Accessions Of Emmer Wheat Research Articles

Evaluation of Genetic Diversity in Wild Emmer Wheat (Triticum dicoccoides) and Durum Wheat (Triticum durum) Accessions Using CAAT and SCoT Markers

Wild emmer is one of the progenitors of wheat, with a high genetic potential for breeding. Continuous evaluations of emmer and other progenitor species are necessary for long-term improvement in yield, agronomic, and stress-related traits. For this purpose, genetic diversity and relationships among 43 wild emmer (Triticum dicoccoides) and 5 durum wheat (Triticum durum) accessions were determined using two DNA marker systems, CAAT box-derived polymorphism (CBDP) and start codon targeted (SCoT) markers. CAAT and SCoT markers generated 63 and 76 polymorphic bands, averaging 9 and 7.6 bands per primer, respectively. The discriminating power, effective multiplex ratio, expected heterozygosity, mean heterozygosity, marker index, polymorphism information content, and resolving power parameters obtained for both marker systems showed the high efficiency of these markers in detecting genetic variation in wild emmer and durum wheat. The results showed that CAAT and SCoT markers with average polymorphism are suitable marker systems for detecting genetic variation between a pool of accessions or populations. These markers would be employed for gene-targeted breeding, and the results indicate that genetic analysis with these markers would be practicable for agricultural improvement and development initiatives.

Association mapping of tan spot and septoria nodorum blotch resistance in cultivated emmer wheat.

A total of 65 SNPs associated with resistance to tan spot and septoria nodorum blotch were identified in a panel of 180 cultivated emmer accessions through association mapping Tan spot and septoria nodorum blotch (SNB) are foliar diseases caused by the respective fungal pathogens Pyrenophora tritici-repentis and Parastagonospora nodorum that affect global wheat production. To find new sources of resistance, we evaluated a panel of 180 cultivated emmer wheat (Triticum turgidum ssp. dicoccum) accessions for reactions to four P. tritici-repentis isolates Pti2, 86-124, 331-9 and DW5, two P. nodorum isolate, Sn4 and Sn2000, and four necrotrophic effectors (NEs) produced by the pathogens. About 8-36% of the accessions exhibited resistance to the four P. tritici-repentis isolates, with five accessions demonstrating resistance to all isolates. For SNB, 64% accessions showed resistance to Sn4, 43% to Sn2000 and 36% to both isolates, with Spain (11% accessions) as the most common origin of resistance. To understand the genetic basis of resistance, association mapping was performed using SNP (single nucleotide polymorphism) markers generated by genotype-by-sequencing and the 9K SNP Infinium array. A total of 46 SNPs were significantly associated with tan spot and 19 SNPs with SNB resistance or susceptibility. Six trait loci on chromosome arms 1BL, 3BL, 4AL (2), 6BL and 7AL conferred resistance to two or more isolates. Known NE sensitivity genes for disease development were undetected except Snn5 for Sn2000, suggesting novel genetic factors are controlling host-pathogen interaction in cultivated emmer. The emmer accessions with the highest levels of resistance to the six pathogen isolates (e.g., CItr 14133-1, PI 94634-1 and PI 377672) could serve as donors for tan spot and SNB resistance in wheat breeding programs.

Evaluation and Identification of Powdery Mildew Resistance Genes in Aegilops tauschii and Emmer Wheat Accessions.

Powdery mildew, caused by Blumeria graminis f. sp. tritici (Bgt), is a serious threat to wheat (Triticum aestivum L.) production. Narrow genetic basis of common wheat boosted the demand for diversified donors against powdery mildew. Aegilops tauschii Coss (2n = 2x = DD) and emmer wheat (2n = 4x = AABB), as the ancestor species of common wheat, are important gene donors for genetic improvement of common wheat. In this study, a total of 71 Ae. tauschii and 161 emmer wheat accessions were first evaluated for their powdery mildew resistance using the Bgt isolate E09. Thirty-three Ae. tauschii (46.5%) and 108 emmer wheat accessions (67.1%) were resistant. Then, all these accessions were tested by the diagnostic markers for 21 known Pm genes. The results showed that Pm2 alleles were detected in all the 71 Ae. tauschii and only Pm4 alleles were detected in 20 of 161 emmer wheat accessions. After haplotype analysis, we identified four Pm4 alleles (Pm4a, Pm4b, Pm4d, and Pm4f) in the emmer wheat accessions and three Pm2 alleles (Pm2d, Pm2e, and Pm2g) in the Ae. tauschii. Further resistance spectrum analysis indicated that these resistance accessions displayed different resistance reactions to different Bgt isolates, implying they may have other Pm genes apart from Pm2 and/or Pm4 alleles. Notably, a new Pm2 allele, Pm2S, was identified in Ae. tauschii, which contained a 64-bp deletion in the first exon and formed a new termination site at the 513th triplet of the shifted reading frame compared with reported Pm2 alleles. The phylogenetic tree of Pm2S showed that the kinship of Pm2S was close to Pm2h. To efficiently and accurately detect Pm2S and distinguish with other Pm2 alleles in Ae. tauschii background, a diagnostic marker, YTU-QS-3, was developed, and its effectiveness was verified. This study provided valuable Pm alleles and enriched the genetic diversity of the powdery mildew resistance in wheat improvement.

Identification of robust yield quantitative trait loci derived from cultivated emmer for durum wheat improvement.

Durum wheat (Triticum turgidum ssp. durum L.) is an important world food crop used to make pasta products. Compared to bread wheat (Triticum aestivum L.), fewer studies have been conducted to identify genetic loci governing yield-component traits in durum wheat. A potential source of diversity for durum is its immediate progenitor, cultivated emmer (T. turgidum ssp. dicoccum). We evaluated two biparental populations of recombinant inbred lines (RILs) derived from crosses between the durum lines Ben and Rusty and the cultivated emmer wheat accessions PI 41025 and PI 193883, referred to as the Ben×PI 41025 (BP025) and Rusty×PI 193883 (RP883) RIL populations, respectively. Both populations were evaluated under field conditions in three seasons with an aim to identify quantitative trait loci (QTLs) associated with yield components and seed morphology that were expressed in multiple environments. A total of 44 and 34 multi-environment QTLs were identified in the BP025 and RP883 populations, respectively. As expected, genetic loci known to govern domestication and development were associated with some of the QTLs, but novel QTLs derived from the cultivated emmer parents and associated with yield components including spikelet number, grain weight, and grain size were identified. These QTLs offer new target loci for durum wheat improvement, and toward that goal, we identified five RILs with increased grain weight and size compared to the durum parents. These materials along with the knowledge of stable QTLs and associated markers can help to expedite the development of superior durum varieties.

QTL mapping for kernel-related traits in a durum wheat x T. dicoccum segregating population.

Durum wheat breeding relies on grain yield improvement to meet its upcoming demand while coping with climate change. Kernel size and shape are the determinants of thousand kernel weight (TKW), which is a key component of grain yield, and the understanding of the genetic control behind these traits supports the progress in yield potential. The present study aimed to dissect the genetic network responsible for kernel size components (length, width, perimeter, and area) and kernel shape traits (width-to-length ratio and formcoefficient) as well as their relationships with kernel weight, plant height, and heading date in durum wheat. Quantitative Trait Locus (QTL) mapping was performed on a segregating population of 110 recombinant inbred lines, derived from a cross between the domesticated emmer wheat accession MG5323 and the durum wheat cv. Latino, evaluated in four different environments. A total of 24 QTLs stable across environments were found and further grouped in nine clusters on chromosomes 2A, 2B, 3A, 3B, 4B, 6B, and 7A. Among them, a QTL cluster on chromosome 4B was associated with kernel size traits and TKW, where the parental MG5323 contributed the favorable alleles, highlighting its potential to improve durum wheat germplasm. The physical positions of the clusters, defined by the projection on the T. durum reference genome, overlapped with already known genes (i.e., BIG GRAIN PROTEIN 1 on chromosome 4B). These results might provide genome-based guidance for the efficient exploitation of emmer wheat diversity in wheat breeding, possibly through yield-related molecular markers.

Characterizing Agronomic and Shoot Morphological Diversity across 263 Wild Emmer Wheat Accessions

Wild emmer, the direct progenitor of modern durum and bread wheat, has mostly been studied for grain quality, biotic, and abiotic stress-related traits. Accordingly, it should also have a certain amount of diversity for morphological and agronomic traits. Despite having a high chance of huge diversity, it has not been deeply explored. In the current study, 263 wild emmer accessions collected from different regions of Israel, Turkey, Lebanon, and Syria were characterized for a total of 19 agronomic and shoot morphological traits. Three trials were carried out in Western Australia, which demonstrated a large variation in these traits. The average phenotypic diversity (H’) was 0.91 as quantified by Shannon’s diversity index. A high heritability was recorded for most of the traits, where biomass/plant and yield/plant were identified as the most potential traits. Correlation analysis revealed several significant associations between traits, including significant positive correlation between yield and tiller number, first leaf area, spike length, and biomass/plant. The principal component analysis (PCA) demonstrated that most of the traits contributed to the overall observed variability. The cluster analysis categorized 263 accessions into five clusters on average. On the other hand, accessions were categorized into eight populations based on the collection region and a comparative analysis demonstrated considerable variations between populations for plant height, spike length, and flag leaf area. Despite the low yield, several wild emmer accessions demonstrated superior performance compared to modern bread wheat cultivars, when selection was based on combining yield with multiple traits. These observations indicate that wild emmer contains a broad gene pool for several agronomic and shoot morphological traits, which can be utilized for bread and durum wheat improvement.

Genome-wide association analyses of leaf rust resistance in cultivated emmer wheat.

Fifteen and eleven loci, with most loci being novel, were identified to associate with seedling and adult resistances, respectively, to the durum-specific races of leaf rust pathogen in cultivated emmer. Leaf rust, caused by Puccinia triticina (Pt), constantly threatens durum (Triticum turgidum ssp. durum) and bread wheat (Triticum aestivum) production worldwide. A Pt race BBBQD detected in California in 2009 poses a potential threat to durum production in North America because resistance source to this race is rare in durum germplasm. To find new resistance sources, we assessed a panel of 180 cultivated emmer wheat (Triticum turgidum ssp. dicoccum) accessions for seedling resistance to BBBQD and for adult resistance to a mixture of durum-specific races BBBQJ, CCMSS, and MCDSS in the field, and genotyped the panel using genotype-by-sequencing (GBS) and the 9K SNP (Single Nucleotide Polymorphism) Infinium array. The results showed 24 and nine accessions consistently exhibited seedling and adult resistance, respectively, with two accessions providing resistance at both stages. We performed genome-wide association studies using 46,383 GBS and 4,331 9K SNP markers and identified 15 quantitative trait loci (QTL) for seedling resistance located mostly on chromosomes 2B and 6B, and 11 QTL for adult resistance on 2B, 3B and 6A. Of these QTL, one might be associated with leaf rust resistance (Lr) gene Lr53, and two with the QTL previously reported in durum or hexaploid wheat. The remaining QTL are potentially associated with new Lr genes. Further linkage analysis and gene cloning are necessary to identify the causal genes underlying these QTL. The emmer accessions with high levels of resistance will be useful for developing mapping populations and adapted durum germplasm and varieties with resistance to the durum-specific races.

Identification of stable QTL controlling multiple yield components in a durum × cultivated emmer wheat population under field and greenhouse conditions.

Crop yield gains are needed to keep pace with a growing global population and decreasing resources to produce food. Cultivated emmer wheat is a progenitor of durum wheat and a useful source of genetic variation for trait improvement in durum. Here, we evaluated a recombinant inbred line population derived from a cross between the North Dakota durum wheat variety Divide and the cultivated emmer wheat accession PI 272527 consisting of 219 lines. The population was evaluated in 3 field environments and 2 greenhouse experiments to identify quantitative trait locus associated with 11 yield-related traits that were expressed in a consistent manner over multiple environments. We identified 27 quantitative trait locus expressed in at least 2 field environments, 17 of which were also expressed under greenhouse conditions. Seven quantitative trait locus regions on chromosomes 1B, 2A, 2B, 3A, 3B, 6A, and 7B had pleiotropic effects on multiple yield-related traits. The previously cloned genes Q and FT-B1, which are known to be associated with development and morphology, were found to consistently be associated with multiple traits across environments. PI 272527 contributed beneficial alleles for quantitative trait locus associated with multiple traits, especially for seed morphology quantitative trait locus on chromosomes 1B, 2B, and 6A. Three recombinant inbred lines with increased grain size and weight compared to Divide were identified and demonstrated the potential for improvement of durum wheat through deployment of beneficial alleles from the cultivated emmer parent. The findings from this study provide knowledge regarding stable and robust quantitative trait locus that breeders can use for improving yield in durum wheat.

Emmer Wheat Eco-Geographic and Genomic Congruence Shapes Phenotypic Performance under Mediterranean Climate.

Emmer wheat (Triticum turgidum ssp. dicoccum) is one of the world’s oldest domesticated crops, and it harbors a potentially rich reservoir of agronomic and nutritional quality trait variations. The growing global demand for plant-based health-food niche markets has promoted new commercial interest in ancient grains, including Emmer wheat. Although T. dicoccum can also perform well under harsh environments, its cultivation along the Mediterranean agro-ecosystems is sparse. Here, we analyze a unique tetraploid wheat collection (n = 121) representing a wide geographic range of Emmer accessions, using 9897 DArTseq markers and on-field phenotypic characterization to quantify the extent of diversity among populations and the interactions between eco-geographic, genetic, and phenotypic attributes. Population genomic inferences based on the DArTseq data indicated that the collection could be split into four distinguished clusters in accordance with their eco-geographic origin although significant phenotypic variation was observed within clusters. Superior early vegetative vigor, shorter plant height, and early phenology were observed among emmer wheat accessions from Ethiopia compared to accessions from northern regions. This adaptive advantage highlights the potential of emmer wheat as an exotic germplasm for wheat improvement through breeding. The direct integration of such germplasm into conventional or organic farming agro-systems under the Mediterranean basin climate is also discussed.

Harnessing the diversity of wild emmer wheat for genetic improvement of durum wheat

Key messageThe multiple derivative lines (MDLs) characterized in this study offer a promising strategy for harnessing the diversity of wild emmer wheat for durum and bread wheat improvement.Crop domestication has diminished genetic diversity and reduced phenotypic plasticity and adaptation. Exploring the adaptive capacity of wild progenitors offer promising opportunities to improve crops. We developed a population of 178 BC1F6 durum wheat (Triticum turgidum ssp. durum) lines by crossing and backcrossing nine wild emmer wheat (T. turgidum ssp. dicoccoides) accessions with the common durum wheat cultivar ‘Miki 3’. Here, we describe the development of this population, which we named as multiple derivative lines (MDLs), and demonstrated its suitability for durum wheat breeding. We genotyped the MDL population, the parents, and 43 Sudanese durum wheat cultivars on a Diversity Array Technology sequencing platform. We evaluated days to heading and plant height in Dongola (Sudan) and in Tottori (Japan). The physical map length of the MDL population was 9 939 Mb with an average of 1.4 SNP/Mb. The MDL population had greater diversity than the Sudanese cultivars. We found high gene exchange between the nine wild emmer accessions and the MDL population, indicating that the MDL captured most of the diversity in the wild emmer accessions. Genome-wide association analysis identified three loci for days to heading on chromosomes 1A and 5A in Dongola and one on chromosome 3B in Tottori. For plant height, common genomic loci were found on chromosomes 4A and 4B in both locations, and one genomic locus on chromosome 7B was found only in Dongola. The results revealed that the MDLs are an effective strategy towards harnessing wild emmer wheat diversity for wheat genetic improvement.

Comparison of the Agronomic, Cytological, Grain Protein Characteristics, as Well as Transcriptomic Profile of Two Wheat Lines Derived From Wild Emmer

Two advanced wheat lines BAd7-209 and BAd23-1 without the functional gene GPC-B1 were obtained from a cross between common wheat cultivar Chuannong 16 (CN16) and wild emmer wheat accession D97 (D97). BAd7-209 showed superior quality parameters than those of BAd23-1 and CN16. We found that the components of glutenins and gliadins in BAd7-209 and BAd23-1 were similar, whereas BAd7-209 had higher amount of glutenins and gliadins than those of BAd23-1. RNA sequencing analysis on developing grains of BAd7-209 and BAd23-1 as well as their parents revealed 382 differentially expressed genes (DEGs) between the high–grain protein content (GPC) (D97 + BAd7-209) and the low-GPC (CN16 + BAd23-1) groups. DEGs were mainly associated with transcriptional regulation of the storage protein genes, protein processing in endoplasmic reticulum, and protein export pathways. The upregulated gluten genes and transcription factors (e.g., NAC, MYB, and bZIP) may contribute to the high GPC in BAd7-209. Our results provide insights into the potential regulation pathways underlying wheat grain protein accumulation and contribute to make use of wild emmer for wheat quality improvement.

Fine mapping of powdery mildew resistance gene MlWE74 derived from wild emmer wheat (Triticum turgidum ssp. dicoccoides) in an NBS-LRR gene cluster.

Powdery mildew resistance gene MlWE74, originated from wild emmer wheat accession G-748-M, was mapped in an NBS-LRR gene cluster of chromosome 2BS. Wheat powdery mildew, caused by Blumeria graminis f. sp. tritici (Bgt), is a globally devastating disease. Wild emmer wheat (Triticum turgidum var. dicoccoides) is a valuable genetic resource for improving disease resistance in common wheat. A powdery mildew resistance gene was transferred to hexaploid wheat line WE74 from wild emmer accession G-748-M. Genetic analysis revealed that the powdery mildew resistance in WE74 is controlled by a single dominant gene, herein temporarily designated MlWE74. Bulked segregant analysis (BSA) and molecular mapping delimited MlWE74 to the terminal region of chromosome 2BS flanking by markers WGGBD412 and WGGBH346 within a genetic interval of 0.25cM and corresponding to 799.9kb genomic region in the Zavitan reference sequence. Sequence annotation revealed two phosphoglycerate mutase-like genes, an alpha/beta-hydrolases gene, and five NBS-LRR disease resistance genes that could serve as candidates for map-based cloning of MlWE74. The geographical location analysis indicated that MlWE74 is mainly distributed in Rosh Pinna and Amirim regions, in the northern part of Israel, where environmental conditions are favorable to the occurrence of powdery mildew. Moreover, the co-segregated marker WGGBD425 is helpful in marker-assisted transfer of MlWE74 into elite cultivars.

Genetic variation and genetic control of intraspikelet differences in grain weight and seed dormancy in wild and domesticated emmer wheats.

Seed dormancy, a vital strategy for wild plant species to adapt to an unpredictable environment in their natural habitats, was eliminated from cereals during the domestication process. Intraspikelet differences in grain size and seed dormancy have been observed in wild emmer wheat. To elucidate the genetic variation of these intraspikelet differences and to determine their genetic control, grain weight ratio (first florets/second florets) (GWR), germination rate, and germination index (GI) were analyzed in 67 wild and 82 domesticated emmer wheat accessions, as well as F1 hybrids, F2 populations, and F3-F6 populations derived from reciprocal crosses between wild and domesticated lines. Only the grains on the first florets of two-grained spikelets in wild accessions had varying degrees of dormancy with GI ranging from 0 to 1, which positively correlated with their GWR. This implies that wild emmer populations comprised genotypes with varying degrees of dormancy, including nondormant genotypes. According to segregations observed in F2 populations, the intraspikelet grain weight difference was controlled by two independently inherited loci. Furthermore, low-GWR populations with low or high GI values could be selected in F5 and F6 generations, implying that the major loci associated with dormancy might be independent of intraspikelet grain weight difference.

Geographical distribution and adaptive variation of VRN-A3 alleles in worldwide polyploid wheat (Triticum spp.) species collection.

The distribution of early flowering alleles of VRN-A3 was found to be biased to low latitudes, and these alleles may contribute to environmental adaptability to low latitudes in cultivated emmer wheat. In wheat (Triticum spp.), the flowering time is an important trait for successful seed production and yield by adapting to the regional environment. An early flowering allele of VRN-A3 with 7- and 25-bp insertions in the promoter region (Vrn-A3a-h1) has recently been reported from the analysis of an emmer wheat (Triticum turgidum L. ssp. dicoccum) accession, TN26. This early flowering allele of VRN-A3 might be associated with the regional adaptation of wheat. In this study, we elucidated its geographic distribution to assess the importance of the early flowering allele of VRN-A3 in worldwide wheat collection. From sequence analysis, we identified six VRN-A3 alleles with the 7- and 25-bp insertions, namely, Vrn-A3a-h2, Vrn-A3a-h3, Vrn-A3a-h4, Vrn-A3a-h5, Vrn-A3a-h6, and Vrn-A3c-h2 from wild emmer wheat, while we identified two VRN-A3 alleles with these insertions, Vrn-A3a-h2 and Vrn-A3c-h1 from cultivated tetraploid and hexaploid wheat species in addition to Vrn-A3a-h1. Among VRN-A3 alleles distributed in cultivated wheat, we found that Vrn-A3a-h2 promoted early heading, whereas Vrn-A3c-h1 did not affect heading time. Our analysis showed that the distribution of early flowering alleles of VRN-A3 dominated in cultivated emmer wheat in Ethiopia and India, which actually showed an early flowering phenotype. This implied that the early flowering alleles of VRN-A3 contribute to adaptability to a low-latitude environment in cultivated emmer wheat. We could not find durum (T. turgidum L. ssp. durum) and bread wheat (T. aestivum L. ssp. aestivum) accessions with these early flowering alleles. Our findings indicated that Vrn-A3a-h1 and Vrn-A3a-h2 were useful for breeding of early flowering cultivars in durum and bread wheat varieties.

Fine mapping of a powdery mildew resistance gene MlIW39 derived from wild emmer wheat (Triticum turgidum ssp. dicoccoides).

Powdery mildew resistance gene MlIW39, originated from wild emmer wheat accession IW39, was mapped to a 460.3 kb genomic interval on wheat chromosome arm 2BS. Wheat powdery mildew, caused by Blumeria graminis f. sp. tritici (Bgt), is destructive disease and a significant threat to wheat production globally. The most effective way to control this disease is genetic resistance. However, when resistance genes become widely deployed in agriculture, their effectiveness is compromised by virulent variants that were previously minor components of the pathogen population or that arise from mutation. This necessitates continual search for new sources of resistance in both wheat and its near relatives. In this study, we produced a common wheat line 8D49 (87-1/IW39//2*87-1), which has all-stage immunity to Bgt isolate E09 and many other Chinese Bgt isolates, by transferring powdery mildew resistance from Israeli wild emmer wheat (WEW) accession IW39 to the susceptible common wheat line 87-1. Genetic analysis indicated that the powdery mildew resistance in 8D49 was controlled by a single dominant gene, temporarily designated MlIW39. Genetic linkage analyses with molecular markers showed that MlIW39 was located in a 0.7 cm genetic region between markers QB-3-16 and 7Seq546 on the short arm of chromosome 2B. Fine mapping using three large F2 populations delimited MlIW39 to a physical interval of approximately 460.3 kb region in the WEW reference genome (Zavitan v1.0) that contained six annotated protein-coding genes, four of which had gene structures similar to known disease resistance genes. This provides a foundation for map-based cloning of MlIW39. Markers 7Seq622 and 7Seq727 co-segregating with MlIW39 can be utilized for marker-assisted selection in further genetic studies and wheat breeding.

Emmer wheat as a source for trait improvement in durum wheat: a study of general and specific combining ability

Emmer wheat (Triticum. turgidum ssp. dicoccum) as an ancestor of bread and durum wheats, can be a potential resource to restore genetic diversity in modern durum wheats. In order to estimate the combining ability, the type of gene action, heritability, and other genetic parameters of agronomic traits, a full diallel cross (12 × 12) was made between eight durum cultivars and four emmer wheat accessions. The F1 hybrids (132 combinations) and their parents were evaluated for important traits that account for productivity during two cropping seasons. Considerable genetic diversity among the parents and the hybrids was evident, with most of the hybrids showing higher grain yields than their respective durum parent. High general combining ability for all of the measured traits, and higher Baker ratios for most of the traits indicated that additive gene action was involved. Based on the Griffing diallel method, the specific combining ability was significant for most of the measured traits. The Hayman analysis revealed the presence of partial dominance gene action for traits such as the number of tillers per plant (NT), grain weight per spike (GWS), harvest index (HI), days to heading (DH), and number of kernels per spike (NKS). However, plant height (PH), days to maturity (DM), peduncle length (PL), and grain yield (GY) were under the influence of the over-dominance gene action. The narrow-sense heritability for GWS, NKS, kernel diameter (KD), and HI was relatively high and these four were positively correlated with grain yield. Therefore, selection for these four traits in early generations may indirectly improve yield. The results indicate that Iranian emmer wheats are a good source of wild type alleles and valuable QTLs to improve the elite durum wheat cultivars.

Mapping and characterization of two stem rust resistance genes derived from cultivated emmer wheat accession PI 193883.

Two stem rust resistance genes identified on chromosome arms 2BL and 6AL of the cultivated emmer wheat accession PI 193883 can be used for protecting modern varieties against Ug99 strains. The wheat research community consistently strives to identify new genes that confer resistance to stem rust caused by the fungal pathogen Puccinia graminis f. sp. tritici Eriks & E. Henn (Pgt). In the current study, our objective was to identify and genetically characterize the stem rust resistance derived from the cultivated emmer accession PI 193883. A recombinant inbred line population developed from a cross between the susceptible durum wheat line Rusty and PI 193883 was genotyped and evaluated for reaction to Pgt races TTKSK, TRTTF, and TMLKC. Two QTLs conferring resistance were identified on chromosome arms 2BL (QSr.fcu-2B) and 6AL (QSr.fcu-6A). The stem rust resistance gene (Sr883-2B) underlying QSr.fcu-2B was recessive, and based on its physical location it is located proximal to the Sr9 region. QSr.fcu-6A was located in the Sr13 region, but PI 193883 is known to carry the susceptible haplotype S4 for Sr13, indicating that the gene underlying QSr.fcu-6A (Sr883-6A) is likely a new allele of Sr13 or a gene residing close to Sr13. Three IWGSC scaffold-based simple sequence repeat (SSR) and two SNP-based semi-thermal asymmetric reverse PCR (STARP) markers were developed for the Sr883-2B region, and one STARP marker was developed for Sr883-6A. Sr883-2B was epistatic to Sr883-6A for reaction to TTKSK and TRTTF, and the two genes had additive effects for TMLKC. These two genes and the markers developed in this research provide additional resources and tools for the improvement in stem rust resistance in durum and common wheat breeding programs.

The novel function of the Ph1 gene to differentiate homologs from homoeologs evolved in Triticum turgidum ssp. dicoccoides via a dramatic meiosis-specific increase in the expression of the 5B copy of the C-Ph1 gene.

The Ph1 gene is the principal regulator of homoeologous chromosome pairing control (HECP) that ensures the diploid-like meiotic chromosome pairing behavior of polyploid wheat. The HECP control was speculated to have evolved after the first event of polyploidization. With the objective to accurately understand the evolution of the HECP control, wild emmer wheat accessions previously known to differ for HECP control were characterized for the structure and expression of the candidatePh1 gene, C-Ph1. The C-TdPh1-5A and 5B gene copies of emmer wheat showed 98 and 99% DNA sequence similarity respectively with the corresponding hexaploid wheat copies. Further, the C-TdPh1-5B carried the C-Ph1-5B specific structural changes and transcribed three splice variants as observed in the hexaploid wheat. Further, single nucleotide changes differentiating accessions varying for HECP control were identified. Analyzed by quantitative expression analysis, the wild emmer accessions with HECP control showed ~ 10,000-fold higher transcript abundance of the C-TdPh1-5B copy during prophase-I compared to accessions lacking the control. Differential transcriptional regulation of C-TdPh1-5B splice variants further revealed that C-Ph1-5Balt1 variant is mainly responsible for differential accumulation of C-Ph1-5B copy in accessions with HECP control. Taken together, these results showed that the HECP control evolved via transcriptional regulation of splice variants during meiosis.

Genome Based Meta-QTL Analysis of Grain Weight in Tetraploid Wheat Identifies Rare Alleles of GRF4 Associated with Larger Grains.

The domestication and subsequent genetic improvement of wheat led to the development of large-seeded cultivated wheat species relative to their smaller-seeded wild progenitors. While increased grain weight (GW) continues to be an important goal of many wheat breeding programs, few genes underlying this trait have been identified despite an abundance of studies reporting quantitative trait loci (QTL) for GW. Here we perform a QTL analysis for GW using a population of recombinant inbred lines (RILs) derived from the cross between wild emmer wheat accession ‘Zavitan’ and durum wheat variety ‘Svevo’. Identified QTLs in this population were anchored to the recent Zavitan reference genome, along with previously published QTLs for GW in tetraploid wheat. This genome-based, meta-QTL analysis enabled the identification of a locus on chromosome 6A whose introgression from wild wheat positively affects GW. The locus was validated using an introgression line carrying the 6A GW QTL region from Zavitan in a Svevo background, resulting in >8% increase in GW compared to Svevo. Using the reference sequence for the 6A QTL region, we identified a wheat ortholog to OsGRF4, a rice gene previously associated with GW. The coding sequence of this gene (TtGRF4-A) contains four single nucleotide polymorphisms (SNPs) between Zavitan and Svevo, one of which reveals the Zavitan allele to be rare in a core collection of wild emmer and completely absent from the domesticated emmer genepool. Similarly, another wild emmer accession (G18-16) was found to carry a rare allele of TtGRF4-A that also positively affects GW and is characterized by a unique SNP absent from the entire core collection. These results exemplify the rich genetic diversity of wild wheat, posit TtGRF4-A as a candidate gene underlying the 6A GW QTL, and suggest that the natural Zavitan and G18-16 alleles of TtGRF4-A have potential to increase wheat yields in breeding programs.

Genetic Mapping of Loci for Resistance to Stem Rust in a Tetraploid Wheat Collection.

Stem rust, caused by Puccinia graminis f. sp. tritici (Pgt), is a major biotic constraint to wheat production worldwide. Disease resistant cultivars are a sustainable means for the efficient control of this disease. To identify quantitative trait loci (QTLs) conferring resistance to stem rust at the seedling stage, an association mapping panel consisting of 230 tetraploid wheat accessions were evaluated for reaction to five Pgt races under greenhouse conditions. A high level of phenotypic variation was observed in the panel in response to all of the races, allowing for genome-wide association mapping of resistance QTLs in wild, landrace, and cultivated tetraploid wheats. Twenty-two resistance QTLs were identified, which were characterized by at least two marker-trait associations. Most of the identified resistance loci were coincident with previously identified rust resistance genes/QTLs; however, six regions detected on chromosomes 1B, 5A, 5B, 6B, and 7B may be novel. Availability of the reference genome sequence of wild emmer wheat accession Zavitan facilitated the search for candidate resistance genes in the regions where QTLs were identified, and many of them were annotated as NOD (nucleotide binding oligomerization domain)-like receptor (NLR) genes or genes related to broad spectrum resistance.