Molecular mapping of powdery mildew resistance genes in wheat: A review

Powdery mildew, caused by Blumeria graminis f. sp. tritici (syn. Erysiphe graminis f. sp. tritici), is one of the most important diseases of common wheat (Triticum aestivum L.) worldwide. Molecular mapping and cloning of genes for resistance to powdery mildew in hexaploid wheat will facilitate the study of molecular mechanisms underlying resistance to powdery mildew diseases and help understand the structure and function of powdery mildew resistance genes, and permit marker-assisted selection in breeding programs. So far, 48 genes/alleles for resistance to powdery mildew at 32 loci have been identified and located on 16 different chromosomes, of which 21 resistance genes/alleles have been tagged by restriction fragment length polymorphisms (RFLPs), random-amplified polymorphic DNAs (RAPDs), amplified fragment length polymorphisms (AFLPs), sequence characterized amplified regions (SCARs), sequence-tagged sites (STS) or simple sequence repeats (SSRs). Several quantitative trait loci (QTLs) for adult plant resistance (APR) to powdery mildew have been associated with molecular markers. The detailed information on chromosomal location and molecular mapping of these genes has been reviewed. Isolation of powdery mildew resistance genes and development of valid molecular markers for pyramiding resistance genes in breeding programs is also discussed.     

Stem solidness and its relationship to grain yield in 17 spring wheat crosses

Summary Stem solidness in the wheat plant provides resistance to the wheat stem sawfly, a pest of wheat in Montana and North Dakota, but some agronomists have been concerned that stem solidness might be related to low grain yields. We evaluated 17 spring wheat crosses for stem solidness, grain yield, and other agronomic traits in F₂ through F₄ generations from 1972 to 1975.Highly significant correlation (0.735) and regression (0.731) coefficients between F₂ and F₃ generations verify previous studies showing that stem solidness is highly heritable and that selection in F₂ should be successful. Solid-stemmed F₄ composites yielded more than hollow stemmed composites at Bozeman and equal yields were obtained at Huntley, indicating that stem solidness is not a deterrent to high grain yield.Joint contribution of Federal Research, Science and Education Administration, U.S. Department of Agriculture and the Montana Agricultural Experiment Station, Bozeman, MT 59717. Published with approval of the Director of the Montana Agricultural Experiment Station as Paper No. 815, Journal Series.     

Generation mean analysis to identify and partition the components of genetic resistance to Septoria nodorum in wheat

Summary Foliar symptom severity of seedlings artificially inoculated with S. nodorum were used to idenify the type of gne action controlling resistance to this pathogen in the early generations of two wheat crosses. In both crosses a resistant spring wheat cultivar was crossed to a susceptible cultivar. Reciprocal crosses were included in the analysis to determine if the cytoplasm contributed in any significant degree to the level of resistance present.Results indicated that resistance was polygenic and that it could be explaned prinerpally by additive gene effects. Some differences in reciprocal crosses were evident, but a significant role for the eytoplasm in resistance is not indicated.Cooperative investigations of the Agricultural Research Service of the U. S. Department of Agriculture and the Montana Agricultural Experiment Station. Journal Paper No. 1210.     

Isolation of wheat mutants with increased resistance to powdery mildew from small induced variant populations

Small variant wheat populations created by induced mutagenesis (n = 69) or adventitious regeneration (n = 66) were intensively screened for an altered response (compared to the parent variety ‘Guardian’) to the causal pathogen of powdery mildew in wheat, Blumeria graminis f. sp. tritici. Intensive field screening following natural infection of replicated plots of wheat lines over two years revealed a total of 13 mutants exhibiting significantly greater resistance than ‘Guardian’: eight from induced mutagenesis (11.6%) of the M2 population and five from adventitious regeneration (7.6%). Complete resistance was identified in two lines, (one (M66) developed following induced mutagenesis, and the other (SC240) by adventitious regeneration). The complete resistance in the induced mutant was stable over two generations and was associated with a high frequency of leaf flecking, and consequently a low grain yield. Resistance in SC240 proved to be unstable; SC240 exhibited complete resistance to powdery mildew in the SC₂ and SC₃ generations, but only 20% of the SC₄ plants were completely resistant, while the remainder were indistinguishable in mildew response to ‘Guardian’. The mildew response of all the SC₅ generation of SC240 was not significantly different from ‘Guardian’. Yield analysis of the thirteen mutants with increased resistance in the presence of powdery mildew indicated that eleven exhibitedgrain yields at least as high as that of ‘Guardian’, while the mutant M19 exhibited a yield significantly higher than that of ‘Guardian’. This revised version was published online in July 2006 with corrections to the Cover Date.     

Molecular characterization of a novel powdery mildew resistance gene Pm30 in wheat originating from wild emmer

Powdery mildew caused by Erysiphe graminis f. sp. tritici is one of the most important wheat diseases in many regions of theworld. A powdery mildew resistance gene, originating from wild emmerwheat (Triticum dicoccoides) accession `C20', from Rosh Pinna, Israel,was successfully transferred to hexaploid wheat through crossing andbackcrossing. Genetic analysis indicated that a single dominant genecontrols the powdery mildew resistance at the seedling stage. SegregatingBC₁F₂ progenies of the cross 87-1/C20//2*8866 wereused for bulked segregant analysis (BSA). The PCR approach was used togenerate polymorphic DNA fragments between the resistant and susceptibleDNA pools by use of 10-mer random primers, STS primers, and wheatmicrosatellite primers. Three markers, Xgwm159/430,Xgwm159/460, and Xgwm159/500, were found to be linked tothe resistance gene. After evaluating the polymorphic markers in twosegregating populations, the distance between the markers and the mildewresistance gene was estimated to be 5–6 cM. By means of ChineseSpring nullisomic-tetrasomics and ditelosomics, the polymorphic markersand the resistance gene were assigned to chromosome arm 5BS and werephysically mapped on the gene rich regions of fragment length (FL) 0.41–0.43 by Chinese Spring deletion lines. As no powdery mildew resistancegene has been reported on chromosome arm 5BS, the mildew resistancegene originating from C20 should be a new gene and is designated Pm30.     

Identification of QTLs for powdery mildew and scald resistance in barley

A population of 103 recombinant inbred lines (RILs, F₉-derived lines) developed from the two-row spring barley cross L94 × ‘Vada’ was evaluated under field conditions for resistance against powdery mildew (Blumeria graminis f.sp. hordei) and scald (Rhynchosporium secalis). Apart from the major resistance gene mlo on chromosome 4 (4H), three QTLs (Rbgq1, Rbgq2 and Rbgq3) for resistance against powdery mildew were detected on chromosomes 2 (2H), 3 (3H), and 7 (5H), respectively. Rbgq1 and Rbgq2 have not been reported before, and did not map to a chromosome region where a major gene for powdery mildew had been reported. Four QTLs (Rrsq1, Rrsq2, Rrsq3 and Rrsq4) for resistance against scald were detected on chromosomes 3 (3H), 4 (4H) and 6 (6H). All four mapped to places where QTLs for scald resistance had been reported before in different populations.     

Two unnecessary powdery mildew resistance genes in a synthetic rye population are neutral on fitness

A synthetic winter rye population was produced with two race-specific powdery mildew resistance genes, one dominant (Rm1) and the other (rm2) recessive, each at a frequency of about 0.50. The population was advanced by open-pollination in an isolated plot under mildew-free conditions for eight years. Samples of generations Syn-0 through Syn-7 were inoculated in the laboratory with two mildew isolates, one avirulent to either resistance gene, the other virulent to Rm1 and avirulent to rm2, to discriminate resistant and susceptible phenotypes. From the proportions of resistant plants, frequencies of Rm1 and rm2 were calculated and the fitness of carriers of resistance alleles was estimated in relation to carriers of susceptibility alleles at the two loci using continuous models and linear regression analyses. Frequencies of the two resistance genes oscillated only weakly over the eight generations. Coefficients of selection against Rm1-and rm2rm2 genotypes were –0.04 and –0.02, respectively, and not significantly different from zero. Thus the two resistance genes were selectively neutral. It is concluded that pyramiding of major powdery mildew resistance genes in rye varieties should not reduce their yield potential in the absence of mildew.     

Interspecific hybrids of Zinnia elegans Jacq. and Z. angustifolia HBK: embryology,morphology and powdery mildew resistance

Summary Reciprocal crosses between 13 cultivars of Zinnia elegans Jacq. and one source of Z. angustifolia HBK resulted in interspecific hybrids. Florets examined 14 days after pollination showed that all 13 cross combinations of Z. angustifolia x Z. elegans yielded embryos in 47.2% of the florets pollinated. Six cross combinations of Z. elegans x Z. angustifolia produced embryos in 4.6% of the total Z. elegans florets pollinated. Interspecific crosses resulted in seed-propagated hybrids when Z. angustifolia was the maternal parent. Hybrids were obtained from reciprocal crosses between the two species when embryos were excised and cultured. Morphologically, most hybrids resembled Z. angustifolia but were sterile. Colchicine treatments of selected sterile hybrids resulted in shoots producing fertile inflorescences. Preliminary tests showed that hybrids were resistant to powdery mildew (Erysiphe cichoracearum DC ex Mecat).Scientific Article No. A 3108, Contribution No. 6173 of the Maryland Agricultural Experiment Station, Department of Horticulture.     

New molecular markers linked to qualitative and quantitative powdery mildew and scald resistance genes in barley for dry areas

A partial genetic linkage map was constructed on 71 doubled-haploid lines derived from a cross between the barley lines Tadmor and WI2291 with 181 molecular markers. The segregating population was used to detect markers linked to the gene Mlg conferring resistance to powdery mildew (Erysiphe graminis f. sp. hordei) and to genes for quantitative resistance to scald (Rhynchosporium secalis). The gene Mlg on chromosome 4H was flanked by two AFLP markers at a distance of 2.0 and 2.4 cM, respectively. QTLs for resistance to scald were detected on chromosomes 2H and 3H. This association of molecular markers with qualitative and quantitative disease resistance loci represents a valuable starting-point for marker-assisted selection. This revised version was published online in July 2006 with corrections to the Cover Date.     

Identification of Resistance Genes Against Powdery Mildew in Four Accessions of Hordeum Vulgare SSP. Spontaneum

Summary Four newly detected accessions of wild barley (Hordeum vulgare ssp. spontaneum) resistant to powdery mildew caused by Blumeria graminis f. sp. hordei were studied with the aim of finding the number of genes/loci conferring the resistance of individual accessions, the type of inheritance of the genes and their relationships to the Mla locus. F₂ populations after crosses between the winter variety ‘Tiffany’ and four wild barley accessions and use of microsatellite DNA markers were focused on the identification of individual resistance genes/loci by means of their chromosomal locations. In PI466495, one locus conferring powdery mildew resistance was identified in highly significant linkage with the marker Bmac0213. This location is consistent with the known locus Mla on chromosome 1HS. In the other three accessions the resistance was determined by two independent loci. In PI466197, PI466297 and PI466461, one locus was identified on chromosome 1HS and three new loci were revealed on chromosomes 2HS (highly significant linkage with Bmac0134), 7HS (highly significant linkage with Bmag0021) and 7HL (significant linkage with EBmac0755). Our prospective aim is identification of further linked DNA markers and the exact location of the resistance genes on the barley chromosomes.     

The oat line Pc54 as a source of resistance to crown rust,stem rust and powdery mildew in Europe

Summary The oat line Pc54 was found to be resistant to powdery mildew under both field and glasshouse conditions. The ratio of resistant to susceptible F₂ and F₂ progeny of a cross between a selection from the Pc54 line (Cc7422) and a susceptible cultivar (Selma) showed that, in addition to carrying the crown rust resistance gene Pc54 and the pg15 gene for stem rust resistance, the mildew resistance of the Pc54 line was conditioned by a single incompletely dominant gene along with additional factors which modified the expression of resistance. Previous results, that there was no linkage between genes Pc54 and Pg15, were confirmed. In addition, there was no evidence of linkage between the mildew resistance gene and gene Pc54. Evaluation of selections from within the Pc54 line showed that the expression of both stem rust and mildew resistance was modified by, or linked to, plant height. The effectiveness of genes Pc54 and Pg15, as measured by virulence frequencies, in central and eastern Europe is described.     

Characterisation of new oat germplasm for resistance to powdery mildew

Summary Wild oat accessions from the Institute of Plant Genetics and Crop Plant Research (IPK) Gatersleben were selected as potential sources of resistance to powdery mildew (Erysiphe graminis f.sp. avenae). The Avena strigosa accessions AVE 128, AVE 488 and AVE 264, along with A. occidentalis accession CAV 3889 showed high levels of mildew resistance in tests at early and late growth stages over two years; the latter is a particularly useful source because it is a hexaploid species. Moderate levels of resistance were also found in A. fatua accession AVE 1981, AVE 2032 and A.sterilis accession AVE 1373 but this could be due to the later maturity of these genotypes.     

Genetic analysis of adult plant resistance to powdery mildew in pea (Pisum sativum L.)

Summary An analysis of adult plant resistance of powdery mildew in 15 F₁, F₂ and F₃ populations of pea derived from crossing 15 diverse and susceptible lines with one resistant line revealed that resistance to powdery mildew is controlled by duplicate recessive genes. The genes were designated as er₁ and er₂.Disease reaction showed independent segregation with three known markers in the resistant parent, namely, af (afila, chromosome 1), st (stipule reduced, chromosome 3) and tl (clavicula, chromosome 7).Contribution form the Department of Genetics and Plant Breeding Banaras Hindu University, Varanasi-221005, India.     

Identification and chromosomal locations of novel genes for resistance to powdery mildew and stripe rust in a wheat line 101-3

Wheat powdery mildew and stripe rust, caused by Blumeria graminis f.sp.tritici (syn. Erysiphe graminis f.sp.tritici) and Puccinia striiformis Westend., respectively, are two important fungal diseases of wheat in many regions in the world that cause significant annual yield losses. In the present study, a dominant powdery mildew and a dominant stripe rust resistance gene in wheat line 101-3 which derived from the progenies of the wide cross between common wheat and Dasypyrum villosum Candary L., was located on chromosome 6B and 1B, respectively, by monosomic analyses. The two genes are different from known resistance genes on chromosome 6B for powdery mildew and 1B for stripe rusts, suggesting that the two genes might be novel resistance genes for powdery mildew and stripe rust, respectively. It is uncertain whether the two genes are allelic or lined with other resistance genes located on chromosome 6B for powdery mildew and 1B for stripe rust. Further allelism tests are necessary to determine the relationships between the resistance gene and other genes located on chromosome 6B for powdery mildew and 1B for stripe rust through molecular markers.     

Flag leaf area in five spring wheat crosses and the relationship to grain yield

Summary Five spring wheat crosses were evaluated over a 6-year period using comparisons between F₂ and F₃ data and between near-isogenic F₄ populations selected for flag leaf area. Nonsignificant r values for F₂ vs. F₃ flag leaf measurements may be due to the effect of environment on flag leaf area, but are probably also an indication of low heritability for this plant character. Near-isogenic populations selected on the basis of flag leaf area showed little difference in grain yield, an indication that other plant parts must be more influential in determining grain yield. Flag leaf area, by itself, appears not to be a good index to plant performance.Joint contribution of ARS/USDA and the Montana Agricultural Experiment Station. Published with approval of the Director of the Montana Agricultural Experiment Station as Paper No. 725. Journal Series. Use of data from Research Centers at Moccasin. Havre. Huntley, and Kalispell is gratefully acknowledged.     

The effect of the ‘Golden Promise’ dwarfing gene on partial resistance to leaf rust and powdery mildew in spring barley

Summary F₂-plant progenies, derived from seven crosses between susceptible erectoides breeding lines and conventional, nutans breeding lines with partial resistance to leaf rust incited by Puccinia hordei Otth., derived from Vada x Cebada Capa, were examined in a field and a glasshouse experiment. In the field experiment, the nutans plant progenies generally were more resistant to leaf rust and powdery mildew than the erectoides progenies. In the glasshouse experiment, the components of partial resistance to leaf rust of six erectoides and six nutans progenies derived from one cross were studied. The nutans progenies generally had lower infection frequencies than erectoides progenies with a similar infection level in the field experiment. The variation for leaf rust and powdery mildew infection in the field was similar for the nutans and erectoides progenies. This suggests that erectoides lines with a fairly high level of partial resistance to leaf rust and powdery mildew can be selected from these populations.     

Hordeum chilense resistance to powdery mildew and its potential use in cereal breeding

Summary Hordeum chilense is a wild barley with high crossability with Triticum, Hordeum and Secale. Its amphiploid with wheat, tritordeum, has potential as a new crop. H. chilense is highly resistant to the powdery mildew diseases of both wheat and barley. Whereas tritordeum is resistant to barley powdery mildew, its reaction to wheat powdery mildew is similar to that of its wheat parent. However H. chilense contributes to a reduced density of mildew colonies. This quantitative resistance of tritordeum is diluted at higher ploidy levels.     

Search for novel genes for resistance to powdery mildew (Sphaerotheca fuliginae) in cucumber (Cucumis sativus)

Summary The current powdery mildew (Sphaerotheca fuligninea) resistant cucumber varieties suffer from leaf chlorosis during autumn, winter and early spring cultivation in the Netherlands. Therefore screening was carried out for novel powdery mildew resistance genes. From 177 accessions, derived from different sources, 108 accessions proved to be partially resistant to S. fuliginea. Crosses were made with 53 resistant accessions to distinguish the presence of novel genes. It is likely that the accessions C. sativus 2145, C. sativus LV 41, PI 188807, Vladivostokij, White and Yellow 1 have one or more recessive powdery mildew resistance genes, different from powdery mildew resistance genes of the line NPI, which was used for variety breeding. Powdery mildew resistance tests with S. fuliginea give similar results in different regions of the world.Abbreviations pmr powdery mildew resistance     

Single-gene resistance to powdery mildew in zucchini squash (Cucurbita pepo)

Cucurbita pepo (pumpkin, squash,gourd) is an economically important species that is susceptible to the cucurbit powdery mildew fungus, Podosphaera xanthii(syn. Sphaerotheca fuliginea). ‘True French’, an open-pollinated cultivar of the Zucchini Group of C. pepo, was crossed with an unnamed powdery-mildew resistant straight neck-type accession, the resistance of which was apparently derived from an interspecific cross with a resistant wild species of Cucurbita,and resistant plants were selected in the F₂ generation. This was followed by six cycles of backcross-pedigree selection for resistance, and resulted in the development of an accession true-breeding for resistance to powdery mildew and nearly isogenic to ‘True French’. The resistant and susceptible near-isogenics were crossed and seeds of the filial and backcross generations were produced. Plants of the parental accessions and their progenies were grown together in a controlled-environment chamber, exposed to the pathogenic fungus, and scored as resistant, partially resistant, or susceptible 27–33 days after sowing. The results indicated that resistance is conferred by a single incompletely dominant gene, designated Pm-0. This revised version was published online in August 2006 with corrections to the Cover Date.     

Spectrum of resistance conferred by ML-O powdery mildew resistance genes in barley

Summary Ten barley mutants and five Ethiopian barley lines representing 11 independently arisen powdery mildew resistance genes in the ml-o locus were tested at the seedling stage to cultures of the powdery mildew fungus from Europe, Israel, USA. Canada, and Japan. They were resistant with infection type 0/(4) in all tests. They were also resistant to field populations of the pathogen when scored in disease nurseries at more than 78 locations in 29 countries in Europe, the Near East, North and South America. New Zealand, and Japan. This indicates that the 11 genes confer the same, world-wide spectrum of powdery mildew resistance. They have no effect on several other barley diseases such as stripe rust and leaf rust.Part of the research reported here was carried out under IAEA Research Agreement No 1043 and Research Contract No 139-74-1 BIO DK with the European Atomic Energy Community.