Key message A high-resolution genetic map combined with haplotype analyses identified a wheat ortholog of rice gene APO1 as the best candidate gene for a 7AL locus affecting spikelet number per spike. Abstract A better understanding of the genes controlling differences in wheat grain yield components can accelerate the improvements required to satisfy future food demands. In this study, we identified a promising candidate gene underlying a quantitative trait locus (QTL) on wheat chromosome arm 7AL regulating spikelet number per spike (SNS). We used large heterogeneous inbred families ( > 10,000 plants) from two crosses to map the 7AL QTL to an 87-kb region (674,019,191–674,106,327 bp, RefSeq v1.0) containing two complete and two partial genes. In this region, we found three major haplotypes that were designated as H1, H2 and H3. The H2 haplotype contributed the high-SNS allele in both H1 × H2 and H2 × H3 segregating populations. The ancestral H3 haplotype is frequent in wild emmer (48%) but rare (~ 1%) in cultivated wheats. By contrast, the H1 and H2 haplotypes became predominant in modern cultivated durum and common wheat, respectively. Among the four candidate genes, only TraesCS7A02G481600 showed a non-synonymous polymorphism that differentiated H2 from the other two haplotypes. This gene, designated here as WHEAT ORTHOLOG OF APO1 ( WAPO1 ), is an ortholog of the rice gene ABERRANT PANICLE ORGANIZATION 1 ( APO1 ), which affects spikelet number. Taken together, the high-resolution genetic map, the association between polymorphisms in the different mapping populations with differences in SNS, and the known role of orthologous genes in other grass species suggest that WAPO-A1 is the most likely candidate gene for the 7AL SNS QTL among the four genes identified in the candidate gene region. Electronic supplementary material The online version of this article (10.1007/s00122-019-03382-5) contains supplementary material, which is available to authorized users.

Summary Awns are stiff, hair‐like structures which grow from the lemmas of wheat (Triticum aestivum) and other grasses that contribute to photosynthesis and play a role in seed dispersal. Variation in awn length in domesticated wheat is controlled primarily by three major genes, most commonly the dominant awn suppressor Tipped1 (B1). This study identifies a transcription repressor responsible for awn inhibition at the B1 locus. Association mapping was combined with analysis in biparental populations to delimit B1 to a distal region of 5AL colocalized with QTL for number of spikelets per spike, kernel weight, kernel length, and test weight. Fine‐mapping located B1 to a region containing only two predicted genes, including C2H2 zinc finger transcriptional repressor TraesCS5A02G542800 upregulated in developing spikes of awnless individuals. Deletions encompassing this candidate gene were present in awned mutants of an awnless wheat. Sequence polymorphisms in the B1 coding region were not observed in diverse wheat germplasm whereas a nearby polymorphism was highly predictive of awn suppression. Transcriptional repression by B1 is the major determinant of awn suppression in global wheat germplasm. It is associated with increased number of spikelets per spike and decreased kernel size.

Allopolyploidy greatly expands the range of possible regulatory interactions among functionally redundant homoeologous genes. However, connection between the emerging regulatory complexity and expression and phenotypic diversity in polyploid crops remains elusive. Here, we use diverse wheat accessions to map expression quantitative trait loci (eQTL) and evaluate their effects on the population-scale variation in homoeolog expression dosage. The relative contribution of cis- and trans-eQTL to homoeolog expression variation is strongly affected by both selection and demographic events. Though trans-acting effects play major role in expression regulation, the expression dosage of homoeologs is largely influenced by cis-acting variants, which appear to be subjected to selection. The frequency and expression of homoeologous gene alleles showing strong expression dosage bias are predictive of variation in yield-related traits, and have likely been impacted by breeding for increased productivity. Our study highlights the importance of genomic variants affecting homoeolog expression dosage in shaping agronomic phenotypes and points at their potential utility for improving yield in polyploid crops.

Background Genetic variation in growth over the course of the season is a major source of grain yield variation in wheat, and for this reason variants controlling heading date and plant height are among the best-characterized in wheat genetics. While the major variants for these traits have been cloned, the importance of these variants in contributing to genetic variation for plant growth over time is not fully understood. Here we develop a biparental population segregating for major variants for both plant height and flowering time to characterize the genetic architecture of the traits and identify additional novel QTL. Results We find that additive genetic variation for both traits is almost entirely associated with major and moderate-effect QTL, including four novel heading date QTL and four novel plant height QTL. FT2 and Vrn-A3 are proposed as candidate genes underlying QTL on chromosomes 3A and 7A, while Rht8 is mapped to chromosome 2D. These mapped QTL also underlie genetic variation in a longitudinal analysis of plant growth over time. The oligogenic architecture of these traits is further demonstrated by the superior trait prediction accuracy of QTL-based prediction models compared to polygenic genomic selection models. Conclusions In a population constructed from two modern wheat cultivars adapted to the southeast U.S., almost all additive genetic variation in plant growth traits is associated with known major variants or novel moderate-effect QTL. Major transgressive segregation was observed in this population despite the similar plant height and heading date characters of the parental lines. This segregation is being driven primarily by a small number of mapped QTL, instead of by many small-effect, undetected QTL. As most breeding populations in the southeast U.S. segregate for known QTL for these traits, genetic variation in plant height and heading date in these populations likely emerges from similar combinations of major and moderate effect QTL. We can make more accurate and cost-effective prediction models by targeted genotyping of key SNPs.

Key message We discovered a natural FT-A2 allele that increases grain number per spike in both pasta and bread wheat with limited effect on heading time. Abstract Increases in wheat grain yield are necessary to meet future global food demands. A previous study showed that loss-of-function mutations in FLOWERING LOCUS T2 (FT2) increase spikelet number per spike (SNS), an important grain yield component. However, these mutations were also associated with reduced fertility, offsetting the beneficial effect of the increases in SNS on grain number. Here, we report a natural mutation resulting in an aspartic acid to alanine change at position 10 (D10A) associated with significant increases in SNS and no negative effects on fertility. Using a high-density genetic map, we delimited the SNS candidate region to a 5.2-Mb region on chromosome 3AS including 28 genes. Among them, only FT-A2 showed a non-synonymous polymorphism (D10A) present in two different populations segregating for the SNS QTL on chromosome arm 3AS. These results, together with the known effect of the ft-A2 mutations on SNS, suggest that variation in FT-A2 is the most likely cause of the observed differences in SNS. We validated the positive effects of the A10 allele on SNS, grain number, and grain yield per spike in near-isogenic tetraploid wheat lines and in an hexaploid winter wheat population. The A10 allele is present at very low frequency in durum wheat and at much higher frequency in hexaploid wheat, particularly in winter and fall-planted spring varieties. These results suggest that the FT-A2 A10 allele may be particularly useful for improving grain yield in durum wheat and fall-planted common wheat varieties.

Awns are stiff, hair-like structures that grow from the spikelets of wheat ( Triticum aestivum L.) and other grasses. In wild wheat relatives, awns play a role in seed dispersal. Awn suppression in domesticated wheat, where awns are shortened or even eliminated entirely, is variably adaptive with both awned and awnless types under widespread cultivation. Although the B1 locus on the long arm of chromosome 5A is a major determinant of awn suppression, no underlying gene or mechanism of action has been identified. Using association mapping, we identified a SNP marker located on the distal end of 5 AL (5A28417) predictive of awn status in a panel of 640 U.S. winter wheat breeding lines, indicating that B1 was the determinant of awn suppression in this germplasm. A nalysis of historical data available for the panel determined 5A28417 was also significantly associated with grain test weight. Evaluation of spike morphology and kernel traits was undertaken in a 341 RIL population developed from a cross between awned soft winter wheat cultivar LA95135 and awnless cultivar SS-MPV57. Awn suppression in the population co-segregated with 5A28417 a nd was co-located with QTL for number of spikelets per spike, kernel weight and kernel length. Fine-mapping located B1 to a region containing only two predicted genes, including a C2H2 zinc finger transcription factor 219 bp from 5A28417 that we named AWNS-A1 . Deletions encompassing both genes were present in awned mutants of the awnless cultivar Brundage. Polymorphisms in the AWNS-A1 coding region were not observed in diverse wheat germplasm. However, sequencing of wheat lines representing different marker haplotypes in the surrounding region identified a deletion 3 kb downstream of AWNS-A1. A marker for this deletion was highly predictive of awn suppression in a collection of diverse wheat accessions, and indicates that AWNS-A1 is likely the major determinant of awn suppression in global wheat germplasm. AWNS-A1 is more highly expressed in developing spikes of awnless individuals, suggesting a mechanism for awn suppression. M. 2010. Source-sink relationship between photosynthetic organs and grain yield attributes during grain filling stage in spring wheat (Triticum aestivum). Int. J. Agric. Biol 12 (4): 509-515. . 2016. Detection of QTLs for traits associated with pre-harvest sprouting resistance in bread wheat (Triticum aestivum L.). Breeding science 66 (2): .260-270. . 2009 . Registration of 'Willow Creek'forage wheat. Journal of plant registrations 3 (2): 185-190. Elbaum R., Zaltzman L., Burgert I. and Fratzl P. 2007. The role of wheat awns in the seed dispersal unit. Science 316 (5826): 884-886. Elshire R.J., Glaubitz J.C., Sun, Q., Poland J.A., Kawamoto K., Buckler E.S. and Mitchell S.E. 2011. A robust, simple genotyping-by-sequencing (GBS) approach for high diversity species. PloS one 6 (5): p.e19379. Evans L.T., Bingham J., Jackson P. and Sutherland, J. 1972. Effect of awns and drought on the supply of photosynthate and its distribution within wheat ears. Annals of App...

Increases in wheat grain yield are necessary to meet future global food demands. A previous study showed that loss-of-function mutations in FLOWERING LOCUS T2 ( FT2 ) increase spikelet number per spike (SNS), an important grain yield component. Unfortunately, associated reductions in fertility offset potential increases in grain number. Here, we report a natural mutation resulting in an aspartic acid to alanine change at position 10 (D10A) associated with significant increases in SNS and no negative effects on fertility. Using a high-density genetic map, we delimited the SNS candidate region to a 5.2 Mb region on chromosome 3AS including 28 genes. Among them, only FT-A2 showed a non-synonymous polymorphism (D10A) present in two different populations segregating for the SNS QTL on chromosome arm 3AS. These results, together with the known effect of the ft-A2 mutations on SNS, suggest that variation in FT-A2 is the most likely cause of the observed differences in SNS. We validated the positive effects of the A10 allele on SNS, grain number, and grain yield per spike in near-isogenic tetraploid wheat lines and in an hexaploid winter wheat population. The A10 allele is present at very low frequency in durum wheat and at much higher frequency in hexaploid wheat, particularly in winter and fall-planted spring varieties. These results suggest that the FT-A2 A10 allele may be particularly useful for improving grain yield in durum wheat and fall planted common wheat varieties.

Wheat (Triticum aestivum) yield is impacted by a diversity of developmental processes which interact with the environment during plant growth. This complex genetic architecture complicates identifying quantitative trait loci (QTL) that can be used to improve yield. Trait data collected on individual processes or components of yield have simpler genetic bases and can be used to model how QTL generate yield variation. The objectives of this experiment were to identify QTL affecting spike yield, evaluate how their effects on spike yield proceed from effects on component phenotypes, and to understand how the genetic basis of spike yield variation changes between environments. A 358 F5:6 RIL population developed from the cross of LA-95135 and SS-MPV-57 was evaluated in two replications at five locations over the 2018 and 2019 seasons. The parents were two soft red winter wheat cultivars differing in flowering, plant height, and yield component characters. Data on yield components and plant growth were used to assemble a structural equation model (SEM) to characterize the relationships between QTL, yield components and overall spike yield. The effects of major QTL on spike yield varied by environment, and their effects on total spike yield were proportionally smaller than their effects on component traits. This typically resulted from contrasting effects on component traits, where an increase in traits associated with kernel number was generally associated with a decrease in traits related to kernel size. In all, the complete set of identified QTL was sufficient to explain most of the spike yield variation observed within each environment. Still, the relative importance of individual QTL varied dramatically. Path analysis based on coefficients estimated through SEM demonstrated that these variations in effects resulted from both different effects of QTL on phenotypes and environment-by-environment differences in the effects of phenotypes on one another, providing a conceptual model for yield genotype-by-environment interactions in wheat.