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Pea seed-borne mosaic virus (PSbMV), a nonpersistently aphid-transmitted potyvirus, has been reported in field pea (Pisum sativum L.)-growing regions worldwide. In 2014, PSbMV was first identified in field peas in North Dakota, U.S.A. Susceptibility and yield losses attributed to PSbMV infection are influenced by viral pathotype and host genotype. Isolate ND14-1, recovered from North Dakota infected seed and identified as pathotype 4 (P4), was mechanically inoculated onto 20 field pea cultivars under greenhouse conditions. PSbMV susceptibility, number of seeds and pods per plant, yield, symptom expression, and PSbMV seed transmission rates were assessed by cultivar. A risk assessment was developed based on cultivar susceptibility, yield reduction, and PSbMV seed transmission. Risk factors were weighted based on perceived importance to commercial field pea producers. Three cultivars were classified as low risk, seven cultivars were classified as intermediate risk, and 10 cultivars were classified as high risk. Two of the low-risk cultivars, Aragorn and Cruiser, were confirmed to be resistant to this isolate of PSbMV. Cultivar Arcadia was susceptible to PSbMV infection with mild expression of symptoms, but was classified as low risk based on a low seed transmission rate and diminished yield losses. This risk assessment could prove a useful tool for growers in field pea cultivar selection where PSbMV is prevalent.

Anthracnose, caused by the fungal pathogen Colletotrichum lindemuthianum, is a damaging seed-transmitted disease of dry beans that causes reduced seed quality and yield. Seed-to-seedling transmission of C. lindemuthianum has been documented as high as 15% in asymptomatic seeds under greenhouse conditions. Increasing pathogen colonization in seeds has been correlated with increasing anthracnose seed symptoms via quantitative PCR (qPCR), but stem colonization has not been quantified. Previous studies also have characterized seed yield and quality losses caused by planting C. lindemuthianum–infected seeds, but none evaluated the effect of growing asymptomatic seeds on disease and plant development under field conditions. A real-time qPCR assay was developed in this study and used to detect C. lindemuthianum in the stems of seedlings as early as 15 days after planting. Field trials measured the seed-to-seedling transmission of C. lindemuthianum across levels of anthracnose symptoms in seeds ranging from healthy to severely discolored. Results from these two field trials indicated that emergence and yield decreased and foliar symptoms, pathogen detection, and incidence of symptoms on progeny seeds increased as the severity of infection in planted seeds increased. In both years, planting asymptomatic seeds resulted in higher anthracnose severity than planting healthy seeds. Yield, seed weight, and incidence of symptoms on progeny seeds were not higher in asymptomatic seeds than in healthy seeds in 2014, when moderate disease pressure was observed. However, these factors were significantly different in 2015, when anthracnose severity was driven up to 75% by conducive weather conditions. This serves as a strong warning to growers that planting seed grown in a field where anthracnose was present, even if those seeds are asymptomatic, can result in yield and quality losses. Planting certified dry bean seed is always recommended.

Dry bean (Phaseolus vulgaris L.) production in many regions is threatened by white mold (WM) [Sclerotinia sclerotiorum (Lib.) de Bary]. Seed yield losses can be up to 100% under conditions favorable for the pathogen. The low heritability, polygenic inheritance, and cumbersome screening protocols make it difficult to breed for improved genetic resistance. Some progress in understanding genetic resistance and germplasm improvement has been accomplished, but cultivars with high levels of resistance are yet to be released. A WM multiparent advanced generation inter-cross (MAGIC) population (n = 1060) was developed to facilitate mapping and breeding efforts. A seedling straw test screening method provided a quick assay to phenotype the population for response to WM isolate 1980. Nineteen MAGIC lines were identified with improved resistance. For genome-wide association studies (GWAS), the data was transformed into three phenotypic distributions-quantitative, polynomial, and binomial-and coupled with ∼52,000 single-nucleotide polymorphisms (SNPs). The three phenotypic distributions identified 30 significant genomic intervals [-log

Dry bean (Phaseolus vulgaris L.) is an important worldwide legume crop with low to moderate levels of resistance to common bacterial blight (CBB) caused by Xanthomonas axonopodis pv. phaseoli. A total of 852 genotypes (cultivars, preliminary and advanced breeding lines) from the North Dakota State University dry bean breeding program were tested for their effectiveness as populations for genome-wide association studies (GWAS) to identify genomic regions associated with resistance to CBB, to exploit the associated markers for marker-assisted breeding (MAB), and to identify candidate genes. The genotypes were evaluated in a growth chamber for disease resistance at both the unifoliate and trifoliate stages. At the unifoliate stage, 35% of genotypes were resistant, while 25% of genotypes were resistant at the trifoliate stage. Libraries generated from each genotype were sequenced using the Illumina platform. After filtering for sequence quality, read depth, and minor allele frequency, 41,998 single-nucleotide polymorphisms (SNPs) and 30,285 SNPs were used in GWAS for the Middle American and Andean gene pools, respectively. One region near the distal end of Pv10 near the SAP6 molecular marker from the Andean gene pool explained 26.7–36.4% of the resistance variation. Three to seven regions from the Middle American gene pool contributed to 25.8–27.7% of the resistance, with the most significant peak also near the SAP6 marker. Six of the eight total regions associated with CBB resistance are likely the physical locations of quantitative trait loci identified from previous genetic studies. The two new locations associated with CBB resistance are located at Pv10:22.91–23.36 and Pv11:52.4. A lipoxgenase-1 ortholog on Pv10 emerged as a candidate gene for CBB resistance. The state of one SNP on Pv07 was associated with susceptibility. Its subsequent use in MAB would reduce the current number of lines in preliminary and advanced field yield trial by up to 14% and eliminate only susceptible genotypes. These results provide a foundational SNP data set, improve our understanding of CBB resistance in dry bean, and impact resource allocation within breeding programs as breeding populations may be used for dual purposes: cultivar development as well as genetic studies.

Multiple plant pathogens cause root rot on field pea, including Aphanomyces euteiches, Rhizoctonia solani and numerous Fusarium spp. In North Dakota, root rot pathogens have been a contributing fac...

Meiotic crossovers (COs) are not uniformly distributed across the genome. Factors affecting this phenomenon are not well understood. Although many species exhibit large differences in CO numbers between sexes, sex-specific aspects of CO landscape are particularly poorly elucidated. Here, we conduct high-resolution CO mapping in maize. Our results show that CO numbers as well as their overall distribution are similar in male and female meioses. There are, nevertheless, dissimilarities at local scale. Male and female COs differ in their locations relative to transcription start sites in gene promoters and chromatin marks, including nucleosome occupancy and tri-methylation of lysine 4 of histone H3 (H3K4me3). Our data suggest that sex-specific factors not only affect male–female CO number disparities but also cause fine differences in CO positions. Differences between male and female CO landscapes indicate that recombination has distinct implications for population structure and gene evolution in male and in female meioses.

The species cytoplasm specific (scs) genes affect nuclear-cytoplasmic interactions in interspecific hybrids. A radiation hybrid (RH) mapping population of 188 individuals was employed to refine the location of the scs ae locus on Triticum aestivum chromosome 1D. “Wheat Zapper,” a comparative genomics tool, was used to predict synteny between wheat chromosome 1D, Oryza sativa, Brachypodium distachyon, and Sorghum bicolor. A total of 57 markers were developed based on synteny or literature and genotyped to produce a RH map spanning 205.2 cR. A test-cross methodology was devised for phenotyping of RH progenies, and through forward genetic, the scs ae locus was pinpointed to a 1.1 Mb-segment containing eight genes. Further, the high resolution provided by RH mapping, combined with chromosome-wise synteny analysis, located the ancestral point of fusion between the telomeric and centromeric repeats of two paleochromosomes that originated chromosome 1D. Also, it indicated that the centromere of this chromosome is likely the result of a neocentromerization event, rather than the conservation of an ancestral centromere as previously believed. Interestingly, location of scs locus in the vicinity of paleofusion is not associated with the expected disruption of synteny, but rather with a good degree of conservation across grass species. Indeed, these observations advocate the evolutionary importance of this locus as suggested by “Maan’s scs hypothesis.”

In this study we assess the genetic architecture of bread-making quality traits in spring wheat (Triticum aestivum L.). A mapping population derived from BR34 and 'Grandin' was used to measure 20 end-use quality traits including six kernel, seven milling and fl our, four dough mixing strength, and three bread-making traits. A total of 31 quantitative trait loci (QTL) signifi cantly associated with all but two traits were identifi ed. These QTL were clustered in fi ve chromosomal regions, namely 1BS, 1DL, 4BL, 5BL, and 6AS, and explained a large proportion of trait variation with favorable alleles contributed by both parents. The 1DL cluster containing the high molecular weight glutenin gene, Glu-D1, had a large genetic infl uence on dough mixing strength and bread-making performance. Most of the QTL affecting kernel traits were clustered on 6AS. Inconsistency of QTL locations detected from different environments was observed for the fl our and milling traits and was likely due to genotype × environment interaction (G × E) effects. Despite high heritabilities estimated for the 20 quality traits evaluated, no QTL were found for fl our brightness and bake water absorption, suggesting that these traits may be controlled by QTL with small effects that could not be detected due to the small population size. Because of the complex inheritance of these traits, it will be necessary to validate these QTL in different spring wheat backgrounds evaluated in similar growth conditions as used in this study before the marker information can be used for breeding applications.

Wheat stem rust caused by Puccinia graminis f. sp. tritici, can cause significant yield losses. To combat the disease, breeders have deployed resistance genes both individually and in combinations to increase resistance durability. A new race, TTKSK (Ug99), identified in Uganda in 1999 is virulent on most of the resistance genes currently deployed, and is rapidly spreading to other regions of the world. It is therefore important to identify, map, and deploy resistance genes that are still effective against TTKSK. One of these resistance genes, Sr13, was previously assigned to the long arm of chromosome 6A, but its precise map location was not known. In this study, the genome location of Sr13 was determined in four tetraploid wheat (T. turgidum ssp. durum) mapping populations involving the TTKSK resistant varieties Kronos, Kofa, Medora and Sceptre. Our results showed that resistance was linked to common molecular markers in all four populations, suggesting that these durum lines carry the same resistance gene. Based on its chromosome location and infection types against different races of stem rust, this gene is postulated to be Sr13. Sr13 was mapped within a 1.2-2. 8 cM interval (depending on the mapping population) between EST markers CD926040 and BE471213, which corresponds to a 285-kb region in rice chromosome 2, and a 3.1-Mb region in Brachypodium chromosome 3. These maps will be the foundation for developing high-density maps, identifying diagnostic markers, and positional cloning of Sr13. © 2010 US Government.

Plant disease resistance is often conferred by genes with nucleotide binding site (NBS) and leucine-rich repeat (LRR) or serine/threonine protein kinase (S/TPK) domains. Much less is known about mechanisms of susceptibility, particularly to necrotrophic fungal pathogens. The pathogens that cause the diseases tan spot and Stagonospora nodorum blotch on wheat produce effectors (host-selective toxins) that induce susceptibility in wheat lines harboring corresponding toxin sensitivity genes. The effector ToxA is produced by both pathogens, and sensitivity to ToxA is governed by the Tsn1 gene on wheat chromosome arm 5BL. Here, we report the cloning of Tsn1 , which was found to have disease resistance gene-like features, including S/TPK and NBS-LRR domains. Mutagenesis revealed that all three domains are required for ToxA sensitivity, and hence disease susceptibility. Tsn1 is unique to ToxA-sensitive genotypes, and insensitive genotypes are null. Sequencing and phylogenetic analysis indicated that Tsn1 arose in the B-genome diploid progenitor of polyploid wheat through a gene-fusion event that gave rise to its unique structure. Although Tsn1 is necessary to mediate ToxA recognition, yeast two-hybrid experiments suggested that the Tsn1 protein does not interact directly with ToxA. Tsn1 transcription is tightly regulated by the circadian clock and light, providing further evidence that Tsn1 -ToxA interactions are associated with photosynthesis pathways. This work suggests that these necrotrophic pathogens may thrive by subverting the resistance mechanisms acquired by plants to combat other pathogens.

The wheat-Stagonospora nodorum pathosystem involves a number of pathogen-produced host-selective toxins that interact with host genes in an inverse gene-for-gene manner to cause disease. The wheat intervarietal recombinant inbred population derived from BR34 and Grandin (BG population) segregates for the toxin sensitivity genes Tsn1, Snn2, and Snn3, which confer sensitivity to the toxins ToxA, SnTox2, and SnTox3, respectively. Here, we report the addition of 141 molecular markers to the BG population linkage maps, the identification and/or development of markers tightly linked to Tsn1 and Snn2, and the validation of the markers using a set of diverse wheat accessions. The BG population maps now contain 787 markers, and new simple sequence repeat (SSR) markers closely linked to Snn2 on chromosome arm 2DS were identified. In an effort to target more markers to the Snn2 locus, STS markers were developed from 2DS bin-mapped ESTs resulting in the development and mapping of 36 markers mostly to the short arms of group 2 chromosomes. Together, SSR and EST-STS markers delineated Snn2 to a 4.0 cM interval. SSRs developed in related work for Tsn1 were mapped in the BG population and delineated the gene to a 1.0 cM interval. Evaluation of the markers for Tsn1 and Snn2 in a diverse set of wheat genotypes validated their utility for marker-assisted selection, which is particularly efficient for removing toxin sensitivity alleles from elite germplasm and varieties.

The cereal crops wheat, rice, maize, and sorghum show conservation of large syntenic blocks in spite of more than 40-fold variation in genome and 20-fold variation in chromosome size. It has been proposed that independent mutations at orthologous loci in traits such as shattering, tough fruiting cases (glumes, in the case of wheat), and threshing may have led to domestication-driven convergent evolution. A different picture is emerging from comparative mapping and cloning of these genes in different cereal crops. It appears that these spike traits are controlled by multiple genetic pathways, and mutations at different loci have been selected during domestication-driven evolution.

Cereal grains are a significant source of ochratoxin A (OTA) in the human diet. Multiple ochratoxigenic Aspergillus and Penicillium spp. have been reported as contaminants on various cereal grains around the world, although relatively few species dominate in any given location. Efforts to mitigate the risk of fungal contamination and OTA accumulation can be made pre- and postharvest. Still, a rapid and reliable screening method is sought that can be used to predict the OTA level of a sample and to inform risk assessments prior to processing. In this study, we assessed the efficacy of two OTA-related indices for OTA level prediction. Infestation rates were determined by direct plating for freshly harvested and stored barley, durum, and hard red spring wheat samples (n = 139) with known OTA levels. Presumptive ochratoxigenic isolates were tested for their ability to produce OTA. The nonribosomal peptide synthase (otanpsPN) involved in OTA biosynthesis was used to quantify ochratoxigenic fungi in barley and wheat. Viable Penicillium verrucosum was present in 45% of the samples. In total, 62.7% (n = 110) of the P. verrucosum isolates tested produced OTA on dichloran yeast extract sucrose 18% glycerol agar. Both OTA level and infestation rate (r = 0.30), as well as OTA level and otanpsPN concentration (r = 0.56), were weakly correlated. Neither infestation rate nor otanpsPN concentration is a reliable predictor of OTA level in a sample.

Wild related Triticum species have been and will be a useful reservoir of genetic diversity for solving problems in the development of improved durum (Triticum turgidum L. var. durum) and bread wheat (T. aestivum L.) cultivars. The alien germplasm may be used to reduce vulnerability of cultivars to pests, and improve agronomic fitness and grain quality. However, lack of genetic recombination and hybrid sterility are two obstacles to the use of alien germplasms. Nuclear-cytoplasmic (NC) incompatibility is known to exist between the T. longissimum S. & M. cytoplasm and T. turgidum nucleus. A two-gene system has been found that restores fertility in this situation: the species cytoplasm specific (scs) gene and the vitality (Vi) gene. This gene system can also be used for the production of hybrid wheat. Effects of these genes on euplasmic (true cytoplasm) durum wheat under a field environment had not been previously investigated. In this study, lines with two copies of the scs(ti) gene, derived from T. timopheevii Zhuk., were compared to lines without the copy of the gene, the parents, and four durum cultivars for five agronomic characteristics: days to heading, plant height, lodging resistance, grain yield, and kernel test weight. Comparison of genotypes homozygous for the scs(ti) gene and those containing no scs(ti) gene indicated minor differences among them, confirming that the scs(ti) gene does not confer any detrimental effects in the euplasmic situation. Thus, the scs(ti) gene could be useful in the production of hybrids in durum wheat.

The action of species cytoplasm specific (scs) gene(s) can be observed when a durum (Triticum turgidum L.) nucleus is placed in the Aegilops longissimum S. & M. cytoplasm. This alloplasmic combination, (lo) durum, results in nonviable progeny. A scs gene derived from T. timopheevii Zhuk. (scs(ti)) produced compatibility with the (lo) cytoplasm. The resulting hemizygous (lo) scs(ti)- durum line was male sterile and when crossed to normal durum produced a 1:1 ratio of plump, viable (PV) seeds with scs(ti) and shriveled inviable (SIV) seeds without scs(ti). In a systematic characterization of durum lines an unusual line was identified that when crossed to (lo) scs(ti)- produced all PV seeds. When planted these PV seeds segregated at a 1:1 ratio of normal vigor plants (NVPs) and low vigor plants (LVPs). The LVP senescence before full maturity. The NVPs were male sterile and when crossed to common durum lines resulted in all plump seeds that again segregated at a 1:1 ratio of NVPs to LVPs. The crosses of these NVPs to common durum lines resulted in a 1:1 ratio of PV to SIV seeds. This study was extended to 317 individuals segregating for scs(ti) and the new locus, derived from durum wheat (scs(d)), establishing the allelic relationship of these two genes.

The compatibility-inducing action of the scsti (species cytoplasm-specific gene derived from Triticum timopheevii) and Vi (vitality) genes can be observed when a durum (T. turgidum) nucleus is placed in T. longissimum cytoplasm. These two genes restore compatibility between an otherwise incompatible nucleus and cytoplasm. The objective of this study was to localize the scsti gene on a linkage map of chromosome 1A, which could eventually be used to clone the gene. The mapping population consisted of 110 F2 individuals derived from crossing a Langdon-T. dicoccoides chromosome 1A substitution line with a euplasmic (normal cytoplasm) line homozygous for the scsti gene. Through a series of testcrosses the genotypes of the 110 individuals were determined: 22 had two copies, 59 had one copy, and 29 had no copy of the scsti gene. Data from RFLP, AFLP, and microsatellite analysis were used to create a linkage map. The flanking marker loci found for the scsti gene were Xbcd12 and Xbcd1449-1A.2 with distances of 2.3 and 0.6 cM, respectively. Nearly 10% of individuals in this population were double recombinant for a genetic interval of

Radiation has been used as a mean to break and transfer fragments of DNA from one plant species to another. Early examples include the experiments by Sears, (Brookhaven Symp Biol 9:1–22, 1956) to transfer rust resistance genes from Aegilops umbellulata to wheat. Radiation found its niche as a mutagen due to advances in nuclear technology and formation of the International Atomic Energy Agency and their sponsorship of developing mutation breeding through “Mutation Enhanced Technologies for Agriculture”. Mutation breeding has resulted in the release of several important cultivars. Although radiation was used in plants for the mutation and introgression of genes from related species (Sears, Brookhaven Symp Biol 9:1–22, 1956; Driscoll and Jensen, Genetics 48:459–468, 1963; Riley and Law, Stadler Genet Symp 16:301–322, 1984; Sears, Crop Sci 33:897–901, 1993), this approach was not used for mapping. This aspect of radiation application was first utilized in animal cell culture lines to generate radiation hybrid (RH) panels. In the beginning these panels were generated for single chromosomes but evolved to the development of whole genome panels. This technology matured in animal systems with the onset of genomics era by its use in the development of high resolution RH-based physical maps for many species before or during the development of complete genome sequence information. The advantages of this system are: (1) radiation-induced breaks are independent of recombination events providing higher and more uniform resolution, (2) radiation dosage could be adjusted to provide varied resolution without greatly affecting the population size and (3) all markers regardless of their polymorphism can be mapped on RH panels. Plant scientists followed these studies by the development of RH panels for individual chromosomes or whole genomes. However, early RH panels in plant systems were of low to medium resolution and of limited use in physical mapping. Recently, RH panels have been produced resulting in map resolutions of 200–400 Kb. These high resolution panels promise the same value as animal systems in helping generate a complete genome sequence with a fraction of the cost of traditional methods. But the use of radiation in plants has matured to go beyond physical mapping by its application to gene cloning and forward/reverse genetic studies. These applications take advantage of plasticity offered by many plant species in tolerating radiation to produce seed and live progeny. This ability allows scientists to phenotype RH lines and to associate the phenotypic data with the genotypic data. The great potential of this system is just being realized.

The Q gene is largely responsible for the widespread cultivation of wheat because it confers the free-threshing character. It also pleiotropically influences many other domestication-related traits such as glume shape and tenacity, rachis fragility, spike length, plant height, and spike emergence time. We isolated the Q gene and verified its identity by analysis of knockout mutants and transformation. The Q gene has a high degree of similarity to members of the AP2 family of transcription factors. The Q allele is more abundantly transcribed than q, and the two alleles differ for a single amino acid. An isoleucine at position 329 in the Q protein leads to an abundance of homodimer formation in yeast cells, whereas a valine in the q protein appears to limit homodimer formation. Ectopic expression analysis allowed us to observe both silencing and overexpression effects of Q. Rachis fragility, glume shape, and glume tenacity mimicked the q phenotype in transgenic plants exhibiting post-transcriptional silencing of the transgene and the endogenous Q gene. Variation in spike compactness and plant height were associated with the level of transgene transcription due to the dosage effects of Q. The q allele is the more primitive, and the mutation that gave rise to Q occurred only once leading to the world's cultivated wheats.

The objectives of this study were to develop a high-density chromosome bin map of homoeologous group 7 in hexaploid wheat (Triticum aestivum L.), to identify gene distribution in these chromosomes, and to perform comparative studies of wheat with rice and barley. We mapped 2148 loci from 919 EST clones onto group 7 chromosomes of wheat. In the majority of cases the numbers of loci were significantly lower in the centromeric regions and tended to increase in the distal regions. The level of duplicated loci in this group was 24% with most of these loci being localized toward the distal regions. One hundred nineteen EST probes that hybridized to three fragments and mapped to the three group 7 chromosomes were designated landmark probes and were used to construct a consensus homoeologous group 7 map. An additional 49 probes that mapped to 7AS, 7DS, and the ancestral translocated segment involving 7BS also were designated landmarks. Landmark probe orders and comparative maps of wheat, rice, and barley were produced on the basis of corresponding rice BAC/PAC and genetic markers that mapped on chromosomes 6 and 8 of rice. Identification of landmark ESTs and development of consensus maps may provide a framework of conserved coding regions predating the evolution of wheat genomes.

Background Development of a high quality reference sequence is a daunting task in crops like wheat with large (~17Gb), highly repetitive (>80%) and polyploid genome. To achieve complete sequence assembly of such genomes, development of a high quality physical map is a necessary first step. However, due to the lack of recombination in certain regions of the chromosomes, genetic mapping, which uses recombination frequency to map marker loci, alone is not sufficient to develop high quality marker scaffolds for a sequence ready physical map. Radiation hybrid (RH) mapping, which uses radiation induced chromosomal breaks, has proven to be a successful approach for developing marker scaffolds for sequence assembly in animal systems. Here, the development and characterization of a RH panel for the mapping of D-genome of wheat progenitor Aegilops tauschii is reported. Results Radiation dosages of 350 and 450 Gy were optimized for seed irradiation of a synthetic hexaploid (AABBDD) wheat with the D-genome of Ae. tauschii accession AL8/78. The surviving plants after irradiation were crossed to durum wheat (AABB), to produce pentaploid RH1s (AABBD), which allows the simultaneous mapping of the whole D-genome. A panel of 1,510 RH1 plants was obtained, of which 592 plants were generated from the mature RH1 seeds, and 918 plants were rescued through embryo culture due to poor germination (1 seeds. This panel showed a homogenous marker loss (2.1%) after screening with SSR markers uniformly covering all the D-genome chromosomes. Different marker systems mostly detected different lines with deletions. Using markers covering known distances, the mapping resolution of this RH panel was estimated to be cM/cR ratio of 1:5.2 and 15 distinct bins. Additionally, with this small set of lines, almost all the tested ESTs could be mapped. A set of 399 most informative RH lines with an average deletion frequency of ~10% were identified for developing high density marker scaffolds of the D-genome. Conclusions The RH panel reported here is the first developed for any wild ancestor of a major cultivated plant species. The results provided insight into various aspects of RH mapping in plants, including the genetically effective cell number for wheat (for the first time) and the potential implementation of this technique in other plant species. This RH panel will be an invaluable resource for mapping gene based markers, developing a complete marker scaffold for the whole genome sequence assembly, fine mapping of markers and functional characterization of genes and gene networks present on the D-genome.