High-density single nucleotide polymorphism (SNP) genotyping arrays are a powerful tool for studying genomic patterns of diversity, inferring ancestral relationships between individuals in populations and studying marker–trait associations in mapping experiments. We developed a genotyping array including about 90 000 gene-associated SNPs and used it to characterize genetic variation in allohexaploid and allotetraploid wheat populations. The array includes a significant fraction of common genome-wide distributed SNPs that are represented in populations of diverse geographical origin. We used density-based spatial clustering algorithms to enable high-throughput genotype calling in complex data sets obtained for polyploid wheat. We show that these model-free clustering algorithms provide accurate genotype calling in the presence of multiple clusters including clusters with low signal intensity resulting from significant sequence divergence at the target SNP site or gene deletions. Assays that detect low-intensity clusters can provide insight into the distribution of presence–absence variation (PAV) in wheat populations. A total of 46 977 SNPs from the wheat 90K array were genetically mapped using a combination of eight mapping populations. The developed array and cluster identification algorithms provide an opportunity to infer detailed haplotype structure in polyploid wheat and will serve as an invaluable resource for diversity studies and investigating the genetic basis of trait variation in wheat.

SummaryBread wheat (Triticum aestivum) is a globally important crop, accounting for 20% of the calories consumed by mankind. We sequenced its large and challenging 17 Gb hexaploid genome using 454 pyrosequencing and compared this with the sequences of diploid ancestral and progenitor genomes. Between 94,000-96,000 genes were identified, and two-thirds were assigned to the A, B and D genomes. High-resolution synteny maps identified many small disruptions to conserved gene order. We show the hexaploid genome is highly dynamic, with significant loss of gene family members upon polyploidization and domestication, and an abundance of gene fragments. Several classes of genes involved in energy harvesting, metabolism and growth are among expanded gene families that could be associated with crop productivity. Our analyses, coupled with the identification of extensive genetic variation, provide a new resource for accelerating gene discovery and improving this major crop.

SummaryIn wheat, a lack of genetic diversity between breeding lines has been recognized as a significant block to future yield increases. Species belonging to bread wheat's secondary and tertiary gene pools harbour a much greater level of genetic variability, and are an important source of genes to broaden its genetic base. Introgression of novel genes from progenitors and related species has been widely employed to improve the agronomic characteristics of hexaploid wheat, but this approach has been hampered by a lack of markers that can be used to track introduced chromosome segments. Here, we describe the identification of a large number of single nucleotide polymorphisms that can be used to genotype hexaploid wheat and to identify and track introgressions from a variety of sources. We have validated these markers using an ultra‐high‐density Axiom® genotyping array to characterize a range of diploid, tetraploid and hexaploid wheat accessions and wheat relatives. To facilitate the use of these, both the markers and the associated sequence and genotype information have been made available through an interactive web site.

Contents Summary286I.Introduction287II.The pollen–stigma interaction289III.Pollen–stigma interactions in species with wet stigmas290IV.Pollen–stigma interactions in species with dry stigmas295V.Is there any consensus among cell signalling pathways regulating pollen–stigma interactions?299VI.Incompatibility and the pollen–stigma interaction300VII.New directions in pollen–stigma interaction research302VIII.Future prospects310Acknowledgements312References312 Summary Siphonogamy, the delivery of nonmotile sperm to the egg via a pollen tube, was a key innovation that allowed flowering plants (angiosperms) to carry out sexual reproduction on land without the need for water. This process begins with a pollen grain (male gametophyte) alighting on and adhering to the stigma of a flower. If conditions are right, the pollen grain germinates to produce a pollen tube. The pollen tube invades the stigma and grows through the style towards the ovary, where it enters an ovule, penetrates the embryo sac (female gametophyte) and releases two sperm cells, one of which fertilizes the egg, while the other fuses with the two polar nuclei of the central cell to form the triploid endosperm. The events before fertilization (pollen–pistil interactions) comprise a series of complex cellular interactions involving a continuous exchange of signals between the haploid pollen and the diploid maternal tissue of the pistil (sporophyte). In recent years, significant progress has been made in elucidating the molecular identity of these signals and the cellular interactions that they regulate. Here we review our current understanding of the cellular and molecular interactions that mediate the earliest of these interactions between the pollen and the pistil that occur on or within the stigma – the ‘pollen–stigma interaction’.

SummaryTargeted selection and inbreeding have resulted in a lack of genetic diversity in elite hexaploid bread wheat accessions. Reduced diversity can be a limiting factor in the breeding of high yielding varieties and crucially can mean reduced resilience in the face of changing climate and resource pressures. Recent technological advances have enabled the development of molecular markers for use in the assessment and utilization of genetic diversity in hexaploid wheat. Starting with a large collection of 819 571 previously characterized wheat markers, here we describe the identification of 35 143 single nucleotide polymorphism‐based markers, which are highly suited to the genotyping of elite hexaploid wheat accessions. To assess their suitability, the markers have been validated using a commercial high‐density Affymetrix Axiom® genotyping array (the Wheat Breeders’ Array), in a high‐throughput 384 microplate configuration, to characterize a diverse global collection of wheat accessions including landraces and elite lines derived from commercial breeding communities. We demonstrate that the Wheat Breeders’ Array is also suitable for generating high‐density genetic maps of previously uncharacterized populations and for characterizing novel genetic diversity produced by mutagenesis. To facilitate the use of the array by the wheat community, the markers, the associated sequence and the genotype information have been made available through the interactive web site ‘CerealsDB’.

SummaryFood security is a global concern and substantial yield increases in cereal crops are required to feed the growing world population. Wheat is one of the three most important crops for human and livestock feed. However, the complexity of the genome coupled with a decline in genetic diversity within modern elite cultivars has hindered the application of marker-assisted selection (MAS) in breeding programmes. A crucial step in the successful application of MAS in breeding programmes is the development of cheap and easy to use molecular markers, such as single-nucleotide polymorphisms. To mine selected elite wheat germplasm for intervarietal single-nucleotide polymorphisms, we have used expressed sequence tags derived from public sequencing programmes and next-generation sequencing of normalized wheat complementary DNA libraries, in combination with a novel sequence alignment and assembly approach. Here, we describe the development and validation of a panel of 1114 single-nucleotide polymorphisms in hexaploid bread wheat using competitive allele-specific polymerase chain reaction genotyping technology. We report the genotyping results of these markers on 23 wheat varieties, selected to represent a broad cross-section of wheat germplasm including a number of elite UK varieties. Finally, we show that, using relatively simple technology, it is possible to rapidly generate a linkage map containing several hundred single-nucleotide polymorphism markers in the doubled haploid mapping population of Avalon · Cadenza.

SummaryDespite some notable successes, only a fraction of the genetic variation available in wild relatives has been utilized to produce superior wheat varieties. This is as a direct result of the lack of availability of suitable high‐throughput technologies to detect wheat/wild relative introgressions when they occur. Here, we report on the use of a new SNP array to detect wheat/wild relative introgressions in backcross progenies derived from interspecific hexaploid wheat/Ambylopyrum muticum F1 hybrids. The array enabled the detection and characterization of 218 genomewide wheat/Am. muticum introgressions, that is a significant step change in the generation and detection of introgressions compared to previous work in the field. Furthermore, the frequency of introgressions detected was sufficiently high to enable the construction of seven linkage groups of the Am. muticum genome, thus enabling the syntenic relationship between the wild relative and hexaploid wheat to be determined. The importance of the genetic variation from Am. muticum introduced into wheat for the development of superior varieties is discussed.

Interspecific hybridization and polyploidy are pivotal processes in plant evolution and speciation. The fate of new hybrid and polyploid taxa is determined by their ability to reproduce either sexually or asexually. Hybrids and allopolyploids with odd chromosome numbers are frequently sterile but some establish themselves through asexual reproduction (vegetative or apomixis). This allows novel genotypes to become established by isolating them from gene flow and leads to complex patterns of variation. The genus Sorbus is a good example of taxonomic complexity arising from the combined effects of hybridization, polyploidy and apomixis. The Avon Gorge in South-west Britain contains the greatest diversity of Sorbus in Europe, with three endemic species and four putative endemic novel hybrids among its 15 native Sorbus taxa. We used a combination of nuclear microsatellite and chloroplast DNA markers to investigate the evolutionary relationships among these Sorbus taxa within the Avon Gorge. We confirm the genetic identity of putative novel taxa and show that hybridization involving sexual diploid species, primarily S. aria and S. torminalis and polyploid facultative apomictic species from subgenus Aria, has been responsible for generating this biodiversity. Importantly our data show that this creative evolutionary process is ongoing within the Avon Gorge. Conservation strategies for the rare endemic Sorbus taxa should therefore consider all Sorbus taxa within the Gorge and must strive to preserve this evolutionary process rather than simply the individual rare taxa that it produces.