Meta-QTL analysis in wheat: progress, challenges and opportunities.

Wheat, an important cereal crop globally, faces major challenges due to increasing global population and changing climates. The production and productivity are challenged by several biotic and abiotic stresses. There is also a pressing demand to enhance grain yield and quality/nutrition to ensure global food and nutritional security. To address these multifaceted concerns, researchers have conducted numerous meta-QTL (MQTL) studies in wheat, resulting in the identification of candidate genes that govern these complex quantitative traits. MQTL analysis has successfully unraveled the complex genetic architecture of polygenic quantitative traits in wheat. Candidate genes associated with stress adaptation have been pinpointed for abiotic and biotic traits, facilitating targeted breeding efforts to enhance stress tolerance. Furthermore, high-confidence candidate genes (CGs) and flanking markers to MQTLs will help in marker-assisted breeding programs aimed at enhancing stress tolerance, yield, quality and nutrition. Functional analysis of these CGs can enhance our understanding of intricate trait-related genetics. The discovery of orthologous MQTLs shared between wheat and other crops sheds light on common evolutionary pathways governing these traits. Breeders can leverage the most promising MQTLs and CGs associated with multiple traits to develop superior next-generation wheat cultivars with improved trait performance. This review provides a comprehensive overview of MQTL analysis in wheat, highlighting progress, challenges, validation methods and future opportunities in wheat genetics and breeding, contributing to global food security and sustainable agriculture.

Introgression of Heterotic Genomic Segments from Brassica carinata into Brassica juncea for Enhancing Productivity.

Interspecific hybridization resulted in the creation of B. juncea introgression lines (ILs) generated from B. carinata with increased productivity and adaptability. Forty ILs were crossed with their respective B. juncea recipient parents to generate introgression line hybrids (ILHs) and the common tester (SEJ 8) was used to generate test hybrids (THs). Mid-parent heterosis in ILHs and standard heterosis in THs were calculated for eight yield and yield-related traits. Heterotic genomic regions were dissected using ten ILs with significant mid-parent heterosis in ILHs and standard heterosis in THs for seed yield. A high level of heterosis for seed yield was contributed by 1000 seed weight (13.48%) in D31_ILHs and by total siliquae/plant (14.01%) and siliqua length (10.56%) in PM30_ILHs. The heterotic ILs of DRMRIJ 31 and Pusa Mustard 30 were examined using polymorphic SNPs between the parents, and a total of 254 and 335 introgressed heterotic segments were identified, respectively. This investigation discovered potential genes, viz., PUB10, glutathione S transferase, TT4, SGT, FLA3, AP2/ERF, SANT4, MYB, and UDP-glucosyl transferase 73B3 that were previously reported to regulate yield-related traits. The heterozygosity of the FLA3 gene significantly improved siliqua length and seeds per siliqua in ILHs of Pusa Mustard 30. This research proved that interspecific hybridization is an effective means of increasing the diversity of cultivated species by introducing new genetic variants and improving the level of heterosis.

Spring Wheat's Ability to Utilize Nitrogen More Effectively Is Influenced by Root Phene Variation.

Genetic improvement for nitrogen use efficiency (NUE) can play a very crucial role in sustainable agriculture. Root traits have hardly been explored in major wheat breeding programs, more so in spring germplasm, largely because of the difficulty in their scoring. A total of 175 advanced/improved Indian spring wheat genotypes were screened for root traits and nitrogen uptake and nitrogen utilization at varying nitrogen levels in hydroponic conditions to dissect the complex NUE trait into its component traits and to study the extent of variability that exists for those traits in Indian germplasm. Analysis of genetic variance showed a considerable amount of genetic variability for nitrogen uptake efficiency (NUpE), nitrogen utilization efficiency (NUtE), and most of the root and shoot traits. Improved spring wheat breeding lines were found to have very large variability for maximum root length (MRL) and root dry weights (RDW) with strong genetic advance. In contrast to high nitrogen (HN), a low nitrogen (LN) environment was more effective in differentiating wheat genotypes for NUE and its component traits. Shoot dry weight (SDW), RDW, MRL, and NUpE were found to have a strong association with NUE. Further study revealed the role of root surface area (RSA) and total root length (TRL) in RDW formation as well as in nitrogen uptake and therefore can be targeted for selection to further the genetic gain for grain yield under high input or sustainable agriculture under limited inputs.

Genome-wide association studies reveal putative QTLs for physiological traits under contrasting phosphorous conditions in wheat (Triticum aestivum L.).

A Genome-wide association (GWAS) study was conducted for phosphorous (P)-use responsive physiological traits in bread wheat at the seedling stage under contrasting P regimes. A panel of 158 diverse advanced breeding lines and released varieties, and a set of 10,800 filtered single nucleotide polymorphism (SNP) markers were used to study marker-trait associations over the eight shoot traits. Principle component analysis separated the two environments (P regimes) because of the differential response of the traits indicating the essentiality of the separate breeding programmes for each environment. Significant variations for genotypic, environmental, and genotype × environment (GEI) effects were observed for all the traits in the combined analysis of variance with moderately high broad sense heritability traits (0.50-0.73). With the different algorithms of association mapping viz., BLINK, FarmCPU, and MLM, 38 unique QTLs under non-limiting P (NLP) and 45 QTLs for limiting P (LP) conditions for various shoot traits were identified. Some of these QTLs were captured by all three algorithms. Interestingly, a Q.iari.dt.sdw.1 on chromosome 1D was found to explain the significant variations in three important physiological traits under non-limiting phosphorus (NLP) conditions. We identified the putative candidate genes for QTLs namely Q.iari.dt.chl.1, Q.iari.dt.sdw.16, Q.iari.dt.sdw.9 and Q.iari.dt.tpc.1 which are potentially involved in the mechanism regulating phosphorus use efficiency through improved P absorption due to improved root architectural traits and better mobilization such as sulfotransferase involved in postembryonic root development, WALLS ARE THIN1 (WAT1), a plant-specific protein that facilitates auxin export; lectin receptor-like kinase essentially involved in plant development, stress response during germination and lateral root development and F-box component of the SKP-Cullin-F box E3 ubiquitin ligase complex and strigolactone signal perception. Expression profiling of putative genes located in identified genomic regions against the wheat expression atlas revealed their significance based on the expression of these genes for stress response and growth development processes in wheat. Our results thus provide an important insight into understanding the genetic basis for improving PUE under phosphorus stress conditions and can shape the future breeding programme by developing and integrating molecular markers for these difficult-to-score important traits.

Deciphering the environmental impact on spike architectural traits for grain yield consolidation in bread wheat (T. aestivum L.).

The realization of grain yield in wheat is decided by source-sink balance under prevailing environmental conditions. Management conditions like changing the sowing time influence the source-sink capacity through modification in agronomic traits. Therefore, this experiment was conducted to decipher the influence of spike architectural traits (SATs) on grain yield and to open avenues for further grain yield enhancement. Comparatively early sowing over timely sowing gives the advantage of realizing higher grain yield with a positive relationship with SATs namely spike length, spikelets per spike, individual spike weight, individual grain weight, number of grains per spikelet, grain length, and grain width of upper and lower spike portion. Confirmatory factorial analysis revealed that spike length, spikelets per spike, individual spike weight, grains per spikelet were having a significant effect in deciding grain yield in early sown. The presence of a significant effect of genotype by environment interaction over grain yield and SATs allows the exploitation of available genotypic and environmental variability for further yield enhancement. GGE analysis on transformed and standardized grain yield-trait (GY-trait) combinations was used in the selection of genotypes having high GY-trait combinations for both sowing times. In early sowing, WG 11 was the best for high GY with high individual spike weight; grain length and grain width at lower and upper parts of the spike; and shorter days to 50% flowering. Genotypes exclusively having the high GY-trait combination along with low values of remaining GY-trait combinations were also selected with genotype focused GGE approach.

Genetic Gain in Yield and Associated Changes in Agronomic Traits in Wheat Cultivars Developed Between 1900 and 2016 for Irrigated Ecosystems of Northwestern Plain Zone of India.

Knowledge about the yield gain over the years due to associated changes in the yield component traits is essential for a critical understanding of yield-limiting factors. To estimate genetic gain in grain yield (GY) and component agronomic traits of wheat varieties released between 1900 and 2016 for northwestern plain zone (NWPZ) of India and to identify agronomic and/or genetic basis of the realized gains, two sets of wheat varieties comprising mega varieties and two recently developed varieties were evaluated under timely sown, tilled, and early sown conservation agriculture (CA) conditions for four consecutive years under irrigated conditions. The average annual genetic gain in GY since 1,905 under timely sown irrigated conditions was found to be 0.544% yr-1 over the average of all varieties and 0.822% yr-1 (24.27 kg ha-1 yr-1) over the first released variety, NP4. The realized mean yield increased from 2,950 kg ha-1 of the variety NP4 released in 1,905-5,649 kg ha-1 of HD3086 released in 2014. Regression analysis revealed a linear reduction in height and peduncle length (PL) over the years with a simultaneous and linear increase in biomass at the rate of 43.9 kg ha-1 yr-1 or relatively at 0.368% yr-1 mainly because of delayed heading and increased crop duration. Regression analysis showed no linear trend for tiller number and thousand-grain weight (TGW). Though harvest index (HI) was found to linearly increase relatively at the rate of 0.198% per annum, polynomial regression improved the fitness of data with the indication of no increase in HI since 1982. Interestingly, genetic gain evaluation under early sown CA conditions for 4 years showed similar relative gain (RG) [a relative improvement in varieties across breeding periods (BP)] (0.544% yr-1) but with a higher absolute value (29.28 kg ha-1 yr-1). Major mega varieties like Kalyan Sona, HD2009, PBW 343, HD2967, and HD3086, which occupied a comparatively larger area, were found highly plastic to the improvements in the production environment under timely sown conditions.

Deciphering the change in root system architectural traits under limiting and non-limiting phosphorus in Indian bread wheat germplasm.

The root system architectures (RSAs) largely decide the phosphorus use efficiency (PUE) of plants by influencing the phosphorus uptake. Very limited information is available on wheat's RSAs and their deciding factors affecting phosphorus uptake efficiency (PupE) due to difficulties in adopting scoring values used for evaluating root traits. Based on our earlier research experience on nitrogen uptake efficiency screening under, hydroponics and soil-filled pot conditions, a comprehensive study on 182 Indian bread wheat genotypes was carried out under hydroponics with limited P (LP) and non-limiting P (NLP) conditions. The findings revealed a significant genetic variation, root traits correlation, and moderate to high heritability for RSAs traits namely primary root length (PRL), total root length (TRL), total root surface area (TSA), root average diameter (RAD), total root volume (TRV), total root tips (TRT) and total root forks (TRF). In LP, the expressions of TRL, TRV, TSA, TRT and TRF were enhanced while PRL and RAD were diminished. An almost similar pattern of correlations among the RSAs was also observed in both conditions except for RAD. RAD exhibited significant negative correlations with PRL, TRL, TSA, TRT and TRF under LP (r = -0.45, r = -0.35, r = -0.16, r = -0.30, and r = -0.28 respectively). The subclass of TRL, TSA, TRV and TRT representing the 0-0.5 mm diameter had a higher root distribution percentage in LP than NLP. Comparatively wide range of H' value i.e. 0.43 to 0.97 in LP than NLP indicates that expression pattern of these traits are highly influenced by the level of P. In which, RAD (0.43) expression was reduced in LP, and expressions of TRF (0.91) and TSA (0.97) were significantly enhanced. The principal component analysis for grouping of traits and genotypes over LP and NLP revealed a high PC1 score indicating the presence of non-crossover interactions. Based on the comprehensive P response index value (CPRI value), the top five highly P efficient wheat genotypes namely BW 181, BW 103, BW 104, BW 143 and BW 66, were identified. Considering the future need for developing resource-efficient wheat varieties, these genotypes would serve as valuable genetic sources for improving P efficiency in wheat cultivars. This set of genotypes would also help in understanding the genetic architecture of a complex trait like P use efficiency.

Enhancing the Nutritional Quality of Major Food Crops Through Conventional and Genomics-Assisted Breeding.

Nutritional stress is making over two billion world population malnourished. Either our commercially cultivated varieties of cereals, pulses, and oilseed crops are deficient in essential nutrients or the soils in which these crops grow are becoming devoid of minerals. Unfortunately, our major food crops are poor sources of micronutrients required for normal human growth. To overcome the problem of nutritional deficiency, greater emphasis should be laid on the identification of genes/quantitative trait loci (QTLs) pertaining to essential nutrients and their successful deployment in elite breeding lines through marker-assisted breeding. The manuscript deals with information on identified QTLs for protein content, vitamins, macronutrients, micro-nutrients, minerals, oil content, and essential amino acids in major food crops. These QTLs can be utilized in the development of nutrient-rich crop varieties. Genome editing technologies that can rapidly modify genomes in a precise way and will directly enrich the nutritional status of elite varieties could hold a bright future to address the challenge of malnutrition.

Application of Genomics Tools in Wheat Breeding to Attain Durable Rust Resistance.

Wheat is an important source of dietary protein and calories for the majority of the world's population. It is one of the largest grown cereal in the world occupying over 215 M ha. Wheat production globally is challenged by biotic stresses such as pests and diseases. Of the 50 diseases of wheat that are of economic importance, the three rust diseases are the most ubiquitous causing significant yield losses in the majority of wheat production environments. Under severe epidemics they can lead to food insecurity threats amid the continuous evolution of new races of the pathogens, shifts in population dynamics and their virulence patterns, thereby rendering several effective resistance genes deployed in wheat breeding programs vulnerable. This emphasizes the need to identify, characterize, and deploy effective rust-resistant genes from diverse sources into pre-breeding lines and future wheat varieties. The use of genetic resistance has been marked as eco-friendly and to curb the further evolution of rust pathogens. Deployment of multiple rust resistance genes including major and minor genes in wheat lines could enhance the durability of resistance thereby reducing pathogen evolution. Advances in next-generation sequencing (NGS) platforms and associated bioinformatics tools have revolutionized wheat genomics. The sequence alignment of the wheat genome is the most important landmark which will enable genomics to identify marker-trait associations, candidate genes and enhanced breeding values in genomic selection (GS) studies. High throughput genotyping platforms have demonstrated their role in the estimation of genetic diversity, construction of the high-density genetic maps, dissecting polygenic traits, and better understanding their interactions through GWAS (genome-wide association studies) and QTL mapping, and isolation of R genes. Application of breeder's friendly KASP assays in the wheat breeding program has expedited the identification and pyramiding of rust resistance alleles/genes in elite lines. The present review covers the evolutionary trends of the rust pathogen and contemporary wheat varieties, and how these research strategies galvanized to control the wheat killer genus Puccinia. It will also highlight the outcome and research impact of cost-effective NGS technologies and cloning of rust resistance genes amid the public availability of common and tetraploid wheat reference genomes.

Yield-enhancing heterotic QTL transferred from wild species to cultivated rice Oryza sativa L.

Utilization of "hidden genes" from wild species has emerged as a novel option for enrichment of genetic diversity for productivity traits. In rice we have generated more than 2000 lines having introgression from 'A' genome-donor wild species of rice in the genetic background of popular varieties PR114 and Pusa44 were developed. Out of these, based on agronomic acceptability, 318 lines were used for developing rice hybrids to assess the effect of introgressions in heterozygous state. These introgression lines and their recurrent parents, possessing fertility restoration ability for wild abortive (WA) cytoplasm, were crossed with cytoplasmic male sterile (CMS) line PMS17A to develop hybrids. Hybrids developed from recurrent parents were used as checks to compare the performance of 318 hybrids developed by hybridizing alien introgression lines with PMS17A. Seventeen hybrids expressed a significant increase in yield and its component traits over check hybrids. These 17 hybrids were re-evaluated in large-size replicated plots. Of these, four hybrids, viz., ILH299, ILH326, ILH867 and ILH901, having introgressions from O. rufipogon and two hybrids (ILH921 and ILH951) having introgressions from O. nivara showed significant heterosis over parental introgression line, recurrent parents and check hybrids for grain yield-related traits. Alien introgressions were detected in the lines taken as male parents for developing six superior hybrids, using a set of 100 polymorphic simple sequence repeat (SSR) markers. Percent introgression showed a range of 2.24 from in O. nivara to 7.66 from O. rufipogon. The introgressed regions and their putative association with yield components in hybrids is reported and discussed.