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Zymoseptoria triticiis the fungal pathogen responsible for Septoria tritici blotch on wheat. Disease outcome in this pathosystem is partly determined by isolate-specific resistance, where wheat resistance genes recognize specific fungal factors triggering an immune response. Despite the large number of known wheat resistance genes, fungal molecular determinants involved in such cultivar-specific resistance remain largely unknown. We identified the avirulence factorAvrStb9using association mapping and functional validation approaches. PathotypingAvrStb9transgenic strains onStb9cultivars, near isogenic lines and wheat mapping populations, showed thatAvrStb9interacts withStb9resistance gene, triggering an immune response.AvrStb9encodes an unusually large avirulence gene with a predicted secretion signal and a protease domain. It belongs to a S41 protease family conserved across different filamentous fungi in the Ascomycota class and may constitute a core effector.AvrStb9is also conserved among a globalZ. triticipopulation and carries multiple amino acid substitutions caused by strong positive diversifying selection. These results demonstrate the contribution of an ‘atypical’ conserved effector protein to fungal avirulence and the role of sequence diversification in the escape of host recognition, adding to our understanding of host-pathogen interactions and the evolutionary processes underlying pathogen adaptation.Author SummaryFungal avirulence (Avr) genes are involved in gene-for-gene relationships with host resistance genes.Avrgenes may at the same time target host defenses to allow infection and be recognized by a host resistance gene triggering a defense response. The fungusZymoseptoria triticicauses Septoria tritici blotch, a major disease of wheat worldwide.Z. triticipopulations rapidly adapt to selection pressures such as host resistance, leading to resistance breakdown. We report the identification of the avirulence geneAvrStb9based on genetic mapping, sequence polymorphisms and allele swapping.AvrStb9is involved in the interaction withStb9resistance gene following the gene-for-gene model, and its recognition hinders disease symptoms in hosts carrying the corresponding resistance gene. Unlike other knownZ. tritici Avreffectors,AvrStb9encodes for an unusually large Avr protein with a predicted protease S41 domain conserved among diverse ascomycete lineages. We also highlight several gene mutations likely involved in escapingStb9-mediated recognition.

Senescence is a complex trait under genetic and environmental control, in which resources are remobilized from vegetative tissue into grain. Delayed senescence, or ‘staygreen’ traits, can confer stress tolerance, with extended photosynthetic activity hypothetically sustaining grain filling. The genetics of senescence regulation are largely unknown, with senescence variation often correlated with phenological traits. Here, we confirm staygreen phenotypes of two Triticum aestivum cv. Paragon ethyl methane sulfonate mutants previously identified during a forward genetic screen and selected for their agronomic performance, similar phenology, and differential senescence phenotypes. Grain filling experiments confirmed a positive relationship between onset of senescence and grain fill duration, reporting an associated ~14% increase in final dry grain weight for one mutant (P, Two staygreen mutants were identified for which grain fill duration and grain weight were enhanced. Independent amino acid substitutions within NAC subdomain IV of NAM-1 homoeologues are proposed as causative.

Plant and inflorescence architecture determine the yield potential of crops. Breeders have harnessed natural diversity for inflorescence architecture to improve yields, and induced genetic variation could provide further gains. Wheat is a vital source of protein and calories; however, little is known about the genes that regulate the development of its inflorescence. Here, we report the identification of semidominant alleles for a class III homeodomain-leucine zipper transcription factor

The main purposes of the study were (i) to develop a mapping population, (ii) to construct its genetic map for further identification of genes associated with important agronomic traits. To the best of our knowledge this is the first segregating population and genetic map developed for Kazakh bread wheat. The work is an example of how plant breeding programs in Kazakhstan have started successfully using next generation plant breeding methods to carry out systematic plant breeding to enhance final grain harvest. The KASP technology and SNP DNA-markers have been exploited to genotype and build a genetic map. The total length of the map was 1376 cM. A total 157, which spanned A, B and D subgenomes, out of the initial 178 SNP markers used formed 26 linkage groups leaving 1 duplicated and 20 unassigned markers. As threshold distance between markers was set ≤30 cM, two linkage groups were obtained for chromosomes such as 2А, 2В, 2D, 3A, 5A, 6B and 7A. Kosambi mapping function was employed to calculate recombination units. RILs were developed through SSD method up to F4 generation. Almost 97% of identified alleles were useful in evaluating the population’s genetic diversity; the remaining 3% showed no outcome. As a result, 77 DNA markers were mapped for A, 74 for B and 27 for D genomes. The mapping population will be genotyped using a high marker density array platform such as Illumina iSelect to obtain a genetic map with a relatively high coverage. Then, the population and high-resolution genetic map will be used to identify genes influencing wheat adaptation in Kazakhstan.

Senescence is a highly quantitative trait, but in wheat the genetics underpinning senescence regulation remain relatively unknown. To select senescence variation, and ultimately identify novel genetic regulators, accurate characterisation of senescence phenotypes is essential. When investigating senescence, phenotyping efforts often focus on, or are limited to, visual assessment of the flag leaves. However, senescence is a whole plant process, involving remobilisation and translocation of resources into the developing grain. Furthermore, the temporal progression of senescence poses challenges regarding trait quantification and description, whereupon the different models and approaches applied result in varying definitions of apparently similar metrics.To gain a holistic understanding of senescence we phenotyped flag leaf and peduncle senescence progression, alongside grain maturation. Reviewing the literature, we identified techniques commonly applied in quantification of senescence variation and developed simple methods to calculate descriptive and discriminatory metrics. To capture senescence dynamism, we developed the idea of calculating thermal time to different flag leaf senescence scores, for which between year Spearman’s rank correlations of r ≥ 0.59, P < 4.7 × 10−5(TT70), identify as an accurate phenotyping method. Following our experience of senescence trait genetic mapping, we recognised the need for singular metrics capable of discriminating senescence variation, identifying Thermal Time to Flag Leaf Senescence score of 70 (TT70) and Mean Peduncle senescence (MeanPed) scores as most informative. Moreover, grain maturity assessments confirmed a previous association between our staygreen traits and grain fill extension, illustrating trait functionality.Here we review different senescence phenotyping approaches and share our experiences of phenotyping two independent RIL populations segregating for staygreen traits. Together, we direct readers towards senescence phenotyping methods we found most effective, encouraging their use when investigating and discriminating senescence variation of differing genetic bases, and to aid trait selection and weighting in breeding and research programs alike.

Senescence is a complex quantitative trait under genetic and environmental control, involving the remobilisation of resources from vegetative tissue into grain. Delayed senescence, or ‘staygreen’ traits, are associated with conferring stress tolerance, with extended photosynthetic activity hypothesised to sustain grain filling. The genetics of senescence regulation are largely unknown, with senescence variation often correlated with phenological traits. Here, we confirm staygreen phenotypes of twoTriticum aestivumcv. Paragon EMS mutants previously identified during a forward genetic screen and selected for their agronomic performance, similar phenology and differential senescence phenotypes. Through grain filling experiments, we confirm a positive relationship between onset of senescence and grain fill duration, reporting an associated ∼14 % increase in final dry grain weight for one mutant,P< 0.05. Recombinant Inbred Line (RIL) populations segregating for senescence were developed for trait mapping purposes, and phenotyped over multiple years under field conditions. Staygreen traits were mapped using exome-capture enabled bulk segregant analysis (BSA), whereupon senescence was quantified, and metrics compared to qualify senescence traits and aid RIL selection. Using BSA we mapped our two staygreen traits to two independent, dominant, loci of 4.8 and 16.7 Mb in size encompassing 56 and 142 genes. Combining single marker association analysis with variant effect prediction, we identified SNPs encoding self-validating mutations located inNAM-1homoeologues and propose these as gene candidates.SummaryApplying an exome-capture enabled Bulk Segregant Analysis, we identified novelNAM-1alleles underpinning two staygreen traits associated with enhancing grain fill duration and grain weight

International audience; Dietary fibre (DF) has multiple health benefits and wheat grains are major sources of DF for human health. However, DF is depleted in white wheat flour which is more widely consumed than wholegrain. The major DF component in white flour is the cell wall polysaccharide arabinoxylan (AX). We have identified the Chinese wheat cultivar Yumai 34 as having unusually high contents of AX in both water-soluble and insoluble forms. We have therefore used populations generated from crosses between Yumai 34 and four other wheat cultivars, three with average contents of AX (Ukrainka, Altigo and Claire) and one also having unusually high AX (Valoris), in order to map QTLs for soluble AX (determined as relative viscosity of aqueous extracts of wholemeal flours) and total AX (determined by enzyme fingerprinting of white flour). A number of QTL were mapped, but most were only detected in one or two crosses. However, all four crosses showed strong QTLs for high RV/total AX on chromosome 1B, with Yumai 34 being the increasing parent, and a KASP marker for the Yumai 34 high AX allele was validated by analysis of high AX lines derived from Yumai 34 but selected by biochemical analysis. A QTL for RV was also mapped on chromosome 6B in Yumai 34 x Valoris, with Valoris being the increasing allele, which is consistent with the observation of transgressive segregation for this population. Association studies in an independent germplasm panel identified marker trait associations for relative viscosity in these same locations while direct selection for fibre content in breeding resulted in high levels of enrichment for the Yumai 34 1B allele. The data therefore indicate that marker-assisted breeding can be used to develop wheat with high AX fibre in white flour.

Functional analysis of longitudinal phenotypic data for five and 10 time points saturates QTL detection power and genomic predictive ability for canopy height in wheat., Genetic studies increasingly rely on high-throughput phenotyping, but the resulting longitudinal data pose analytical challenges. We used canopy height data from an automated field phenotyping platform to compare several approaches to scanning for quantitative trait loci (QTLs) and performing genomic prediction in a wheat recombinant inbred line mapping population based on up to 26 sampled time points (TPs). We detected four persistent QTLs (i.e. expressed for most of the growing season), with both empirical and simulation analyses demonstrating superior statistical power of detecting such QTLs through functional mapping approaches compared with conventional individual TP analyses. In contrast, even very simple individual TP approaches (e.g. interval mapping) had superior detection power for transient QTLs (i.e. expressed during very short periods). Using spline-smoothed phenotypic data resulted in improved genomic predictive abilities (5–8% higher than individual TP prediction), while the effect of including significant QTLs in prediction models was relatively minor (

Crop productivity must increase at unprecedented rates to meet the needs of the growing worldwide population. Exploiting natural variation for the genetic improvement of crops plays a central role in increasing productivity. Although current genomic technologies can be used for high-throughput identification of genetic variation, methods for efficiently exploiting this genetic potential in a targeted, systematic manner are lacking. Here, we developed a haplotype-based approach to identify genetic diversity for crop improvement using genome assemblies from 15 bread wheat (Triticum aestivum) cultivars. We used stringent criteria to identify identical-by-state haplotypes and distinguish these from near-identical sequences (~99.95% identity). We showed that each cultivar shares ~59 % of its genome with other sequenced cultivars and we detected the presence of extended haplotype blocks containing hundreds to thousands of genes across all wheat chromosomes. We found that genic sequence alone was insufficient to fully differentiate between haplotypes, as were commonly used array-based genotyping chips due to their gene centric design. We successfully used this approach for focused discovery of novel haplotypes from a landrace collection and documented their potential for trait improvement in modern bread wheat. This study provides a framework for defining and exploiting haplotypes to increase the efficiency and precision of wheat breeding towards optimising the agronomic performance of this crucial crop., Brinton, Uauy and colleagues utilize genomic data from the 10+ Wheat Genome Project to develop a useful tool for studying and generating new wheat cultivars. This framework uses advanced exploitation of wheat haplotypes to bring newfound precision and efficiency to wheat breeding.

FLOWERING LOCUS T (FT) is a central integrator of environmental signals that regulates the timing of vegetative to reproductive transition in flowering plants. In model plants, these environmental signals have been shown to include photoperiod, vernalization, and ambient temperature pathways, and in crop species, the integration of the ambient temperature pathway remains less well understood. In hexaploid wheat, at least 5 FT-like genes have been identified, each with a copy on the A, B, and D genomes. Here, we report the characterization of FT-B1 through analysis of FT-B1 null and overexpression genotypes under different ambient temperature conditions. This analysis has identified that the FT-B1 alleles perform differently under diverse environmental conditions; most notably, the FT-B1 null produces an increase in spikelet and tiller number when grown at lower temperature conditions. Additionally, absence of FT-B1 facilitates more rapid germination under both light and dark conditions. These results provide an opportunity to understand the FT-dependent pathways that underpin key responses of wheat development to changes in ambient temperature. This is particularly important for wheat, for which development and grain productivity are sensitive to changes in temperature.

The main purposes of the study were i) to develop a first mapping population for bread wheat grown in Kazakhstan, ii) to construct its genetic map for further identification of genes associated with important agronomic traits.To the best of our knowledge this is the first segregating population and genetic map developed for Kazakh bread wheat. The work is an example of how plant breeding programs in Kazakhstan have started successfully deploying next generation plant breeding methods.The KASP (Compatative Allele Specific PCR) technology of LGC Group and SNP DNA-markers have been exploited to genotype and build a genetic map of the segregating population. The total length of the map was 1376 cM. A total 157 out of initial 178 SNP markers used formed 26 linkage groups leaving 1 duplicated and 20 unassigned markers. The threshold distance between markers was set ≤ 30 cM. Therefore, two linkage groups were obtained for chromosomes such as 2A, 2B, 2D, 3A, 5A, 6B and 7A. Despite one duplicated and 20 unassigned markers, the 157 KASP SNP markers that were mapped spanned A, B and D genomes of wheat. Kosambi Mapping function was employed to calculate recombination units between makers. RILs were developed through SSD method up to F4 generation. Almost 97% of identified alleles were useful in evaluating the population’s genetic diversity; the remaining 3% showed no outcome. As a result, 77 DNA markers were mapped for A, 74 for B and 27 for D genomes. The mapping population will be genotyped using high marker density array planform such as Illumina iSelect to obtain a genetic map with a relatively high coverage. Then, the population and high-resolution genetic map will be used to identify genes influencing wheat adaptation in Kazakhstan.

Dietary fibre (DF) has multiple health benefits, and wheat products are major sources of DF for human health. However, DF is depleted in white flour, which is most widely consumed, compared to wholegrain. The major type of DF in white wheat flour is the cell wall polysaccharide arabinoxylan (AX). Previous studies have identified the Chinese wheat cultivar Yumai 34 as having unusually high contents of AX in both water-soluble and insoluble forms. We have therefore used populations generated from crosses between Yumai 34 and four other wheat cultivars, three with average contents of AX (Ukrainka, Altigo and Claire) and one also having unusually high AX (Valoris), in order to map QTLs for soluble AX (determined as relative viscosity) of aqueous extracts of wholemeal flours) and total AX (determined by enzyme fingerprinting of white flour). A number of QTL were mapped, but most were only detected in one or two crosses. However, all four crosses showed strong QTLs for high RV/total AX on chromosome 1B, with Yumai 34 being the increasing parent, and a KASP marker for the high AX Yumai 34 allele was validated by analysis of high AX lines derived from Yumai 34 but selected by biochemical analysis. A QTL for RV was mapped on chromosome 6B in Yumai 34 × Valoris, with Valoris being the increasing allele, which was consistent with the observation of transgressive segregation for this trait. The data indicate that breeding can be used to develop wheat with high AX fibre in white flour.

Automated phenotyping technologies are capable of providing continuous and precise measurements of traits that are key to today’s crop research, breeding and agronomic practices. In additional to monitoring developmental changes, high-frequency and high-precision phenotypic analysis can enable both accurate delineation of the genotype-to-phenotype pathway and the identification of genetic variation influencing environmental adaptation and yield potential. Here, we present an automated and scalable field phenotyping platform called CropQuant, designed for easy and cost-effective deployment in different environments. To manage infield experiments and crop-climate data collection, we have also developed a web-based control system called CropMonitor to provide a unified graphical user interface (GUI) to enable realtime interactions between users and their experiments. Furthermore, we established a high-throughput trait analysis pipeline for phenotypic analyses so that lightweight machine-learning modelling can be executed on CropQuant workstations to study the dynamic interactions between genotypes (G), phenotypes (P), and environmental factors (E). We have used these technologies since 2015 and reported results generated in 2015 and 2016 field experiments, including developmental profiles of five wheat genotypes, performance-related traits analyses, and new biological insights emerged from the application of the CropQuant platform.

FLOWERING LOCUS T (FT) is a central integrator of environmental signals that regulates the timing of vegetative to reproductive transition in flowering plants. In model plants, these environmental signals have been shown to include photoperiod, vernalization, and ambient temperature pathways, and in crop species, the integration of the ambient temperature pathway remains less well understood. In hexaploid wheat, at least 5 FT‐like genes have been identified, each with a copy on the A, B, and D genomes. Here, we report the characterization of FT‐B1 through analysis of FT‐B1 null and overexpression genotypes under different ambient temperature conditions. This analysis has identified that the FT‐B1 alleles perform differently under diverse environmental conditions; most notably, the FT‐B1 null produces an increase in spikelet and tiller number when grown at lower temperature conditions. Additionally, absence of FT‐B1 facilitates more rapid germination under both light and dark conditions. These results provide an opportunity to understand the FT‐dependent pathways that underpin key responses of wheat development to changes in ambient temperature. This is particularly important for wheat, for which development and grain productivity are sensitive to changes in temperature., The physiology and development of wheat respond dramatically to increases in ambient growth temperature. To combat the adverse effects of this response, we need to understand its genetic and molecular basis. Here, we present the characterization of the FLOWERING LOCUS T (FT)‐B1 mediated temperature responses to flowering, plant architecture, germination, and gene expression. We show that FT‐B1 alleles respond differently under diverse temperatures and that the absence of FT‐B1 enhances germination and increases spikelet production at lower temperatures.

Understanding the genomic complexity of bread wheat is important for unraveling domestication processes, environmental adaptation, and for future of..., Understanding the genomic complexity of bread wheat (Triticum aestivum L.) is a cornerstone in the quest to unravel the processes of domestication and the following adaptation of domesticated wheat to a wide variety of environments across the globe. Additionally, it is of importance for future improvement of the crop, particularly in the light of climate change. Focusing on the adaptation after domestication, a nested association mapping (NAM) panel of 60 segregating biparental populations was developed, mainly involving landrace accessions from the core set of the Watkins hexaploid wheat collection optimized for genetic diversity. A modern spring elite variety, “Paragon,” was used as common reference parent. Genetic maps were constructed following identical rules to make them comparable. In total, 1611 linkage groups were identified, based on recombination from an estimated 126,300 crossover events over the whole NAM panel. A consensus map, named landrace consensus map (LRC), was constructed and contained 2498 genetic loci. These newly developed genetics tools were used to investigate the rules underlying genome fluidity or rigidity, e.g., by comparing marker distances and marker orders. In general, marker order was highly correlated, which provides support for strong synteny between bread wheat accessions. However, many exceptional cases of incongruent linkage groups and increased marker distances were also found. Segregation distortion was detected for many markers, sometimes as hot spots present in different populations. Furthermore, evidence for translocations in at least 36 of the maps was found. These translocations fell, in general, into many different translocation classes, but a few translocation classes were found in several accessions, the most frequent one being the well-known T5B:7B translocation. Loci involved in recombination rate, which is an interesting trait for plant breeding, were identified by QTL analyses using the crossover counts as a trait. In total, 114 significant QTL were detected, nearly half of them with increasing effect from the nonreference parents.

Key message A high level of genetic diversity was found in the A. E. Watkins bread wheat landrace collection. Genotypic information was used to determine the population structure and to develop germplasm resources. Abstract In the 1930s A. E. Watkins acquired landrace cultivars of bread wheat (Triticum aestivum L.) from official channels of the board of Trade in London, many of which originated from local markets in 32 countries. The geographic distribution of the 826 landrace cultivars of the current collection, here called the Watkins collection, covers many Asian and European countries and some from Africa. The cultivars were genotyped with 41 microsatellite markers in order to investigate the genetic diversity and population structure of the collection. A high level of genetic diversity was found, higher than in a collection of modern European winter bread wheat varieties from 1945 to 2000. Furthermore, although weak, the population structure of the Watkins collection reveals nine ancestral geographical groupings. An exchange of genetic material between ancestral groups before commercial wheat-breeding started would be a possible explanation for this. The increased knowledge regarding the diversity of the Watkins collection was used to develop resources for wheat research and breeding, one of them a core set, which captures the majority of the genetic diversity detected. The understanding of genetic diversity and population structure together with the availability of breeding resources should help to accelerate the detection of new alleles in the Watkins collection. Electronic supplementary material The online version of this article (doi:10.1007/s00122-014-2344-5) contains supplementary material, which is available to authorized users.

Vernalization, photoperiod and the relatively poorly defined earliness per se (eps) genes regulate flowering in plants. We report here the validation of a major eps quantitative trait locus (QTL) located on wheat 1DL using near isogenic lines (NILs). We used four independent pairs of NILs derived from a cross between Spark and Rialto winter wheat varieties, grown in both the field and controlled environments. NILs carrying the Spark allele, defined by QTL flanking markers Xgdm111 and Xbarc62, consistently flowered 3–5 days earlier when fully vernalized relative to those with the Rialto. The effect was independent of photoperiod under field conditions, short days (10-h light), long days (16-h light) and very long days (20-h light). These results validate our original QTL identified using doubled haploid (DH) populations. This QTL represents variation maintained in elite north-western European winter wheat germplasm. The two DH lines used to develop the NILs, SR9 and SR23 enabled us to define the location of the 1DL QTL downstream of marker Xgdm111. SR9 has the Spark 1DL arm while SR23 has a recombinant 1DL arm with the Spark allele from Xgdm111 to the distal end. Our work suggests that marker assisted selection of eps effects is feasible and useful even before the genes are cloned. This means eps genes can be defined and positionally cloned in the same way as the photoperiod and vernalization genes have been. This validation study is a first step towards fine mapping and eventually cloning the gene directly in hexaploid wheat.

Highlights • The Rht8 introgression was associated with robust height reduction across different nitrogen and water treatments. • The Rht8 NIL had increased root-lodging resistance at agronomically-relevant nitrogen levels. • Decreased spike length associated with Rht8 resulted in a semi-compact spike. • Yield penalty associated with Rht8 was driven by reduced grain number and spike number. • The yield penalty was overcome at low nitrogen and in irrigated conditions., Reduced height 8 (Rht8) is the main alternative to the GA-insensitive Rht alleles in hot and dry environments where it reduces plant height without yield penalty. The potential of Rht8 in northern-European wheat breeding remains unclear, since the close linkage with the photoperiod-insensitive allele Ppd-D1a is unfavourable in the relatively cool summers. In the present study, two near-isogenic lines (NILs) contrasting for the Rht8/tall allele from Mara in a UK-adapted and photoperiod-sensitive wheat variety were evaluated in trials with varying nitrogen fertiliser (N) treatments and water regimes across sites in the UK and Spain. The Rht8 introgression was associated with a robust height reduction of 11% regardless of N treatment and water regime and the Rht8 NIL was more resistant to root-lodging at agronomically-relevant N levels than the tall NIL. In the UK with reduced solar radiation over the growing season than the site in Spain, the Rht8 NIL showed a 10% yield penalty at standard agronomic N levels due to concomitant reduction in grain number and spike number whereas grain weight and harvest index were not significantly different to the tall NIL. The yield penalty associated with the Rht8 introgression was overcome at low N and in irrigated conditions in the UK, and in the high-temperature site in Spain. Decreased spike length and constant spikelet number in the Rht8 NIL resulted in spike compaction of 15%, independent of N and water regime. The genetic interval of Rht8 overlaps with the compactum gene on 2DS, raising the possibility of the same causative gene. Further genetic dissection of these loci is required.

In bread wheat (Triticum aestivum L.), crop height is an important determinant of agronomic performance. The aim of this study was to identify genes controlling variation in crop height segregating in elite European winter wheat germplasm. Four doubled haploid populations derived from the crosses Avalon × Cadenza, Savannah × Rialto, Spark × Rialto and Charger × Badger were selected, representing wide diversity in European winter wheat breeding programmes. Genetic maps based on simple sequence repeat (SSR) and Diversity Arrays Technology (DArT) markers were constructed for each population. One hundred and four significant quantitative trait loci (QTL) were identified in the four populations. A meta-analysis was conducted and the effects condensed into sixteen meta-QTL on chromosomes 1A, 1B, 1D, 2A (two meta-QTL), 2B, 2D, 3A, 3B, 4B, 4D, 5A, 5B, 6A, 6B and 6D. These include QTL with additive effects equivalent to Rht-D1 and a potentially new allele of Rht8. The description of these effects offers new opportunities for the manipulation of crop height, biomass and yield in wheat breeding programmes.

Grain morphology in wheat (Triticum aestivum) has been selected and manipulated even in very early agrarian societies and remains a major breeding target. We undertook a large-scale quantitative analysis to determine the genetic basis of the phenotypic diversity in wheat grain morphology. A high-throughput method was used to capture grain size and shape variation in multiple mapping populations, elite varieties, and a broad collection of ancestral wheat species. This analysis reveals that grain size and shape are largely independent traits in both primitive wheat and in modern varieties. This phenotypic structure was retained across the mapping populations studied, suggesting that these traits are under the control of a limited number of discrete genetic components. We identified the underlying genes as quantitative trait loci that are distinct for grain size and shape and are largely shared between the different mapping populations. Moreover, our results show a significant reduction of phenotypic variation in grain shape in the modern germplasm pool compared with the ancestral wheat species, probably as a result of a relatively recent bottleneck. Therefore, this study provides the genetic underpinnings of an emerging phenotypic model where wheat domestication has transformed a long thin primitive grain to a wider and shorter modern grain.

The Lr34/Yr18 adult plant resistance gene contributes signifi cantly to durable leaf rust (caused by Puccinia triticina Eriks.) resistance. Simple and robust molecular markers that enable early detection of Lr34/Yr18 are a major advancement in wheat (Triticum aestivum L.) breeding. An insertion/deletion size variant located at the csLV34 locus on chromosome 7D within an intron sequence of a sulfate transporter-like gene tightly linked to the Lr34/Yr18 dual rust resistance gene was used to examine a global collection of wheat cultivars, landraces, and D genome–containing diploid and polyploid species of wheat relatives. Two predominant allelic size variants, csLV34a and b, found among the wheat cultivars showed disparate variation in different wheat growing zones. A strong association was observed between the presence of Lr34/Yr18 and the csLV34b allele and wheat lines known to have Lr34/Yr18 that had the csLV34a allele were rare. All landraces with the exception of those from China were predominantly of the csLV34a type. Only one size variant, csLV34a, was detected among the diploid and polyploid D genome–containing species, indicating that csLV34b arose subsequent to hexaploid bread wheat synthesis. The lineage of the csLV34b allele associated with Lr34/Yr18 in modern wheat cultivars from North and South America, CIMMYT, Australia, and Russia was tracked back to the cultivars Mentana and Ardito developed in Italy by Nazareno Strampelli in the early 1900s. The robustness of the csLV34 marker in postulating the likely occurrence of Lr34/Yr18 across a wide range of wheat germplasm and its utility in wheat breeding was confi rmed.

The sequence-specific amplified polymorphism (S-SAP) method was used to genotype 198 Nordic bread wheat landraces and cultivars from the 19th to the 21st centuries. It was shown that theSukkula-9900-LARD retrotransposon primer was highly suitable for resolving closely related wheat materials. Cluster analysis was generally consistent with pedigree information and revealed a clear separation for growth habit but not for countries. A principal coordinates analysis (PCoA) showed a separation into different time periods (before 1910, 1910–1969 and 1970–2003). These results are consistent with the breeding history and pedigree information, indicating that little hybridization has occurred between winter and spring wheat, in contrast to frequent exchange of germplasm between the Nordic countries. Estimates of gene diversity, the PCoA results, and changes in band frequencies across time indicate that plant breeding has led to substantial genetic shifts in Nordic wheat. Diversity was reduced through selections from landraces during the early 20th century, followed by a period of relatively lower genetic diversity, and a subsequent increase and net gains in diversity from the late 1960s onwards through the use of exotic germplasm. Thus, an anticipated loss of overall genetic diversity was found to be negligible, although allele losses have occurred at specific loci.

Einkorn wheat Triticum monococcum (2n=2x=14, A m A m ) is one of the earliest domesticated crops. However, it was abandoned for cultivation before the Bronze Age and has infrequently been used in wheat breeding. Little is known about the genetic variation in adaptively important biological traits in T. monococcum. A collection of 30 accessions of diverse geographic origins were characterized for phenotypic variation in various agro-morphological traits including grain storage proteins and endosperm texture, nucleotide-binding site (NBS) domain profiles of resistance (R ) genes and resistance gene analogues (RGAs), and germination under salt and drought stresses. Forty-six SSR (single sequence repeat) markers from bread wheat (T. aestivum ,2 n=6x=42, AABBDD) A genome were used to establish trait–marker associations using linear mixed models. Multiple significant associations were identified, some of which were on chromosomal regions containing previously known genetic loci. It is concluded that T. monococcum possesses large genetic diversity in multiple traits. The findings also indicate that the efficiency of association mapping is much higher in T. monococcum than in other plant species. The use of T. monococcum as a reference species for wheat functional genomics is discussed.

SUMMARY Salt-affected soil is a major problem and has led to large areas of the earth's surface being unable to support agriculture. Wheat is not tolerant to high levels of salt; however, a number of its wild relatives, in particular Thinopyrum bessarabicum Savul. and Rayss, which grows on the Black Sea shore in the Crimea, are tolerant to high levels of NaCl. Thus the introgression of genes conferring salt tolerance from Th. bessarabicum into Triticum aestivum is one way in which salt-tolerant wheat varieties can be produced. This paper reviews the progress made in developing salt-tolerant wheat carrying Th. bessarabicum genes and salt-tolerant wheat/Th. bessarabicum amphiploids.

Salt-affected soil is a major world-wide problem with many hectares of land lost to cultivation each year .To combat this problem, the development and assessment of a novel salt tolerant cereal, tritipyrum, was carried out. This is a hybrid between wheat and Thinopyrum bessarabicum. a very salt-tolerant member of the Triticeae. A range of tritipyrums derived from crosses between tetraploid wheat, Triticum durum, and Th. bessarabicum was produced. Although meiosis in ihe tritipyrums was generally regular, chromosome pairing failure was observed in each of the genotypes. In addition, the level of fertility was relatively low, the fertility of bagged spikes ranging from 29% to 51%. One iritipyrum produced multiple seeds in some florets. Three tritipyrums selected at random performed better than their wheat parents in hydroculture experiments in each of three treatments (150, 200 and 250 mol/ m3 Nacl). The potential for the exploitation of tritipyrum in salt-affected soils is discussed.

Vernalization, photoperiod and the relatively poorly defined earliness per se (eps) genes regulate flowering in plants. We report here the validation of a major eps quantitative trait locus (QTL) located on wheat 1DL using near isogenic lines (NILs). We used four independent pairs of NILs derived from a cross between Spark and Rialto winter wheat varieties, grown in both the field and controlled environments. NILs carrying the Spark allele, defined by QTL flanking markers Xgdm111 and Xbarc62, consistently flowered 3–5 days earlier when fully vernalized relative to those with the Rialto. The effect was independent of photoperiod under field conditions, short days (10-h light), long days (16-h light) and very long days (20-h light). These results validate our original QTL identified using doubled haploid (DH) populations. This QTL represents variation maintained in elite north-western European winter wheat germplasm. The two DH lines used to develop the NILs, SR9 and SR23 enabled us to define the location of the 1DL QTL downstream of marker Xgdm111. SR9 has the Spark 1DL arm while SR23 has a recombinant 1DL arm with the Spark allele from Xgdm111 to the distal end. Our work suggests that marker assisted selection of eps effects is feasible and useful even before the genes are cloned. This means eps genes can be defined and positionally cloned in the same way as the photoperiod and vernalization genes have been. This validation study is a first step towards fine mapping and eventually cloning the gene directly in hexaploid wheat. Electronic supplementary material The online version of this article (doi:10.1007/s11032-014-0094-3) contains supplementary material, which is available to authorized users.

Grain yield (GY) and grain protein concentration (GPC) are two major traits contributing to the economic value of the wheat crop. These are, consequently, major targets in wheat breeding programs, but their simultaneous improvement is hampered by the negative correlation between GPC and GY. Identifying the genetic determinants of GPC and GY through quantitative trait loci (QTL) analysis would be one way to identify chromosomal regions, allowing improvement of GPC without reducing GY using marker-assisted selection. Therefore, QTL detection was carried out for GY and GPC using three inter-connected doubled haploid populations grown in a large multi-environment trial network. Chromosomes 2A, 2D, 3B, 7B and 7D showed co-location of QTL for GPC and GY with antagonistic effects, thus contributing to the negative GPC-GY relationship. Nonetheless, genomic regions determining GPC independently of GY across experiments were found on chromosomes 3A and 5D and could help breeders to move the GPC-GY relationship in a desirable direction.

Wheat varieties tend to be chromosomally unstable producing on average 2–3% of plants with abnormal chromosome numbers. A number of semi dwarf wheat varieties, carrying the gibberellic acid insensitive dwarfing genes Rht1 or Rht2, have been seen to produce distinct tall off types due to reduction in dosage of the chromosome carrying the dwarfing gene. The UK variety ‘Brigand’, carrying Rht2 on chromosome 4D, produced very distinct tall off types when this chromosome was reduced in dosage. The frequency of tall off types was sufficiently high to cause the variety to fail United Kingdom statutory uniformity tests. An attempt to prevent the loss of chromosome 4D was made by constructing translocation chromosomes involving the short arm of chromosome 4D, which carries Rht2, and the long arm of chromosome 4S l from Aegilops sharonensis, which carries a gene(s) conferring preferential transmission. The work in this paper describes the field evaluation of two lines carrying 4DS.4DL-4S l L translocations, and demonstrates their success in preventing spontaneously occurring monosomy of chromosome 4D in semi-dwarf wheats.

Increasing photosynthetic capacity by extending canopy longevity during grain filling using slow senescing stay-green genotypes is a possible means to improve yield in wheat. Ethyl methanesulfonate (EMS) mutated wheat lines (Triticum aestivum L. cv. Paragon) were screened for fast and slow canopy senescence to investigate the impact on yield and nitrogen partitioning. Stay-green and fast-senescing lines with similar anthesis dates were characterised in detail. Delayed senescence was only apparent at higher nitrogen supply with low nitrogen supply enhancing the rate of senescence in all lines. In the stay-green line 3 (SG3), on a whole plant basis, tiller and seed number increased whilst thousand grain weight (TGW) decreased; although a greater N uptake was observed in the main tiller, yield was not affected. In fast-senescing line 2 (FS2), yield decreased, principally as a result of decreased TGW. Analysis of N-partitioning in the main stem indicated that although the slow-senescing line had lower biomass and consequently less nitrogen in all plant parts, the proportion of biomass and nitrogen in the flag leaf was greater at anthesis compared to the other lines; this contributed to the grain N and yield of the slow-senescing line at maturity in both the main tiller and in the whole plant. A field trial confirmed senescence patterns of the two lines, and the negative impact on yield for FS2 and a positive impact for SG3 at low N only. The lack of increased yield in the slow-senescing line was likely due to decreased biomass and additionally a possible sink limitation.

The rate of organ emergence and the duration of developmental phases are key components of plant fitness to its environment. Here, we describe a hybrid genetic algorithm for the estimation of the parameters of complex non-linear simulation models that we used to estimate the varietal parameters of a well-evaluated ecophysiological model (Sirius) of wheat phenology. The aim of this study was to analyze the correlations between the varietal parameters of Sirius and to determine the minimum number of parameters that need to be estimated in order to accurately simulate the effect of the genotype and environment on wheat anthesis date. A panel of 16 bread wheat cultivars was grown under controlled conditions with different vernalization and daylength treatments and in a field nursery with different sowing dates in order to characterize their vernalization and photoperiodic requirements and earliness per se. These cultivars were also grown in the field in two consecutive growing seasons in France and in the UK along a 800 km latitude transect, where the dynamics of leaf appearance and anthesis date were determined. Variation in both final leaf number and anthesis date in response to the environment and the genotype was predicted with a mean error of 0.55 leaves and 3.94 d, respectively, after estimation of only three of the seven varietal parameters of Sirius: the phyllochron, the response of vernalization rate to temperature, and the daylength response of leaf production. Among them, the phyllochron was reasonably well estimated and showed a positive association with earliness per se (Spearman's coefficient of rank correlation = 0.59). This study showed that the number of varietal parameters in Sirius was overestimated and that considering only three varietal parameters reduced the correlations between the parameters and the root mean square error of prediction for final leaf number and anthesis date. We conclude that a phenomenogical model of wheat development can be used to estimate key phenological parameters that are difficult to determine in the field; offering the possibility to conduct large-scale quantitative genetic studies to understand better the genetic control of flowering time in cereals.

Twenty-three wheat/alien addition or substitution lines were screened using restriction fragment length polymorphisms for the presence or absence of 4/5 and 4/7 reciprocal translocations in the alien chromosomes. Such translocations have previously been identified in wheat and rye. Group 4 and group 5 Aegilops umbellulata, Triticum urartu, and Thinopyrum bessarabicum chromosomes were found to carry 4/5 translocations. Evidence for a 4/7 translocation was also found in Secale montanum. The presence of the 4/5 translocations in T. urartu indicates that the translocation predates the polyploidization of wheat. The implications of these results are discussed.Key words: wheat/alien addition and substitution lines, RFLPs, translocations, wheat/alien introgression, Triticeae, evolution.

Nitrogen (N) fertilizer represents a significant cost for the grower and may also have environmental impacts through nitrate leaching and N2O (a greenhouse gas) emissions associated with denitrification. The objectives of this study were to analyze the genetic variability in N-use efficiency (grain dry matter (DM) yield per unit N available from soil and fertilizer; NUE) in winter wheat and identify traits for improved NUE for application in breeding. Fourteen UK and French cultivars and two French advanced breeding lines were tested in a 2 year/four site network comprising different locations in France and in the UK. Detailed growth analysis was conducted at anthesis and harvest in experiments including DM and N partitioning. Senescence of either the flag leaf or the whole leaf canopy was assessed from a visual score every 3-4 days from anthesis to complete canopy senescence. The senescence score was fitted against thermal time using a five parameters monomolecular-logistic equation allowing the estimation of the timing of the onset and the rate of post-anthesis senescence. In each experiment, grain yield was reduced under low N (LN), with an average reduction of 2.2 t ha(-1) (29%). Significant N x genotype level interaction was observed for NUE. Crop N uptake at harvest on average was reduced from 227 kg N ha(-1) under high N (HN) to 109 kg N ha(-1) under LN conditions while N-utilization efficiency (grain DM yield per unit crop N uptake at harvest; NUtE) increased from 34.0 to 52.1 kg DM kg(-1) N. Overall genetic variability in NUE under LN related mainly to differences in NUtE rather than N-uptake efficiency (crop N uptake at harvest per unit N available from soil and fertilizer; NUpE). However, at one site there was also a positive correlation between NUpE and NUE at LN in both years. Moreover, across the 2 year/four site network, the N x genotype effect for NUpE partly explained the N x genotype effect for grain yield and NUE. Averaging across the 16 genotypes, the timing of onset of senescence explained 86% of the variation in NUtE amongst site-season-N treatment combinations. The linear regression of onset of senescence on NutE amongst genoytpes was not significant under HN, but at three of the four sites was significant under LN explaining 32-70% of the phenotypic variation amongst genotypes in NutE. Onset of senescence amongst genotypes was negatively correlated with the efficiency with which above-ground N at anthesis was remobilized to the grain under LN. It is concluded that delaying the onset of post-anthesis senescence may be an important trait for increasing grain yield of wheat grown under low N supply. (C) 2011 Elsevier B.V. All rights reserved.

BackgroundPrevious quantitative trait loci (QTLs) studies using the Avalon × Cadenza doubled haploid (DH) population identified eleven QTLs determining plant height, heading date and grain yield. The objectives of this study were: (i) to provide insight into the effects of these QTLs using reciprocal multiple near isogenic lines (NILs) with each pair of alleles compared in both parental backgrounds (Avalon or Cadenza), (ii) quantifying epistasis by looking at the background effects and (iii) predict favourable allelic combinations to develop superior genotypes adapted to a target environment.ResultsTo this aim, a library of 553 BC2NILs and their recurrent parents were tested over two growing seasons (2012/2013 and 2013/2014). The results obtained in the present study validated the plant height, heading date and grain yield QTLs previously identified. Epistatic interactions were detected for the 6B QTL for plant height and heading date, 3A QTL for heading date and grain yield and 2A QTL for grain yield.ConclusionThe marker assisted backcrossing strategy used provided an efficient method of resolving QTL for key agronomic traits in wheat as Mendelian factors determining possible epistatic interactions. The study shows that these QTLs are amenable to marker assisted selection, fine mapping, future positional cloning, and physiological trait dissection.