding value prediction using hyper-spectral image data.

High-throughput phenotyping (HTP) technologies can produce data on thousands of phenotypes per unit being monitored. These data can be used to breed for economically and environmentally relevant traits (e.g., drought tolerance); however, incorporating high-dimensional phenotypes in genetic analyses and in breeding schemes poses important statistical and computational challenges. To address this problem, we developed regularized selection indices; the methodology integrates techniques commonly used in high-dimensional phenotypic regressions (including penalization and rank-reduction approaches) into the selection index (SI) framework. Using extensive data from CIMMYT's (International Maize and Wheat Improvement Center) wheat breeding program we show that regularized SIs derived from hyper-spectral data offer consistently higher accuracy for grain yield than those achieved by standard SIs, and by vegetation indices commonly used to predict agronomic traits. Regularized SIs offer an effective approach to leverage HTP data that is routinely generated in agriculture; the methodology can also be used to conduct genetic studies using high-dimensional phenotypes that are often collected in humans and model organisms including body images and whole-genome gene expression profiles.

Grain yield genetic gains and changes in physiological related traits for CIMMYT's High Rainfall Wheat Screening Nursery tested across international environments.

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

Incorporating Genome-Wide Association Mapping Results Into Genomic Prediction Models for Grain Yield and Yield Stability in CIMMYT Spring Bread Wheat.

Untangling the genetic architecture of grain yield (GY) and yield stability is an important determining factor to optimize genomics-assisted selection strategies in wheat. We conducted in-depth investigation on the above using a large set of advanced bread wheat lines (4,302), which were genotyped with genotyping-by-sequencing markers and phenotyped under contrasting (irrigated and stress) environments. Haplotypes-based genome-wide-association study (GWAS) identified 58 associations with GY and 15 with superiority index Pi (measure of stability). Sixteen associations with GY were "environment-specific" with two on chromosomes 3B and 6B with the large effects and 8 associations were consistent across environments and trials. For Pi, 8 associations were from chromosomes 4B and 7B, indicating 'hot spot' regions for stability. Epistatic interactions contributed to an additional 5-9% variation on average. We further explored whether integrating consistent and robust associations identified in GWAS as fixed effects in prediction models improves prediction accuracy. For GY, the model accounting for the haplotype-based GWAS loci as fixed effects led to up to 9-10% increase in prediction accuracy, whereas for Pi this approach did not provide any advantage. This is the first report of integrating genetic architecture of GY and yield stability into prediction models in wheat.

Validation of Candidate Gene-Based Markers and Identification of Novel Loci for Thousand-Grain Weight in Spring Bread Wheat.

Increased thousand-grain weight (TGW) is an important breeding target for indirectly improving grain yield (GY). Fourteen reported candidate genes known to enhance TGW were evaluated in two independent and existing datasets of wheat at CIMMYT, the Elite Yield Trial (EYT) from 2015 to 2016 (EYT2015-16) and the Wheat Association Mapping Initiative (WAMI) panel, to study their allele effects on TGW and to verify their suitability for marker-assisted selection. Of these, significant associations were detected for only one gene (TaGs3-D1) in the EYT2015-16 and two genes (TaTGW6 and TaSus1) in WAMI. The reported favorable alleles of TaGs3-D1 and TaTGW6 genes decreased TGW in the datasets. A haplotype-based genome wide association study was implemented to identify the genetic determinants of TGW on a large set of CIMMYT germplasm (4,302 lines comprising five EYTs), which identified 15 haplotype blocks to be significantly associated with TGW. Four of them, identified on chromosomes 4A, 6A, and 7A, were associated with TGW in at least three EYTs. The locus on chromosome 6A (Hap-6A-13) had the largest effect on TGW and additionally GY with increases of up to 2.60 g and 258 kg/ha, respectively. Discovery of novel TGW loci described in our study expands the opportunities for developing diagnostic markers and for multi-gene pyramiding to derive new allele combinations for enhanced TGW and GY in CIMMYT wheat.