Willmar L. Leiser

Multiply to conquer: Copy number variations at Ppd-B1 and Vrn-A1 facilitate global adaptation in wheat

BackgroundCopy number variation was found to be a frequent type of DNA polymorphism in the human genome often associated with diseases but its importance in crops and the effects on agronomic traits are still largely unknown.ResultsHere, we employed a large worldwide panel of 1110 winter wheat varieties to assess the frequency and the geographic distribution of copy number variants at the Photoperiod-B1 (Ppd-B1) and the Vernalization-A1 (Vrn-A1) loci as well as their effects on flowering time under field conditions. We identified a novel four copy variant of Vrn-A1 and based on the phylogenetic relationships among the lines show that the higher copy variants at both loci are likely to have arisen independently multiple times. In addition, we found that the frequency of the different copy number variants at both loci reflects the environmental conditions in the varieties’ region of origin and based on multi-location field trials show that Ppd-B1 copy number has a substantial effect on the fine-tuning of flowering time.ConclusionsIn conclusion, our results show the importance of copy number variation at Ppd-B1 and Vrn-A1 for the global adaptation of wheat making it a key factor for wheat success in a broad range of environments and in a wider context substantiate the significant role of copy number variation in crops.

Copy number variations of CBF genes at the Fr-A2 locus are essential components of winter hardiness in wheat.

Winter hardiness is important for the adaptation of wheat to the harsh winter conditions in temperate regions and is thus also an important breeding goal. Here, we employed a panel of 407 European winter wheat cultivars to dissect the genetic architecture of winter hardiness. We show that copy number variation (CNV) of CBF (C-repeat Binding Factor) genes at the Fr-A2 locus is the essential component for winter survival, with CBF-A14 CNV being the most likely causal polymorphism, accounting for 24.3% of the genotypic variance. Genome-wide association mapping identified several markers in the Fr-A2 chromosomal region, which even after accounting for the effects of CBF-A14 copy number explained approximately 15% of the genotypic variance. This suggests that additional, as yet undiscovered, polymorphisms are present at the Fr-A2 locus. Furthermore, CNV of Vrn-A1 explained an additional 3.0% of the genotypic variance. The allele frequencies of all loci associated with winter hardiness were found to show geographic patterns consistent with their role in adaptation. Collectively, our results from the candidate gene analysis, association mapping and genome-wide prediction show that winter hardiness in wheat is a quantitative trait, but with a major contribution of the Fr-A2 locus.

Genetic architecture of male fertility restoration of Triticum timopheevii cytoplasm and fine-mapping of the major restorer locus Rf3 on chromosome 1B

Key messageRestoration of fertility in the cytoplasmic male sterility-inducing Triticum timopheevii cytoplasm can be achieved with the major restorer locus Rf3 located on chromosome 1B, but is also dependent on modifier loci.AbstractHybrid breeding relies on a hybrid mechanism enabling a cost-efficient hybrid seed production. In wheat and triticale, cytoplasmic male sterility based on the T. timopheevii cytoplasm is commonly used, and the aim of this study was to dissect the genetic architecture underlying fertility restoration. Our study was based on two segregating F2 triticale populations with 313 and 188 individuals that share a common female parent and have two different lines with high fertility restoration ability as male parents. The plants were cloned to enable replicated assessments of their phenotype and fertility restoration was evaluated based on seed set or staining for pollen fertility. The traits showed high heritabilities but their distributions differed between the two populations. In one population, a quarter of the lines were sterile, conforming to a 3:1 segregation ratio. QTL mapping identified two and three QTL in these populations, with the major QTL being detected on chromosome 1B. This QTL was collinear in both populations and likely corresponds to Rf3. We found that Rf3 explained approximately 30 and 50% of the genotypic variance, has a dominant mode of inheritance, and that the female parent lacks this locus, probably due to a 1B.1R translocation. Taken together, Rf3 is a major restorer locus that enables fertility restoration of the T. timopheevii cytoplasm, but additional modifier loci are needed for full restoration of male fertility. Consequently, Rf3 holds great potential for hybrid wheat and triticale breeding, but other loci must also be considered, either through marker-assisted or phenotypic selection.

A three-component system incorporating Ppd-D1, copy number variation at Ppd-B1, and numerous small-effect quantitative trait loci facilitates adaptation of heading time in winter wheat cultivars of worldwide origin: Heading time control in wheat

The broad adaptability of heading time has contributed to the global success of wheat in a diverse array of climatic conditions. Here, we investigated the genetic architecture underlying heading time in a large panel of 1,110 winter wheat cultivars of worldwide origin. Genome-wide association mapping, in combination with the analysis of major phenology loci, revealed a three-component system that facilitates the adaptation of heading time in winter wheat. The photoperiod sensitivity locus Ppd-D1 was found to account for almost half of the genotypic variance in this panel and can advance or delay heading by many days. In addition, copy number variation at Ppd-B1 was the second most important source of variation in heading, explaining 8.3% of the genotypic variance. Results from association mapping and genomic prediction indicated that the remaining variation is attributed to numerous small-effect quantitative trait loci that facilitate fine-tuning of heading to the local climatic conditions. Collectively, our results underpin the importance of the two Ppd-1 loci for the adaptation of heading time in winter wheat and illustrate how the three components have been exploited for wheat breeding globally.

Identification of mega-environments in Europe and effect of allelic variation at maturity E loci on adaptation of European soybean

Soybean cultivation holds great potential for a sustainable agriculture in Europe, but adaptation remains a central issue. In this large mega-environment (MEV) study, 75 European cultivars from five early maturity groups (MGs 000-II) were evaluated for maturity-related traits at 22 locations in 10 countries across Europe. Clustering of the locations based on phenotypic similarity revealed six MEVs in latitudinal direction and suggested several more. Analysis of maturity identified several groups of cultivars with phenotypic similarity that are optimally adapted to the different growing regions in Europe. We identified several haplotypes for the allelic variants at the E1, E2, E3 and E4 genes, with each E haplotype comprising cultivars from different MGs. Cultivars with the same E haplotype can exhibit different flowering and maturity characteristics, suggesting that the genetic control of these traits is more complex and that adaptation involves additional genetic pathways, for example temperature requirement. Taken together, our study allowed the first unified assessment of soybean-growing regions in Europe and illustrates the strong effect of photoperiod on soybean adaptation and MEV classification, as well as the effects of the E maturity loci for soybean adaptation in Europe.

Overcoming Phosphorus Deficiency in West African Pearl Millet and Sorghum Production Systems: Promising Options for Crop Improvement.

West Africa (WA) is among the most food insecure regions. Rapid human population growth and stagnating crop yields greatly contribute to this fact. Poor soil fertility, especially low plant available phosphorus (P) is constraining food production in the region. P-fertilizer use in WA is among the lowest in the world due to inaccessibility and high prices, often unaffordable to resource-poor subsistence farmers. This article provides an overview of soil P-deficiency in WA and opportunities to overcome it by exploiting sorghum and pearl millet genetic diversity. The topic is examined from the perspectives of plant breeding, soil science, plant physiology, plant nutrition, and agronomy, thereby referring to recent results obtained in a joint interdisciplinary research project, and reported literature. Specific objectives are to summarize: (1) The global problem of P scarcity and how it will affect WA farmers; (2) Soil P dynamics in WA soils; (3) Plant responses to P deficiency; (4) Opportunities to breed for improved crop adaptation to P-limited conditions; (5) Challenges and trade-offs for improving sorghum and pearl millet adaptation to low-P conditions in WA; and (6) Systems approaches to address soil P-deficiency in WA. Sorghum and pearl millet in WA exhibit highly significant genetic variation for P-uptake efficiency, P-utilization efficiency, and grain yield under P-limited conditions indicating the possibility of breeding P-efficient varieties. Direct selection under P-limited conditions was more efficient than indirect selection under high-P conditions. Combining P-uptake and P-utilization efficiency is recommendable for WA to avoid further soil mining. Genomic regions responsible for P-uptake, P-utilization efficiency, and grain yield under low-P have been identified in WA sorghum and pearl millet, and marker-assisted selection could be possible once these genomic regions are validated. Developing P-efficient genotypes may not, however, be a sustainable solution in itself in the long-term without replenishing the P removed from the system in harvested produce. We therefore propose the use of integrated soil fertility management and systems-oriented management such as enhanced crop-tree-livestock integration in combination with P-use-efficiency-improved varieties. Recycling P from animal bones, human excreta and urine are also possible approaches toward a partially closed and efficient P cycle in WA.

Potential for Marker-Assisted Simultaneous Improvement of Grain and Biomass Yield in Triticale

Triticale is a promising crop for agricultural biomass production but breeding has until now mainly focused on grain yield. Here, we evaluated the potential of marker-assisted simultaneous improvement of grain yield and biomass yield. To this end, we employed a large triticale doubled haploid population with 647 individuals derived from four families that were phenotyped for grain yield and biomass yield, as well as thousand-kernel weight, tiller density, and plant height in multi-environment field trials. Employing an association mapping approach, we identified quantitative trait loci (QTL) for all the five traits. The phenotypic correlation between grain yield and biomass yield was low, and we detected only one overlapping QTL suggesting different genetic architectures underlying both traits. Our results indicate that a marker-based selection for either grain yield or biomass yield does not adversely affect the other traits. Furthermore, an improvement of the multiplicative yield traits can to some extent also be achieved by selection for QTL identified for the component traits. Taken together, our results suggest that marker-assisted breeding can assist the establishment of dual-purpose triticale cultivars with high grain and biomass yield.

Simultaneous improvement of grain yield and protein content in durum wheat by different phenotypic indices and genomic selection

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.AbstractGrain 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.

Defeating the Warrior: genetic architecture of triticale resistance against a novel aggressive yellow rust race

Key messageGenome-wide association mapping of resistance against the novel, aggressive ‘Warrior’ race of yellow rust in triticale revealed a genetic architecture with some medium-effect QTL and a quantitative component, which in combination confer high levels of resistance on both leaves and ears.AbstractYellow rust is an important destructive fungal disease in small grain cereals and the exotic ‘Warrior’ race has recently conquered Europe. The aim of this study was to investigate the genetic architecture of yellow rust resistance in hexaploid winter triticale as the basis for a successful resistance breeding. To this end, a diverse panel of 919 genotypes was evaluated for yellow rust infection on leaves and ears in multi-location field trials and genotyped by genotyping-by-sequencing as well as for known Yr resistance loci. Genome-wide association mapping identified ten quantitative trait loci (QTL) for yellow rust resistance on the leaves and seven of these also for ear resistance. The total genotypic variance explained by the QTL amounted to 44.0% for leaf and 26.0% for ear resistance. The same three medium-effect QTL were identified for both traits on chromosomes 1B, 2B, and 7B. Interestingly, plants pyramiding the resistance allele of all three medium-effect QTL were generally most resistant, but constitute less than 5% of the investigated triticale breeding material. Nevertheless, a genome-wide prediction yielded a higher predictive ability than prediction based on these three QTL. Taken together, our results show that yellow rust resistance in winter triticale is genetically complex, including both medium-effect QTL as well as a quantitative resistance component. Resistance to the novel ‘Warrior’ race of this fungal pathogen is consequently best achieved by recurrent selection in the field based on identified resistant lines and can potentially be assisted by genomic approaches.

Phenotypic and genetic analysis of spike and kernel characteristics in wheat reveals long-term genetic trends of grain yield components

Key messagePhenotypic and genetic analysis of six spike and kernel characteristics in wheat revealed geographic patterns as well as long-term trends arising from breeding progress, particularly in regard to spikelet fertility, i.e. the number of kernels per spikelet, a grain yield component that appears to underlie the increase in the number of kernels per spike.AbstractWheat is a staple crop of global relevance that faces continuous demands for improved grain yield. In this study, we evaluated a panel of 407 winter wheat cultivars for six characteristics of spike and kernel development. All traits showed a large genotypic variation and had high heritabilities. We observed geographic patterns for some traits in addition to long-term trends showing a continuous increase in the number of kernels per spike. This breeding progress is likely due to the increase in spikelet fertility, i.e. the number of kernels per spikelet. While the number of kernels per spike and spikelet fertility were significantly positively correlated, both traits showed a significant negative correlation with thousand-kernel weight. Genome-wide association mapping identified only small- and moderate-effect QTL and an effect of the phenology loci Rht-D1 and Ppd-D1 on some of the traits. The allele frequencies of some QTL matched the observed geographic patterns. The quantitative inheritance of all traits with contributions of additional small-effect QTL was substantiated by genomic prediction. Taken together, our results suggest that some of the examined traits were already the basis of grain yield progress in wheat in the past decades. A more targeted exploitation of the available variation, potentially coupled with genomic approaches, may assist wheat breeding in continuing to increase yield levels globally.