European Winter Wheat Research Articles

Mapping rust resistance in European winter wheat: many QTLs for yellow rust resistance, but only a few well characterized genes for stem rust resistance

Key messageStem rust resistance was mainly based on a few, already known resistance genes; for yellow rust resistance there was a combination of designated genes and minor QTLs.Yellow rust (YR) caused by Puccinia striiformis f. sp. tritici (Pst) and stem rust (SR) caused by Puccinia graminis f. sp. tritici (Pgt) are among the most damaging wheat diseases. Although, yellow rust has occurred regularly in Europe since the advent of the Warrior race in 2011, damaging stem rust epidemics are still unusual. We analyzed the resistance of seven segregating populations at the adult growth stage with the parents being selected for YR and SR resistances across three to six environments (location–year combinations) following inoculation with defined Pst and Pgt races. In total, 600 progenies were phenotyped and 563 were genotyped with a 25k SNP array. For SR resistance, three major resistance genes (Sr24, Sr31, Sr38/Yr17) were detected in different combinations. Additional QTLs provided much smaller effects except for a gene on chromosome 4B that explained much of the genetic variance. For YR resistance, ten loci with highly varying percentages of explained genetic variance (pG, 6–99%) were mapped. Our results imply that introgression of new SR resistances will be necessary for breeding future rust resistant cultivars, whereas YR resistance can be achieved by genomic selection of many of the detected QTLs.

Novel PHOTOPERIOD-1 gene variants associate with yield-related and root-angle traits in European bread wheat

Key messagePHOTOPERIOD-1 homoeologous gene copies play a pivotal role in regulation of flowering time in wheat. Here, we show that their influence also extends to spike and shoot architecture and even impacts root development.The sequence diversity of three homoeologous copies of the PHOTOPERIOD-1 gene in European winter wheat was analyzed by Oxford Nanopore amplicon-based multiplex sequencing and molecular markers in a panel of 194 cultivars representing breeding progress over the past 5 decades. A strong, consistent association with an average 8% increase in grain yield was observed for the PpdA1-Hap1 haplotype across multiple environments. This haplotype was found to be linked in 51% of cultivars to the 2NS/2AS translocation, originally introduced from Aegilops ventricosa, which leads to an overestimation of its effect. However, even in cultivars without the 2NS/2AS translocation, PpdA1-Hap1 was significantly associated with increased grain yield, kernel per spike and kernel per m2 under optimal growth conditions, conferring a 4% yield advantage compared to haplotype PpdA1-Hap4. In contrast to Ppd-B1 and Ppd-D1, the Ppd-A1 gene exhibits novel structural variations and a high number of SNPs, highlighting the evolutionary changes that have occurred in this region over the course of wheat breeding history. Additionally, cultivars carrying the photoperiod-insensitive Ppd-D1a allele not only exhibit earlier heading, but also deeper roots compared to those with photoperiod-sensitive alleles under German conditions. PCR and KASP assays have been developed that can be effectively employed in marker-assisted breeding programs to introduce these favorable haplotypes.

Leveraging the potential of big genomic and phenotypic data for genome-wide association mapping in wheat

Genome-wide association mapping studies (GWAS) based on Big Data are a potential approach to improve marker-assisted selection in plant breeding. The number of available phenotypic and genomic data sets in which medium-sized populations of several hundred individuals have been studied is rapidly increasing. Combining these data and using them in GWAS could increase both the power of QTL discovery and the accuracy of estimation of underlying genetic effects, but is hindered by data heterogeneity and lack of interoperability. In this study, we used genomic and phenotypic data sets, focusing on Central European winter wheat populations evaluated for heading date. We explored strategies for integrating these data and subsequently the resulting potential for GWAS. Establishing interoperability between data sets was greatly aided by some overlapping genotypes and a linear relationship between the different phenotyping protocols, resulting in high quality integrated phenotypic data. In this context, genomic prediction proved to be a suitable tool to study relevance of interactions between genotypes and experimental series, which was low in our case. Contrary to expectations, fewer associations between markers and traits were found in the larger combined data than in the individual experimental series. However, the predictive power based on the marker-trait associations of the integrated data set was higher across data sets. Therefore, the results show that the integration of medium-sized to Big Data is an approach to increase the power to detect QTL in GWAS. The results encourage further efforts to standardize and share data in the plant breeding community.

Leveraging trait and QTL covariates to improve genomic prediction of resistance to Fusarium head blight in Central European winter wheat.

Fusarium head blight (FHB) is a devastating disease of wheat, causing yield losses, reduced grain quality, and mycotoxin contamination. Breeding can mitigate the severity of FHB epidemics, especially with genomics-assisted methods. The mechanisms underlying resistance to FHB in wheat have been extensively studied, including phenological traits and genome-wide markers associated with FHB severity. Here, we aimed to improve genomic prediction for FHB resistance across breeding programs by incorporating FHB-correlated traits and FHB-associated loci as model covariates. We combined phenotypic data on FHB severity, anthesis date, and plant height with genome-wide marker data from five Central European winter wheat breeding programs for genome-wide association studies (GWAS) and genomic prediction. Within all populations, FHB was correlated with anthesis date and/or plant height, and a marker linked to the semi-dwarfing locus Rht-D1 was detected with GWAS for FHB. Including the Rht-D1 marker, anthesis date, and/or plant height as covariates in genomic prediction modeling improved prediction accuracy not only within populations but also in cross-population scenarios.

Baking Quality of Organically Grown European Winter Wheat Germplasm in the South-East of Romania

Baking quality of organic winter wheat remains a significant challenge, primarily due to the usually lower grain protein concentration realized in organic agriculture. This study examined eighty-four winter wheat genotypes from Europe under organic conditions in the south-east of Romania, during two contrasting seasons. Various quality traits, i.e., grain protein content, bread volume, dough strength, breakdown and extensibility were analyzed. All examined traits exhibited significant differences among cultivars. Molecular assays, utilizing markers for Glu-A1, Glu-D1, Glu-B3g and NAM-A1 loci revealed genetic diversity within the germplasm. The study highlighted sixteen genotypes with bread volume statistically significant different from the average trial. In the organic system the Glu-A1x2* allele had positive effects on bread volume, water absorption, dough strength and extensibility. At Glu-D1, the alleles for Dx5 + Dy10 were detected in over 85% cultivars and showed positive effects on dough development time and extensibility. Another locus with positive effect on baking quality was NAM-A1. In this study the haplotype NAM-A1c had the highest frequency (51%), followed by NAM-A1d (38%) and NAM-A1a (11%). Both NAM-A1a and NAM-A1c haplotypes positively affected bread volume, water absorption, dough strength and extensibility. The Glu-B3g allele had a positive effect on development time. However, these genes could not explain the whole variation of quality parameters, as some of the best cultivars had unfavorable alleles while other cultivars carrying positive alleles had low quality. Obviously, other factors strongly influenced the quality parameters in organic agriculture, and further research is needed to understand their nature. Our results suggest the potential for achieving improved breadmaking quality, under organic conditions, by exploring genetic possibilities. However, combining suitable breadmaking quality with high yield will remain a challenge.

High-throughput field phenotyping reveals that selection in breeding has affected the phenology and temperature response of wheat in the stem elongation phase.

Crop growth and phenology are driven by seasonal changes in environmental variables, with temperature as one important factor. However, knowledge about genotype-specific temperature response and its influence on phenology is limited. Such information is fundamental to improve crop models and adapt selection strategies. We measured the increase in height of 352 European winter wheat varieties in 4 years to quantify phenology, and fitted an asymptotic temperature response model. The model used hourly fluctuations in temperature to parameterize the base temperature (Tmin), the temperature optimum (rmax), and the steepness (lrc) of growth responses. Our results show that higher Tmin and lrc relate to an earlier start and end of stem elongation. A higher rmax relates to an increased final height. Both final height and rmax decreased for varieties originating from the continental east of Europe towards the maritime west. A genome-wide association study (GWAS) indicated a quantitative inheritance and a large degree of independence among loci. Nevertheless, genomic prediction accuracies (GBLUPs) for Tmin and lrc were low (r≤0.32) compared with other traits (r≥0.59). As well as known, major genes related to vernalization, photoperiod, or dwarfing, the GWAS indicated additional, as yet unknown loci that dominate the temperature response.

Genetic control and prospects of predictive breeding for European winter wheat’s Zeleny sedimentation values and Hagberg-Perten falling number

Key messageSedimentation values and falling number in the last decades have helped maintain high baking quality despite rigorous selection for grain yield in wheat. Allelic combinations of major loci sustained the bread-making quality while improving grain yield. Glu-D1, Pinb-D1, and non-gluten proteins are associated with sedimentation values and falling number in European wheat.Zeleny sedimentation values (ZSV) and Hagberg-Perten falling number (HFN) are among the most important parameters that help determine the baking quality classes of wheat and, thus, influence the monetary benefits for growers. We used a published data set of 372 European wheat varieties evaluated in replicated field trials in multiple environments. ZSV and HFN traits hold a wide and significant genotypic variation and high broad-sense heritability. The genetic correlations revealed positive and significant associations of ZSV and HFN with each other, grain protein content (GPC) and grain hardness; however, they were all significantly negatively correlated with grain yield. Besides, GPC appeared to be the major predictor for ZSV and HFN. Our genome-wide association analyses based on high-quality SSR, SNP, and candidate gene markers revealed a strong quantitative genetic nature of ZSV and HFN by explaining their total genotypic variance as 41.49% and 38.06%, respectively. The association of known Glutenin (Glu-1) and Puroindoline (Pin-1) with ZSV provided positive analytic proof of our studies. We report novel candidate loci associated with globulins and albumins—the non-gluten monomeric proteins in wheat. In addition, predictive breeding analyses for ZSV and HFN suggest using genomic selection in the early stages of breeding programs with an average prediction accuracy of 81 and 59%, respectively.

Sequence Characterization of Extra-Chromosomal Circular DNA Content in Multiple Blackgrass (Alopecurus myosuroides) Populations.

Alopecurus myosuroides (blackgrass) is a problematic weed of Western European winter wheat, and its success is largely due to widespread multiple-herbicide resistance. Previous analysis of F2 seed families derived from two distinct blackgrass populations exhibiting equivalent non-target site resistance (NTSR) phenotypes shows resistance is polygenic and evolves from standing genetic variation. Using a CIDER-seq pipeline, we show that herbicide-resistant (HR) and herbicide-sensitive (HS) F3 plants from these F2 seed families as well as the parent populations they were derived from carry extra-chromosomal circular DNA (eccDNA). We identify the similarities and differences in the coding structures within and between resistant and sensitive populations. Although the numbers and size of detected eccDNAs varied between the populations, comparisons between the HR and HS blackgrass populations identified shared and unique coding content, predicted genes, and functional protein domains. These include genes related to herbicide detoxification such as Cytochrome P450s, ATP-binding cassette transporters, and glutathione transferases including AmGSTF1. eccDNA content was mapped to the A. myosuroides reference genome, revealing genomic regions at the distal end of chromosome 5 and the near center of chromosomes 1 and 7 as regions with a high number of mapped eccDNA gene density. Mapping to 15 known herbicide-resistant QTL regions showed that the eccDNA coding sequences matched twelve, with four QTL matching HS coding sequences; only one region contained HR coding sequences. These findings establish that, like other pernicious weeds, blackgrass has eccDNAs that contain homologs of chromosomal genes, and these may contribute genetic heterogeneity and evolutionary innovation to rapidly adapt to abiotic stresses, including herbicide treatment.

Genome-wide association mapping for pre-harvest sprouting in European winter wheat detects novel resistance QTL, pleiotropic effects, and structural variation in multiple genomes.

Pre-harvest sprouting (PHS), germination of seeds before harvest, is a major problem in global wheat (Triticum aestivum L.) production, and leads to reduced bread-making quality in affected grain. Breeding for PHS resistance can prevent losses under adverse conditions. Selecting resistant lines in years lacking pre-harvest rain, requires challenging of plants in the field or in the laboratory or using genetic markers. Despite the availability of a wheat reference and pan-genome, linking markers, genes, allelic, and structural variation, a complete understanding of the mechanisms underlying various sources of PHS resistance is still lacking. Therefore, we challenged a population of European wheat varieties and breeding lines with PHS conditions and phenotyped them for PHS traits, grain quality, phenological and agronomic traits to conduct genome-wide association mapping. Furthermore, we compared these marker-trait associations to previously reported PHS loci and evaluated their usefulness for breeding. We found markers associated with PHS on all chromosomes, with strong evidence for novel quantitative trait locus/loci (QTL) on chromosome 1A and 5B. The QTL on chromosome 1A lacks pleiotropic effect, for the QTL on 5B we detected pleiotropic effects on phenology and grain quality. Multiple peaks on chromosome 4A co-located with the major resistance locus Phs-A1, for which two causal genes, TaPM19 and TaMKK3, have been proposed. Mapping markers and genes to the pan-genome and chromosomal alignments provide evidence for structural variation around this major PHS-resistance locus. Although PHS is controlled by many loci distributed across the wheat genome, Phs-A1 on chromosome 4A seems to be the most effective and widely deployed source of resistance, in European wheat varieties.

Genetic Basis of Resistance to Warrior (-) Yellow Rust Race at the Seedling Stage in Current Central and Northern European Winter Wheat Germplasm.

To evaluate genetic variability and seedling plant response to a dominating Warrior (-) race of yellow rust in Northern and Central European germplasm, we used a population of 229 winter wheat cultivars and breeding lines for a genome-wide association study (GWAS). A wide variation in yellow rust disease severity (based on infection types 1-9) was observed in this panel. Four breeding lines, TS049 (from Austria), TS111, TS185, and TS229 (from Germany), and one cultivar, TS158 (KWS Talent), from Germany were found to be resistant to Warrior (-) FS 53/20 and Warrior (-) G 23/19. The GWAS identified five significant SNPs associated with yellow rust on chromosomes 1B, 2A, 5B, and 7A for Warrior (-) FS 53/20, while one SNP on chromosome 5B was associated with disease for Warrior (-) G 23/19. For Warrior (-) FS 53/20, we discovered a new QTL for yellow rust resistance associated with the marker Kukri_c5357_323 on chromosome 1B. The resistant alleles G and T at the marker loci Kukri_c5357_323 on chromosome 1B and Excalibur_c17489_804 on chromosome 5B showed the largest effects (1.21 and 0.81, respectively) on the severity of Warrior (-) FS 53/20 and Warrior (-) G 23/19. Our results provide the basis for knowledge-based resistance breeding in the face of the enormous impact of the Warrior (-) race on wheat production in Europe.

Wheat EARLY FLOWERING 3 affects heading date without disrupting circadian oscillations.

Plant breeders have indirectly selected for variation at circadian-associated loci in many of the world's major crops, when breeding to increase yield and improve crop performance. Using an eight-parent Multiparent Advanced Generation Inter-Cross (MAGIC) population, we investigated how variation in circadian clock-associated genes contributes to the regulation of heading date in UK and European winter wheat (Triticum aestivum) varieties. We identified homoeologues of EARLY FLOWERING 3 (ELF3) as candidates for the Earliness per se (Eps) D1 and B1 loci under field conditions. We then confirmed a single-nucleotide polymorphism within the coding region of TaELF3-B1 as a candidate polymorphism underlying the Eps-B1 locus. We found that a reported deletion at the Eps-D1 locus encompassing TaELF3-D1 is, instead, an allele that lies within an introgression region containing an inversion relative to the Chinese Spring D genome. Using Triticum turgidum cv. Kronos carrying loss-of-function alleles of TtELF3, we showed that ELF3 regulates heading, with loss of a single ELF3 homoeologue sufficient to alter heading date. These studies demonstrated that ELF3 forms part of the circadian oscillator; however, the loss of all homoeologues was required to affect circadian rhythms. Similarly, loss of functional LUX ARRHYTHMO (LUX) in T. aestivum, an orthologue of a protein partner of Arabidopsis (Arabidopsis thaliana) ELF3, severely disrupted circadian rhythms. ELF3 and LUX transcripts are not co-expressed at dusk, suggesting that the structure of the wheat circadian oscillator might differ from that of Arabidopsis. Our demonstration that alterations to ELF3 homoeologues can affect heading date separately from effects on the circadian oscillator suggests a role for ELF3 in cereal photoperiodic responses that could be selected for without pleiotropic deleterious alterations to circadian rhythms.

Genome-wide association study and genomic prediction of resistance to stripe rust in current Central and Northern European winter wheat germplasm

Key messageWe found two loci on chromosomes 2BS and 6AL that significantly contribute to stripe rust resistance in current European winter wheat germplasm.Stripe or yellow rust, caused by the fungus Puccinia striiformis Westend f. sp. tritici, is one of the most destructive wheat diseases. Sustainable management of wheat stripe rust can be achieved through the deployment of rust resistant cultivars. To detect effective resistance loci for use in breeding programs, an association mapping panel of 230 winter wheat cultivars and breeding lines from Northern and Central Europe was employed. Genotyping with the Illumina® iSelect® 25 K Infinium® single nucleotide polymorphism (SNP) genotyping array yielded 8812 polymorphic markers. Structure analysis revealed two subpopulations with 92 Austrian breeding lines and cultivars, which were separated from the other 138 genotypes from Germany, Norway, Sweden, Denmark, Poland, and Switzerland. Genome-wide association study for adult plant stripe rust resistance identified 12 SNP markers on six wheat chromosomes which showed consistent effects over several testing environments. Among these, two marker loci on chromosomes 2BS (RAC875_c1226_652) and 6AL (Tdurum_contig29607_413) were highly predictive in three independent validation populations of 1065, 1001, and 175 breeding lines. Lines with the resistant haplotype at both loci were nearly free of stipe rust symptoms. By using mixed linear models with those markers as fixed effects, we could increase predictive ability in the three populations by 0.13–0.46 compared to a standard genomic best linear unbiased prediction approach. The obtained results facilitate an efficient selection for stripe rust resistance against the current pathogen population in the Northern and Central European winter wheat gene pool.

Long-Amplicon Single-Molecule Sequencing Reveals Novel, Trait-Associated Variants of VERNALIZATION1 Homoeologs in Hexaploid Wheat.

The gene VERNALIZATION1 (VRN1) is a key controller of vernalization requirement in wheat. The genome of hexaploid wheat (Triticum aestivum) harbors three homoeologous VRN1 loci on chromosomes 5A, 5B, and 5D. Structural sequence variants including small and large deletions and insertions and single nucleotide polymorphisms (SNPs) in the three homoeologous VRN1 genes not only play an important role in the control of vernalization requirement, but also have been reported to be associated with other yield related traits of wheat. Here we used single-molecule sequencing of barcoded long-amplicons to assay the full-length sequences (∼13 kbp plus 700 bp from the promoter sequence) of the three homoeologous VRN1 genes in a panel of 192 predominantly European winter wheat cultivars. Long read sequences revealed previously undetected duplications, insertions and single-nucleotide polymorphisms in the three homoeologous VRN1 genes. All the polymorphisms were confirmed by Sanger sequencing. Sequence analysis showed the predominance of the winter alleles vrn-A1, vrn-B1, and vrn-D1 across the investigated cultivars. Associations of SNPs and structural variations within the three VRN1 genes with 20 economically relevant traits including yield, nodal root-angle index and quality related traits were evaluated at the levels of alleles, haplotypes, and copy number variants. Cultivars carrying structural variants within VRN1 genes showed lower grain yield, protein yield and biomass compared to those with intact genes. Cultivars carrying a single vrn-A1 copy and a unique haplotype with a high number of SNPs were found to have elevated grain yield, kernels per spike and kernels per m2 along with lower grain sedimentation values. In addition, we detected a novel SNP polymorphism within the G-quadruplex region of the promoter of vrn-A1 that was associated with deeper roots in winter wheat. Our findings show that multiplex, single-molecule long-amplicon sequencing is a useful tool for detecting variants in target genes within large plant populations, and can be used to simultaneously assay sequence variants among target multiple gene homoeologs in polyploid crops. Numerous novel VRN1 haplotypes and alleles were identified that showed significantly associations to economically important traits. These polymorphisms were converted into PCR or KASP assays for use in marker-assisted breeding.

A catalogue of resistance gene homologs and a chromosome-scale reference sequence support resistance gene mapping in winter wheat.

SummaryA resistance gene atlas is an integral component of the breeder’s arsenal in the fight against evolving pathogens. Thanks to high‐throughput sequencing, catalogues of resistance genes can be assembled even in crop species with large and polyploid genomes. Here, we report on capture sequencing and assembly of resistance gene homologs in a diversity panel of 907 winter wheat genotypes comprising ex situ genebank accessions and current elite cultivars. In addition, we use accurate long‐read sequencing and chromosome conformation capture sequencing to construct a chromosome‐scale genome sequence assembly of cv. Attraktion, an elite variety representative of European winter wheat. We illustrate the value of our resource for breeders and geneticists by (i) comparing the resistance gene complements in plant genetic resources and elite varieties and (ii) conducting genome‐wide associations scans (GWAS) for the fungal diseases yellow rust and leaf rust using reference‐based and reference‐free GWAS approaches. The gene content under GWAS peaks was scrutinized in the assembly of cv. Attraktion.

Dwarfing gene Rht24 does not affect Fusarium head blight resistance in a large European winter wheat diversity panel

Reduced height (Rht) genes are widely used in modern wheat breeding although some confer higher susceptibility to Fusarium head blight (FHB) caused by F. graminearum and other species. Our objective was to test whether the recently identified Rht24b dwarfing allele has a neutral effect on FHB response as reported previously from a single mapping population when unrelated winter wheat cultivars were analyzed. We artificially infected a panel of 420 cultivars divided into four genotypic groups (Rht24a + Rht-D1a, Rht24b + Rht-D1a, Rht24a + Rht-D1b, Rht24b + Rht-D1b) with Fusarium culmorum in five location-year combinations. High and significant (P ≤ 0.001) genetic variance for FHB severity and plant height (PH) was found in the entire panel as well as within the four Rht groups and both traits showed high entry-mean heritabilities of 0.92 and 0.98, respectively. Rht24b had no significant effect on FHB severity whereas Rht-D1b increased FHB susceptibility by 37%. The 29 most resistant cultivars either had the tallness alleles of the above mentioned Rht-D1 gene or Rht24b alone. The Rht24b + Rht-D1b combination had no significantly higher FHB severity than Rht-D1b alone. However, Rht24b reduced average PH only by 6.8 cm, whereas Rht-D1b conferred a reduction of 13.6 cm. For breeding short, FHB-resistant germplasm the neutral Rht24 gene must be complemented by further QTL or other FHB-neutral Rht genes.

Identification of a novel stripe rust resistance gene from the European winter wheat cultivar 'Acienda': A step towards rust proofing wheat cultivation.

All stage resistance to stripe rust races prevalent in India was investigated in the European winter wheat cultivar ‘Acienda’. In order to dissect the genetic basis of the resistance, a backcross population was developed between ‘Acienda’ and the stripe rust susceptible Indian spring wheat cultivar ‘HD 2967’. Inheritance studies revealed segregation for a dominant resistant gene. High density SNP genotyping was used to map stripe rust resistance and marker regression analysis located stripe rust resistance to the distal end of wheat chromosome 1A. Interval mapping located this region between the SNP markers AX-95162217 and AX-94540853, at a LOD score of 15.83 with a phenotypic contribution of 60%. This major stripe rust resistance locus from ‘Acienda’ has been temporarily designated as Yraci. A candidate gene search in the 2.76 Mb region carrying Yraci on chromosome 1A identified 18 NBS-LRR genes based on wheat RefSeqv1.0 annotations. Our results indicate that as there is no major gene reported in the Yraci chromosome region, it is likely to be a novel stripe rust resistance locus and offers potential for deployment, using the identified markers, to confer all stage stripe rust resistance.

Exploring the legacy of Central European historical winter wheat landraces

Historical wheat landraces are rich sources of genetic diversity offering untapped reservoirs for broadening the genetic base of modern varieties. Using a 20K SNP array, we investigated the accessible genetic diversity in a Central European bread wheat landrace collection with great drought, heat stress tolerance and higher tillering capacity. We discovered distinct differences in the number of average polymorphisms between landraces and modern wheat cultivars, and identified a set of novel rare alleles present at low frequencies in the landrace collection. The detected polymorphisms were unevenly distributed along the wheat genome, and polymorphic markers co-localized with genes of great agronomic importance. The geographical distribution of the inferred Bayesian clustering revealed six genetically homogenous ancestral groups among the collection, where the Central European core bared an admixed background originating from four ancestral groups. We evaluated the effective population sizes (Ne) of the Central European collection and assessed changes in diversity over time, which revealed a dramatic ~ 97% genetic erosion between 1955 and 2015.

Fusarium head blight resistance in European winter wheat: insights from genome-wide transcriptome analysis

BackgroundFusarium head blight (FHB) is a devastating disease of wheat worldwide. Resistance to FHB is quantitatively controlled by the combined effects of many small to medium effect QTL. Flowering traits, especially the extent of extruded anthers, are strongly associated with FHB resistance.ResultsTo characterize the genetic basis of FHB resistance, we generated and analyzed phenotypic and gene expression data on the response to Fusarium graminearum (Fg) infection in 96 European winter wheat genotypes, including several lines containing introgressions from the highly resistant Asian cultivar Sumai3. The 96 lines represented a broad range in FHB resistance and were assigned to sub-groups based on their phenotypic FHB severity score. Comparative analyses were conducted to connect sub-group-specific expression profiles in response to Fg infection with FHB resistance level. Collectively, over 12,300 wheat genes were Fusarium responsive. The core set of genes induced in response to Fg was common across different resistance groups, indicating that the activation of basal defense response mechanisms was largely independent of the resistance level of the wheat line. Fg-induced genes tended to have higher expression levels in more susceptible genotypes. Compared to the more susceptible non-Sumai3 lines, the Sumai3-derivatives demonstrated higher constitutive expression of genes associated with cell wall and plant-type secondary cell wall biogenesis and higher constitutive and Fg-induced expression of genes involved in terpene metabolism. Gene expression analysis of the FHB QTL Qfhs.ifa-5A identified a constitutively expressed gene encoding a stress response NST1-like protein (TraesCS5A01G211300LC) as a candidate gene for FHB resistance. NST1 genes are key regulators of secondary cell wall biosynthesis in anther endothecium cells. Whether the stress response NST1-like gene affects anther extrusion, thereby affecting FHB resistance, needs further investigation.ConclusionInduced and preexisting cell wall components and terpene metabolites contribute to resistance and limit fungal colonization early on. In contrast, excessive gene expression directs plant defense response towards programmed cell death which favors necrotrophic growth of the Fg pathogen and could thus lead to increased fungal colonization.

Establishment and characteristics analysis of a crop–drought vulnerability curve: a case study of European winter wheat

Abstract. As an essential component of drought risk, crop–drought vulnerability refers to the degree of the adverse response of a crop to a drought event. Different drought intensities and environments can cause significant differences in crop yield losses. Therefore, quantifying drought vulnerability and then identifying its spatial characteristics will help understand vulnerability and develop risk-reduction strategies. We select the European winter wheat growing area as the study area and 0.5∘ × 0.5∘ grids as the basic assessment units. Winter wheat drought vulnerability curves are established based on the erosion–productivity impact calculator model simulation. Their loss change and loss extent characteristics are quantitatively analysed by the key points and cumulative loss rate, respectively, and are then synthetically identified via K-means clustering. The results show the following. (1) The regional yield loss rate starts to rapidly increase from 0.13 when the drought index reaches 0.18 and then converts to a relatively stable stage with the value of 0.74 when the drought index reaches 0.66. (2) In contrast to the Pod Plain, the stage transitions of the vulnerability curve lags behind in the southern mountain area, indicating a stronger tolerance to drought. (3) According to the loss characteristics during the initial, development, and attenuation stages, the vulnerability curves can be divided into five clusters, namely low-low-low, low-low-medium, medium-medium-medium, high-high-high, and low-medium-high loss types, corresponding to the spatial distribution from low latitude to high latitude and from mountain to plain. The paper provides ideas for the study of the impact of environment on vulnerability and for the possible application of vulnerability curve in the context of climate change.

Comparing Sentinel-1 and -2 Data and Indices for Agricultural Land Use Monitoring

Agricultural vegetation development and harvest date monitoring over large areas requires frequent remote sensing observations. In regions with persistent cloud coverage during the vegetation season this is only feasible with active systems, such as SAR, and is limited for optical data. To date, optical remote sensing vegetation indices are more frequently used to monitor agricultural vegetation status because they are easily processed, and the characteristics are widely known. This study evaluated the correlations of three Sentinel-2 optical indices with Sentinel-1 SAR indices over agricultural areas to gain knowledge about their relationship. We compared Sentinel-2 Normalized Difference Vegetation Index, Normalized Difference Water Index, and Plant Senescence Radiation Index with Sentinel-1 SAR VV and VH backscatter, VH/VV ratio, and Sentinel-1 Radar Vegetation Index. The study was conducted on 22 test sites covering approximately 35,000 ha of four different main European agricultural land use types, namely grassland, maize, spring barley, and winter wheat, in Lower Saxony, Germany, in 2018. We investigated the relationship between Sentinel-1 and Sentinel-2 indices for each land use type considering three phenophases (growing, green, senescence). The strength of the correlations of optical and SAR indices differed among land use type and phenophase. There was no generic correlation between optical and SAR indices in our study. However, when the data were split by land use types and phenophases, the correlations increased remarkably. Overall, the highest correlations were found for the Radar Vegetation Index and VH backscatter. Correlations for grassland were lower than for the other land use types. Adding auxiliary data to a multiple linear regression analysis revealed that, in addition to land use type and phenophase information, the lower quartile and median SAR values per field, and a spatial variable, improved the models. Other auxiliary data retrieved from a digital elevation model, Sentinel-1 orbit direction, soil type information, and other SAR values had minor impacts on the model performance. In conclusion, despite the different nature of the signal generation, there were distinct relationships between optical and SAR indices which were independent of environmental variables but could be stratified by land use type and phenophase. These relationships showed similar patterns across different test sites. However, a regional clustering of landscapes would significantly improve the relationships.