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Seedlings of durum wheat [ Triticum turgidum subsp. durum (Desf.) Husn] were exposed to zinc nutrition and to ozone (O 3 ) in a factorial combination: adequate (+Zn treatment) or no Zn (−Zn) in the nutrient solution, followed by exposure to either ozone-free air (filtered air, FA) or to 150 nL L −1 ozone (O 3 ) for 4 h. Although omitting Zn from the nutrient solution failed to impose a genuine Zn deficiency, −Zn*FA durum wheat seedlings showed a typical deficiency behaviour, i.e. Zn mobilisation from root to shoot. Such inter-organ Zn redistribution, however, did not occur in −Zn*O 3 plants. Exposure to each stress singly decreased the activity and the protein amount of foliar plasma membrane H + -ATPase, but not stress combination, which even increased the H + -ATPase expression with respect to control. In the −Zn*O 3 plants, moreover, the foliar activities of the plasma membrane-bound NAD(P)H-dependent superoxide synthase and of Cu,Zn-superoxide dismutase, and the transcripts abundance of the luminal binding protein and of the protein disulphide isomerase, were also stimulated. It is proposed that, even in the absence of actual Zn starvation, the perception of deficiency conditions could trigger changes in redox homoeostasis at the plasma membrane level, helpful in compensating an O 3 -dependent oxidative damage.

Purpose Selection for root traits has become a target in (pre-)breeding programs aiming at improving crop ability to capture soil resources. However, the benefit of selected traits in heterogeneous target environments will depend on spatial and temporal interactions between root systems, the soil environment (fertility and water supply) and management (fertiliser placement). Methods We assessed growth and phosphorus acquisition of durum wheat ( Triticum durum L.) lines defined by contrasting seminal root angles (41° ‘Narrow’ vs 82° ‘Wide’), in response to different soil phosphorus placements and seasonal water dynamics. Responses were evaluated in clear pots (seedlings), rhizoboxes (late-tillering stage) and a custom lysimeter system (flowering stage). Results The Narrow genotype increased deep soil exploration (down to 50 cm) during early growth, with a tendency to more rapidly acquire phosphorus placed as a deep fertiliser band (25 cm depth). However, genotypic differences in shoot biomass or phosphorus uptake were not apparent at anthesis, in part due to phosphorus-induced changes in phenological development. Contrastingly, the wide genotype increased root growth in the topsoil (0–10 cm) when phosphorus was stratified in this layer and produced greater biomass at anthesis under these conditions. Shoot and root biomass and phosphorus uptake decreased when the topsoil dried out, with the greatest effect observed for phosphorus stratified in this layer. Conclusion The benefits of the selected root angle trait strongly depend on nutrient and water distributions and dynamics in the target environment. Ideotype breeding efforts and farmer selection of genotypes should consider the context in which genotypes will be deployed. Highlight The interaction between root system architecture and heterogeneous distributions of phosphorus and available water determine the relative performance of durum wheat genotypes with contrasting root angles.

Allopolyploidization has occurred frequently within the Triticum–Aegilops complex which provides a suitable system to investigate how allopolyploidization shapes the expression patterns of duplicated homeologs. We have conducted transcriptome‐profiling of leaves and young inflorescences in wild and domesticated tetraploid wheats, Triticum turgidum ssp. dicoccoides (BBAA) and ssp. durum (BBAA), an extracted tetraploid (BBAA), and a synthetic tetraploid (SˡSˡAA) wheat together with its diploid parents, Aegilops longissima (SˡSˡ) and Triticum urartu (AA). The two diploid species showed tissue‐specific differences in genome‐wide ortholog expression, which plays an important role in transcriptome shock‐mediated homeolog expression rewiring and hence transcriptome asymmetry in the synthetic tetraploid. Further changes of homeolog expression apparently occurred in natural tetraploid wheats, which led to novel transcriptome asymmetry between the two subgenomes. In particular, our results showed that extremely biased homeolog expression can occur rapidly upon the allotetraploidzation and this trend is further enhanced in the course of domestication and evolution of polyploid wheats. Our results suggest that allopolyploidization is accompanied by distinct phases of homeolog expression changes, with parental legacy playing major roles in the immediate rewiring of homeolog expression upon allopolyploidization, while evolution and domestication under allotetraploidy drive further homeolog‐expression changes toward re‐established subgenome expression asymmetry.

Members of the AP2/ERF transcription factor family play critical roles in plant development, biosynthesis of key metabolites, and stress response. A detailed study was performed to identify TtAP2s/ERFs in the durum wheat (Triticum turgidum ssp. durum) genome, which resulted in the identification of 271 genes distributed on chromosomes 1A-7B. By carrying 27 genes, chromosome 6A had the highest number of TtAP2s/ERFs. Furthermore, a duplication assay of TtAP2s/ERFs demonstrated that 70 duplicated gene pairs had undergone purifying selection. According to RNA-seq analysis, the highest expression levels in all tissues and in response to stimuli were associated with DRF and ERF subfamily genes. In addition, the results revealed that TtAP2/ERF genes have tissue-specific expression patterns, and most TtAP2/ERF genes were significantly induced in the root tissue. Additionally, 13 TtAP2/ERF genes (six ERFs, three DREBs, two DRFs, one AP2, and one RAV) were selected for further analysis via qRT-PCR of their potential in coping with drought and salinity stresses. The TtAP2/ERF genes belonging to the DREB subfamily were markedly induced under both drought-stress and salinity-stress conditions. Furthermore, docking simulations revealed several residues in the pocket sites of the proteins associated with the stress response, which may be useful in future site-directed mutagenesis studies to increase the stress tolerance of durum wheat. This study could provide valuable insights for further evolutionary and functional assays of this important gene family in durum wheat.

Key messageSimultaneous improvement of protein content and grain yield by index selection is possible but its efficiency largely depends on the weighting of the single traits. The genetic architecture of these indices is similar to that of the primary traits.Grain yield and protein content are of major importance in durum wheat breeding, but their negative correlation has hampered their simultaneous improvement. To account for this in wheat breeding, the grain protein deviation (GPD) and the protein yield were proposed as targets for selection. The aim of this work was to investigate the potential of different indices to simultaneously improve grain yield and protein content in durum wheat and to evaluate their genetic architecture towards genomics-assisted breeding. To this end, we investigated two different durum wheat panels comprising 159 and 189 genotypes, which were tested in multiple field locations across Europe and genotyped by a genotyping-by-sequencing approach. The phenotypic analyses revealed significant genetic variances for all traits and heritabilities of the phenotypic indices that were in a similar range as those of grain yield and protein content. The GPD showed a high and positive correlation with protein content, whereas protein yield was highly and positively correlated with grain yield. Thus, selecting for a high GPD would mainly increase the protein content whereas a selection based on protein yield would mainly improve grain yield, but a combination of both indices allows to balance this selection. The genome-wide association mapping revealed a complex genetic architecture for all traits with most QTL having small effects and being detected only in one germplasm set, thus limiting the potential of marker-assisted selection for trait improvement. By contrast, genome-wide prediction appeared promising but its performance strongly depends on the relatedness between training and prediction sets.

Summary Durum wheat (Triticum turgidum subsp. durum) is a key crop worldwide, and yet, its improvement and adaptation to emerging environmental threats is made difficult by the limited amount of allelic variation included in its elite pool. New allelic diversity may provide novel loci to international crop breeding through quantitative trait loci (QTL) mapping in unexplored material. Here, we report the extensive molecular and phenotypic characterization of hundreds of Ethiopian durum wheat landraces and several Ethiopian improved lines. We test 81 587 markers scoring 30 155 single nucleotide polymorphisms and use them to survey the diversity, structure, and genome‐specific variation in the panel. We show the uniqueness of Ethiopian germplasm using a siding collection of Mediterranean durum wheat accessions. We phenotype the Ethiopian panel for ten agronomic traits in two highly diversified Ethiopian environments for two consecutive years and use this information to conduct a genome‐wide association study. We identify several loci underpinning agronomic traits of interest, both confirming loci already reported and describing new promising genomic regions. These loci may be efficiently targeted with molecular markers already available to conduct marker‐assisted selection in Ethiopian and international wheat. We show that Ethiopian durum wheat represents an important and mostly unexplored source of durum wheat diversity. The panel analysed in this study allows the accumulation of QTL mapping experiments, providing the initial step for a quantitative, methodical exploitation of untapped diversity in producing a better wheat.

Key messageWe evaluated the potential of wheat wild relatives for the improvement in grain quality characteristics including micronutrients (Fe, Zn) and gluten and identified diploid wheats and the timopheevii lineage as the most promising resources.Domestication enabled the advancement of civilization through modification of plants according to human requirements. Continuous selection and cultivation of domesticated plants induced genetic bottlenecks. However, ancient diversity has been conserved in crop wild relatives. Wheat (Triticum aestivum L.; Triticum durum Desf.) is one of the most important staple foods and was among the first domesticated crop species. Its evolutionary diversity includes diploid, tetraploid and hexaploid species from the Triticum and Aegilops taxa and different genomes, generating an AA, BBAA/GGAA and BBAADD/GGAAAmAm genepool, respectively. Breeding and improvement in wheat altered its grain quality. In this review, we identified evolutionary patterns and the potential of wheat wild relatives for quality improvement regarding the micronutrients Iron (Fe) and Zinc (Zn), the gluten storage proteins α-gliadins and high molecular weight glutenin subunits (HMW-GS), and the secondary metabolite phenolics. Generally, the timopheevii lineage has been neglected to date regarding grain quality studies. Thus, the timopheevii lineage should be subject to grain quality research to explore the full diversity of the wheat gene pool.

Summary The Ethiopian plateau hosts thousands of durum wheat (Triticum turgidum subsp. durum) farmer varieties (FV) with high adaptability and breeding potential. To harness their unique allelic diversity, we produced a large nested association mapping (NAM) population intercrossing fifty Ethiopian FVs with an international elite durum wheat variety (Asassa). The Ethiopian NAM population (EtNAM) is composed of fifty interconnected bi‐parental families, totalling 6280 recombinant inbred lines (RILs) that represent both a powerful quantitative trait loci (QTL) mapping tool, and a large pre‐breeding panel. Here, we discuss the molecular and phenotypic diversity of the EtNAM founder lines, then we use an array featuring 13 000 single nucleotide polymorphisms (SNPs) to characterize a subset of 1200 EtNAM RILs from 12 families. Finally, we test the usefulness of the population by mapping phenology traits and plant height using a genome wide association (GWA) approach. EtNAM RILs showed high allelic variation and a genetic makeup combining genetic diversity from Ethiopian FVs with the international durum wheat allele pool. EtNAM SNP data were projected on the fully sequenced AB genome of wild emmer wheat, and were used to estimate pairwise linkage disequilibrium (LD) measures that reported an LD decay distance of 7.4 Mb on average, and balanced founder contributions across EtNAM families. GWA analyses identified 11 genomic loci individually affecting up to 3 days in flowering time and more than 1.6 cm in height. We argue that the EtNAM is a powerful tool to support the production of new durum wheat varieties targeting local and global agriculture.

Background and aims Characterising the spatial distribution of tree fine roots (diameter≤2 mm) is fundamental for a better understanding of belowground functioning when tree are grown with associated crops in agroforestry systems. Our aim was to compare fine root distributions and orientations in trees grown in an alley cropping agroforestry stand with those in a tree monoculture. Methods Fieldwork was conducted in two adjacent 17 year old hybrid walnut (Juglans regia×nigra L.) stands in southern France: the agroforestry stand was intercropped with durum wheat (Triticum turgidum L. subsp. durum) whereas the tree monoculture had a natural understorey. Root intercepts were mapped to a depth of 150 cm on trench walls in both stands, and to a depth of 400 cm in the agroforestry stand in order to characterise tree root distribution below the crop's maximum rooting depth. Soil cubes were then extracted to assess three dimensional root orientation and to establish a predictive model of root length densities (RLD) derived from root intersection densities (RID). Results In the tree monoculture, root mapping demonstrated a very high tree RID in the top 50 cm and a slight decrease in RID with increasing soil depth. However, in the agroforestry stand, RID was significantly lower at 50 cm, tree roots colonized deeper soil layers and were more vertically oriented. In the agroforestry stand, RID and RLD were greater within the tree row than in the inter-row. Conclusions Fine roots of intercropped walnut trees grew significantly deeper, indicating a strong plasticity in root distribution. This plasticity reduced direct root competition from the crop, enabling trees to access deeper water tables not available to crop roots.

Two major genes for Na + exclusion in durum wheat, Nax1 and Nax2, that were previously identified as the Na + transporters TmHKT1;4-A2 and TmHKT1;5-A, were transferred into bread wheat in order to increase its capacity to restrict the accumulation of Na + in leaves. The genes were crossed from tetraploid durum wheat (Triticum turgidum ssp. durum) into hexaploid bread wheat (Triticum aestivum) by interspecific crossing and marker-assisted selection for hexaploid plants containing one or both genes. Nax1 decreased the leaf blade Na + concentration by 50%, Nax2 decreased it by 30%, and both genes together decreased it by 60%. The signature phenotype of Nax1, the retention of Na + in leaf sheaths resulting in a high Na + sheath:blade ratio, was found in the Nax1 lines. This conferred an extra advantage under a combination of waterlogged and saline conditions. The effect of Nax2 on lowering the Na + concentration in bread wheat was surprising as this gene is very similar to the TaHKT1;5-D Na + transporter already present in bread wheat, putatively at the Kna1 locus. The results indicate that both Nax genes have the potential to improve the salt tolerance of bread wheat.