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Cultivated hexaploid oat has three different sets of nuclear genomes (A, C, D), but its evolutionary history remains elusive. A multiplexed shotgun sequencing procedure was explored to acquire maternal phylogenetic signals from chloroplast and mitochondria genomes of 25 Avena species. Phylogenetic analyses of the acquired organelle SNP data revealed a new maternal pathway towards hexaploids of oat genome evolution involving three diploid species ( A. ventricosa , A. canariensis and A. longiglumis ) and two tetraploid species ( A. insularis and A. agadiriana ). Cultivated hexaploid A. sativa acquired its maternal genome from an AC genome tetraploid close to A. insularis . Both AC genome A. insularis and AB genome A. agadiriana obtained a maternal genome from an ancient A, not C, genome diploid close to A. longiglumis . Divergence dating showed the major divergences of C genome species 19.9–21.2 million years ago (Mya), of the oldest A genome A. canariensis 13–15 Mya, and of the clade with hexaploids 8.5–9.5 Mya. These findings not only advance our knowledge on oat genome evolution, but also have implications for oat germplasm conservation and utilization in breeding.

Background Cultivated hexaploid oat (Common oat; Avena sativa ) has held a significant place within the global crop community for centuries; although its cultivation has decreased over the past century, its nutritional benefits have garnered increased interest for human consumption. We report the development of fully annotated, chromosome-scale assemblies for the extant progenitor species of the A s - and C p -subgenomes, Avena atlantica and Avena eriantha respectively. The diploid Avena species serve as important genetic resources for improving common oat’s adaptive and food quality characteristics. Results The A. atlantica and A. eriantha genome assemblies span 3.69 and 3.78 Gb with an N50 of 513 and 535 Mb, respectively. Annotation of the genomes, using sequenced transcriptomes, identified ~ 50,000 gene models in each species—including 2965 resistance gene analogs across both species. Analysis of these assemblies classified much of each genome as repetitive sequence (~ 83%), including species-specific, centromeric-specific, and telomeric-specific repeats. LTR retrotransposons make up most of the classified elements. Genome-wide syntenic comparisons with other members of the Pooideae revealed orthologous relationships, while comparisons with genetic maps from common oat clarified subgenome origins for each of the 21 hexaploid linkage groups. The utility of the diploid genomes was demonstrated by identifying putative candidate genes for flowering time (HD3A) and crown rust resistance ( Pc 91). We also investigate the phylogenetic relationships among other A- and C-genome Avena species. Conclusions The genomes we report here are the first chromosome-scale assemblies for the tribe Poeae, subtribe Aveninae. Our analyses provide important insight into the evolution and complexity of common hexaploid oat, including subgenome origin, homoeologous relationships, and major intra- and intergenomic rearrangements. They also provide the annotation framework needed to accelerate gene discovery and plant breeding.

The genus Avena (oats) contains diploid, tetraploid and hexaploid species that evolved through hybridization and polyploidization. Four genome types (named A through D) are generally recognized. We used GBS markers to construct linkage maps of A genome diploid ( Avena strigosa x A . wiestii , 2n = 14), and AB genome tetraploid ( A . barbata 2n = 28) oats. These maps greatly improve coverage from older marker systems. Seven linkage groups in the tetraploid showed much stronger homology and synteny with the A genome diploids than did the other seven, implying an allopolyploid hybrid origin of A . barbata from distinct A and B genome diploid ancestors. Inferred homeologies within A . barbata revealed that the A and B genomes are differentiated by several translocations between chromosomes within each subgenome. However, no translocation exchanges were observed between A and B genomes. Comparison to a consensus map of ACD hexaploid A . sativa (2n = 42) revealed that the A and D genomes of A . sativa show parallel rearrangements when compared to the A genomes of the diploids and tetraploids. While intergenomic translocations are well known in polyploid Avena , our results are most parsimoniously explained if translocations also occurred in the A, B and D genome diploid ancestors of polyploid Avena .

Background Hexaploid oat ( Avena sativa L.) is a commercially important cereal crop due to its soluble dietary fiber β-glucan, a hemicellulose known to prevent cardio-vascular diseases. To maximize health benefits associated with the consumption of oat-based food products, breeding efforts have aimed at increasing the β-glucan content in oat groats. However, progress has been limited. To accelerate oat breeding efforts, we leveraged existing breeding datasets (1,230 breeding lines from South Dakota State University oat breeding program grown in multiple environments between 2015 and 2022) to conduct a genome-wide association study (GWAS) to increase our understanding of the genetic control of beta-glucan content in oats and to compare strategies to implement genomic selection (GS) to increase genetic gain for β-glucan content in oat. Results Large variation for β-glucan content was observed with values ranging between 3.02 and 7.24%. An independent GWAS was performed for each breeding panel in each environment and identified 22 loci distributed over fourteen oat chromosomes significantly associated with β-glucan content. Comparison based on physical position showed that 12 out of 22 loci coincided with previously identified β-glucan QTLs, and three loci are in the vicinity of cellulose synthesis genes, Cellulose synthase-like ( Csl ). To perform a GWAS analysis across all breeding datasets, the β-glucan content of each breeding line was predicted for each of the 26 environments. The overall GWAS identified 73 loci, of which 15 coincided with loci identified for individual environments and 37 coincided with previously reported β-glucan QTLs not identified when performing the GWAS in single years. In addition, 21 novel loci were identified that were not reported in the previous studies. The proposed approach increased our ability to detect significantly associated markers. The comparison of multiple GS scenarios indicated that using a specific set of markers as a fixed effect in GS models did not increase the prediction accuracy. However, the use of multi-environment data in the training population resulted in an increase in prediction accuracy (0.61–0.72) as compared to single-year (0.28–0.48) data. The use of USDA-SoyWheOatBar-3 K genotyping array data resulted in a similar level of prediction accuracy as did genotyping-by-sequencing data. Conclusion This study identified and confirmed the location of multiple loci associated with β-glucan content. The proposed genomic strategies significantly increase both our ability to detect significant markers in GWAS and the accuracy of genomic predictions. The findings of this study can be useful to accelerate the genetic improvement of β-glucan content and other traits.

We used next-generation sequencing analysis of the 3′-part of 18S rDNA, ITS1, and a 5′-part of the 5.8S rDNA region to understand genetic variation among seven diploid A-genome Avena species. We used 4–49 accessions per species that represented the As genome (A. atlantica, A. hirtula, and wiestii), Ac genome (A. canariensis), Ad genome (A. damascena), Al genome (A. longiglumis), and Ap genome (A. prostrata). We also took into our analysis one C-genome species, A. clauda, which previously was found to be related to A-genome species. The sequences of 169 accessions revealed 156 haplotypes of which seven haplotypes were shared by two to five species. We found 16 ribotypes that consisted of a unique sequence with a characteristic pattern of single nucleotide polymorphisms and deletions. The number of ribotypes per species varied from one in A. longiglumis to four in A. wiestii. Although most ribotypes were species-specific, we found two ribotypes shared by three species (one for A. damascena, A. hirtula, and A. wiestii, and the second for A. longiglumis, A. atlantica, and A. wiestii), and a third ribotype shared between A. atlantica and A. wiestii. A characteristic feature of the A. clauda ribotype, a diploid C-genome species, is that two different families of ribotypes have been found in this species. Some of these ribotypes are characteristic of Cc-genome species, whereas others are closely related to As-genome ribotypes. This means that A. clauda can be a hybrid between As- and C-genome oats.

Background Oat ( Avena sativa L.) is an important grain and feed crop, playing a significant role in agricultural production. However, lodging remains a key factor limiting oat yield and quality. Recent advances in molecular biology have shed light on the mechanisms of oat lodging resistance, particularly highlighting the critical role of lignin and cellulose content in stem strength. Nevertheless, the molecular regulatory networks governing lignin and cellulose biosynthesis in the basal second internode remain poorly understood. Results In this study, we conducted a comprehensive analysis of phenotypic traits, lignin and cellulose content, and transcriptomic and metabolomic profiles in the basal second internode of two contrasting oat cultivars of lodging-resistant MY1 and lodging-susceptible DY2 during the filling, milk, and dough stages. Both MY1 and DY2 showed the highest levels of lignin and cellulose at milk stage, and MY1 maintained significantly higher lignin and cellulose content compared to DY2 at all developmental stages ( P  < 0.05). In addition, a total of 8,116 differentially expressed genes (DEGs) and 4,374 metabolites were identified. Our results identified six key genes involved in lignin synthesis ( 4CL3 , CAD6 , HRPA2 , CCOAOMT , CCR1 , PRX112 ) and five associated metabolites (L-phenylalanine, sinapropyl alcohol, sinapic acid, ferulic acid, coniferyl alcohol), as well as two cellulose synthesis-related genes ( CESA4 , CESA9 ) and one metabolite (uridine diphosphate glucose). These genes and metabolites were mainly highly expressed and enriched in MY1 during the milk stage, suggesting their potential role in enhancing lodging resistance. Conclusions Our integrated phenotypic traits, lignin and cellulose content, and transcriptomic and metabolomic analysis provides new insights into oat lodging resistance, and the key genes and metabolites identified in this study provide direct targets for improving lodging resistance in oat breeding.

Cytogenetic observations, phylogenetic studies and genome analysis using high-density genetic markers have suggested a tetraploid Avena species carrying the C and D genomes (formerly C and A) to be the donor of all hexaploid oats (AACCDD). However, controversy surrounds which of the three extant CCDD tetraploid species— A . insularis , A . magna and A . murphyi —is most closely related to hexaploid oats. The present work describes a comparative karyotype analysis of these three CCDD tetraploid species and two hexaploid species, A . sativa and A . byzantina . This involved the use of FISH with six simple sequence repeats (SSRs) with the motifs CT, AAC, AAG, ACG, ATC and ACT, two repeated ribosomal sequences, and C genome-specific repetitive DNA. The hybridization pattern of A . insularis with oligonucleotide (AC) 10 was also determined and compared with those previously published for A . sativa and A . byzantina . Significant differences in the 5S sites and SSR hybridization patterns of A . murphyi compared to the other CCDD species rule out its being directly involved in the origin of the hexaploids. In contrast, the repetitive and SSR hybridization patterns shown by the D genome chromosomes, and by most of the C genome chromosomes of A . magna and A . insularis , can be equated with the corresponding chromosomes of the hexaploids. Several chromosome hybridization signals seen for A . insularis , but not for A . magna , were shared with the hexaploid oats species, especially with A . byzantina . These diagnostic signals add weight to the idea that the extant A . insularis , or a direct ancestor of it, is the most closely related progenitor of hexaploid oats. The similarity of the chromosome hybridization patterns of the hexaploids and CCDD tetraploids was taken as being indicative of homology. A common chromosome nomenclature for CCDD species based on that of the hexaploid species is proposed.

Background Glutathione S-transferases (GSTs) are essential multifunctional enzymes. In the face of abiotic stresses such as drought and heavy metal exposure, plants utilize GSTs for detoxification and antioxidant defense, as these enzymes facilitate the conjugation of glutathione (GSH) with toxic compounds. Specific details of this process, however, remain unknown. Results This study identified 118 Avena sativa GST ( AsGST ) genes within the A. sativa genome and classified them into five subfamilies: Tau, Phi, Zeta, Lambda, and EF1Bγ. Phylogenetic analysis revealed that AsGSTs exhibit significant similarity to corresponding GST categories in Arabidopsis thaliana and Oryza sativa , indicating a possible common ancestor. Gene structure and conserved motif analysis demonstrated that AsGST genes within the same subfamily shares similarities in the number and positioning of exons and introns, as well as in motif composition, suggesting that these genes may perform analogous biological functions in A. sativa . The promoter regions of the identified genes are enriched with various cis-acting elements that play roles in plant growth and development, stress response, and hormone signaling. Transcriptomic analysis and real-time quantitative PCR (RT-qPCR) validation indicated that the expression of four AsGST genes ( AsGSTU12 , AsGSTU13 , AsGSTU14 , and AsGSTU15 ) was significantly up-regulated in the roots of A. sativa under both PEG-induced drought stress and CdCl 2 -induced cadmium stress. These genes likely regulate reactive oxygen species (ROS) levels by catalyzing their scavenging through glutathione (GSH) substrates, and may also participate in ABA signaling and the maintenance of osmotic homeostasis. Under cadmium stress, these genes may mitigate cadmium toxicity by enhancing the chelation and sequestration of cadmium via GSH or through its compartmentalization, as evident from the subcellular localization studies. Conclusion This study systematically described the GST gene family in A. sativa , characterized its expression patterns and potential functions in response to drought and cadmium stress, and confirmed the essential role of the AsGST gene family in mediating stress responses. The findings enhance our understanding of the mechanisms underlying stress tolerance and offer valuable genetic resources for breeding stress-tolerant A. sativa . The work also provides a theoretical framework and identifies gene targets for the development of stress-resistant A. sativa varieties.

Plants are subjected to various biotic and abiotic stresses that significantly impact their growth and productivity. To achieve balanced crop growth and yield, including for leafy vegetables, the continuous application of micronutrient is crucial. This study investigates the effects of different concentrations of copper sulphate (0, 75, 125, and 175 ppm) on the morphological and biochemical features of Spinacia oleracea and Avena sativa . Morphological parameters such as plant height, leaf area, root length, and fresh and dry weights were optimized at a concentration of 75 ppm copper sulfate. At this concentration, chlorophyll a & b levels increased significantly in Spinacia oleracea (462.9 and 249.8 𝜇𝑔/𝑔), and Avena sativa (404.7 and 437.63𝜇𝑔/𝑔). However, carotenoid content and sugar levels in Spinacia oleracea were negatively affected, while sugar content in Avena sativa increased at 125 ppm (941.6 µg/ml). Protein content increased in Spinacia oleracea (75 ppm, 180.3 µg/ml) but decreased in Avena sativa . Phenol content peaked in both plants at 75 ppm (362.2 and 244.5 µg/ml). Higher concentrations (175 ppm) of copper sulfate reduced plant productivity and health. Plants exposed to control and optimal concentrations (75 and 125 ppm) of copper sulpate exhibited the best health and growth compared to those subjected to higher concentrations. Maximum plant height, leaf area, root length, fresh and dry weights were observed at lower concentrations (75 and 125 ppm) of copper sulfate, while higher concentrations caused toxicity. Optimal copper sulfate levels enhanced chlorophyll a, chlorophyll b, total chlorophyll, protein, and phenol contents but inhibited sugar and carotenoid contents in both Spinacia oleracea and Avena sativa . Overall, increased copper sulfate treatment adversely affected the growth parameters and biochemical profiles of these plants.

Many herbicide-resistant weed species are polyploids, but far too little about the evolution of resistance mutations in polyploids is understood. Hexaploid wild oat (Avena fatua) is a global crop weed and many populations have evolved herbicide resistance. We studied plastidic acetyl-coenzyme A carboxylase (ACCase)-inhibiting herbicide resistance in hexaploid wild oat and revealed that resistant individuals can express one, two or three different plastidic ACCase gene resistance mutations (Ile-1781-Leu, Asp-2078-Gly and Cys-2088-Arg). Using ACCase resistance mutations as molecular markers, combined with genetic, molecular and biochemical approaches, we found in individual resistant wild-oat plants that (1) up to three unlinked ACCase gene loci assort independently following Mendelian laws for disomic inheritance, (2) all three of these homoeologous ACCase genes were transcribed, with each able to carry its own mutation and (3) in a hexaploid background, each individual ACCase resistance mutation confers relatively low-level herbicide resistance, in contrast to high-level resistance conferred by the same mutations in unrelated diploid weed species of the Poaceae (grass) family. Low resistance conferred by individual ACCase resistance mutations is likely due to a dilution effect by susceptible ACCase expressed by homoeologs in hexaploid wild oat and/or differential expression of homoeologous ACCase gene copies. Thus, polyploidy in hexaploid wild oat may slow resistance evolution. Evidence of coexisting non-target-site resistance mechanisms among wild-oat populations was also revealed. In all, these results demonstrate that herbicide resistance and its evolution can be more complex in hexaploid wild oat than in unrelated diploid grass weeds. Our data provide a starting point for the daunting task of understanding resistance evolution in polyploids.