Regional distribution of resistances to powdery mildew in winter and spring barley cultivars grown in the northern part of France

The spatio-temporal distribution of race-specific resistances to powdery mildew was analysed in northern France (the east, the north and the west of Paris). Resistances were identified in 26 winter and six spring barley cultivars. Seedling leaf segments were inoculated with 20 powdery mildew isolates, chosen to identify 14 resistance alleles. As opposed to other European countries, the resistance alleles differed between winter and spring cultivars grown in the three regions. Most of the winter cultivars had no resistance allele, or only the widespread resistance alleles Mlra and/or Mlh, plus Mlg in the west. Mla9 and Mla13 were also present in the north, but at a low frequency. Spring cultivars carried the alleles Mla7, Mla9, Mla12, Mlk, Mlg or MlLa in the east, where a diversification of resistances has occurred since 1987, particularly because of the use of ‘Volga’ (Mla7, Mlk, Mlg and MlLa). In the north and the west, Mla12 dominated after a decrease in the frequency of Mla7, Mla13 has recently been introduced in the north and the west with the cultivar ‘Vodka’.     

Effect of powdery mildew (Erysiphe graminis f.sp. hordei) on photosynthesis and grain yield of partially resistant genotypes of spring barley (Hordeum vulgare L.)

The effect of powdery mildew on the photosynthesis and grain yield of partially resistant spring barley was investigated. The effect of powdery mildew on the CO₂ exchange rate (CER) of inoculated and non-inoculated leaves was measured in several glasshouse experiments. Grain yield reduction by powdery mildew was determined in three field experiments. The CER of the inoculated leaves was reduced by infection but that of the non-inoculated leaves was not significantly different from that of the corresponding leaves of the control plants. The reduction in CER of the different genotypes was closely related to the percentage leaf area infected with powdery mildew. The correlation between degree of mildew infection and grain yield reduction varied between r = 0.84 (significant at P = 0.01) and r = 0.41 (not significant). Differences in greenness of the crop were correlated better with grain yield reduction (r = 0.66, P = 0.01) than powdery-mildew infection itself. Low percentages of powdery-mildew infection did not lead to significant reduction in grain yield. Partial resistance to powdery mildew could thus be a valuable attribute of future barley varieties.     

Powdery mildew resistance in winter barley cultivars

Powdery mildew is the most common disease of barley in the Czech Republic and winter barley plays a crucial role in the winter survival and reproduction of the pathogen. This study was aimed at identifying resistance genes in winter barley cultivars grown in the Czech Republic from 1971 to 2005 by using the method of postulation. Forty-one cultivars and a parental line were tested at the seedling stage for their reaction to 32 selected isolates of Blumeria graminis f. sp. hordei . In total, 26 resistance spectra were detected and the following 18 resistance genes were found: Mla6 , Mla7 , Mla8 , Mla13 , Mla14 , MlaMu2 , MlaNo3 , MlaRu4 , Mlg , Mlh , MlLa , Mlra , Ml ( Bw ), Ml ( Ch ), Ml ( Dr2 ), Ml ( Dt5 ), Ml ( IM9 ) and Ml ( St ). Two cultivars ('Kiruna' and 'Sorna') exhibited heterogeneity for mildew resistance. Another source of Mla13 and a possible centre of origin of Ml ( Bw ) are discussed.     

Powdery mildew resistance in barley landrace material. I. Screening for resistance

A total of 4,681 accessions of Hordeum vulgare landrace material from Ethiopia, East Mediterranean, Near East, Nepal and China were sown in the field and subjected to the natural powdery mildew epidemic in Denmark. Apparently resistant accessions were selected. Selfed progeny from them were retested and reselected in subsequent years at four locations in Denmark. Finally, 16 promising donors of resistance were retained. They were characterized in the field and tested in the seedling stage for reaction to up to 72 different isolates of the powdery mildew fungus. The absence of the corresponding virulences in the Danish airborne powdery mildew population was ascertained in five years. The resistances in the 16 donors are apparently mutually different and from known sources of powdery mildew resistance in barley. This revised version was published online in July 2006 with corrections to the Cover Date.     

Resistance to powdery mildew in selections from Tunisian landraces of barley

Two-hundred and thirty-two accessions of barley landraces collected from Tunisia were screened for resistance to powdery mildew. A number of race-specific genes were detected using the detached leaf technique. Among the 232 accessions tested, 169 were susceptible to powdery mildew, 20 were resistant and 43 showed differential reactions to the three isolates of powdery mildew used. An attempt was made to determine the number of genes, the types of gene, the types of gene action and the gene loci in 20 resistant accessions. Three types of cross were made: (1) the accessions were crossed to the susceptible variety ‘Pallas’, (2) the accessions were crossed with ‘Pallas’ isolines, and (2) accessions with identical powdery mildew reaction patterns were intercrossed. Three isolates of Erysiphe graminis f. sp. hordei were used: Bzm-1, KM 18-75, R13C. A number of different resistance genes were detected among the 19 resistant accessions. Surprisingly, segregation indicating single genes only were detected with the isolates used. Some of these genes could be associated with loci already known. In 19 cases a dominant and in one a recessive mode of inheritance was detected. The recessive gene was not located at the mlo locus. This investigation represents the first systematic study of race-specific genes for powdery mildew resistance in Tunisian landraces. The newly identified sources of resistance may be used in many strategies of breeding for disease resistance.     

The resistance to leaf rust and powdery mildew of recombinant lines of barley (Hordeum vulgare L.) derived from H. vulgare × H. bulbosum crosses

A set of 23 recombinant lines (RLs) of barley ( Hordeum vulgare L.) derived from H. vulgare  ×  H. bulbosum L. crosses was inoculated with barley leaf rust ( Puccinia hordei ) and powdery mildew ( Blumeria graminis f.sp. hordei ) at the seedling stage to identify their levels and mechanisms of resistance. Eight RLs were studied further in glasshouse and field tests. All three barley parents ('Emir', 'Golden Promise' and 'Vada') were highly susceptible to powdery mildew and leaf rust isolates. Several RLs showed partial resistance expressed as high relative latency periods and low relative infection frequencies against leaf rust. This high level of partial resistance was due to a very high level of early aborting colonies without host cell necrosis. Several RLs showed hypersensitive resistance to some or all isolates. For powdery mildew, one RL was completely resistant to the CC1 isolate and had a hypersensitive resistance to the CO-02 isolate. Three RLs derived from 'Emir' were completely resistant to both powdery mildew isolates, and three more RLs tested in the field had higher levels of partial resistance than their parents. The results indicate that H. bulbosum contains major and minor gene(s) for resistance to leaf rust and powdery mildew that can be transferred to cultivated barley.     

Powdery mildew resistance in Czech and Slovak barley cultivars

Fifteen powdery mildew resistance genes and the gene MlaN81 derived from ‘Nepal 81’were found in 76 Czech and Slovak spring and winter barley cultivars when tested for reaction to a set of powdery mildew isolates. Nine cultivars (‘Donum’, ‘Expres’, ‘Jubilant’, ‘Orbit’, ‘Primus’, ‘Progres’, ‘Stabil’, ‘Vladan’ and ‘Zlatan’) are composed of lines with different resistance genes. The Mlat gene is present in nine cultivars and was transferred from the Anatolian landrace ‘A‐516′. The resistances derived from ‘KM‐1192’and ‘CI 7672’were identical and designated Ml(Kr). Five winter barley cultivars possess the Ml(Bw) resistance. The winter barley line ‘KM‐2099’carries the mlo gene. The parental cultivar ‘Palestine 10’was also tested in which the genes Mlk1, MlLa were identified. The German cultivar ‘Salome’, a parent of seven cultivars tested, probably carries the gene MlLa in addition to mlo and Mla7. The gene mlo6 may be present in the cultivar ‘Heris’. Most of the results were confirmed by the pedigrees of the cultivars.     

Genetics of multiple disease resistance in a doubled-haploid population of barley

The barley accession Q21861 possesses resistance to the stem-rust (Puccinia graminis f.sp. tritici), leaf-rust (P. hordei), and powdery-mildew (Blumeria graminis f.sp. hordei) pathogens. An anther-culture-derived doubled-haploid population was produced from F₁ plants from a cross of this accession and the susceptible breeding line SM89010 as a means of rapidly and efficiently determining the genetics of multiple disease resistance. The doubled-haploid population segregated 1:1 (resistant:susceptible) for resistance to the stem rust pathotype QCC indicating the involvement of a single resistance gene, rpg4. Two-gene (3:1) and one-gene (1:1) segregation ratios were observed for resistance to the stem-rust pathotype MCC at low (23–25°c) and high (27–29°C) temperature, respectively. These different segregation patterns were due to a pathotype × temperature interaction exhibited by rpg4 and Rpg1. another stem-rust-resistance gene present in Q21861. One-gene and two-gene segregation ratios were observed in reaction to the leaf rust and powdery mildew pathogens. These data demonstrate the utility of doubled haploid populations for determining the genetics of multiple disease resistance in barley.     

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.     

Field assessment of partial resistance to powdery mildew in spring barley

Summary Partial resistance to powdery mildew in spring barley was evaluated in three plot types: large isolation plots, in 1.4 m² plots in chessboard design with guard plots of spring wheat and in single rows. Percentage leaf area covered by powdery mildew was scored four to six times during the season and partial resistance was characterized by the area under the disease progress curve. Varietal differences were revealed in al three plot designs, differences between the most resistant and susceptible genotypes being of a factor five. Differences between varieties decreased with decreasing plot size. The relationship between single scores of amount of powdery mildew on the upper four leaves and the area under the disease progress curve was high in all plot designs during the first two to three weeks after heading, allowing selection for the trait by one or two scorings. Differential ranking of varieties between different plot designs was observed, and is assumed to be due to increasing plot interference with reduced plot size and reduced distance between plots. A reliable selection for partial resistance could be made in large isolation plots and in 1.4 m² plots, but hardly in single rows.     

Short Communication Comparison between QTL analysis of powdery mildew resistance in barley based on detached primary leaves and on field data

A quantitative trait loci (QTL) analysis of quantitative powdery mildew resistance was performed on 216 doubled haploid lines derived from a cross between the winter-barley varieties ‘Igri’ and ‘Danilo’ using 67 RFLP loci. Resistance to powdery mildew was determined in the field with natural infection and on detached primary leaves with a specific isolate. The major QTL found in both sets of analysis mapped to the same chromosomal region. No further QTL could be found in the analysis based on detached leaves and one additional minor QTL was found in the analysis based on field data.     

A novel resistance against powdery mildew found in winter barley cultivars

Barley powdery mildew caused by Blumeria graminis f. sp. hordei can be effectively controlled using genetic resistance. Moreover, specific resistances are also important for characterizing cultivars and verifying their origin, purity and authenticity. Winter barley is distinguished by several specific resistances, which are usually absent in spring barley. Besides responses caused by known genes, many cultivars showed a response suggesting the presence of an unknown resistance. Therefore, the aim of this research was firstly, to test winter barley cultivars, suspected to carry an unknown resistance gene, using a large collection of pathogen isolates for their expression of this specific response and to characterise the corresponding resistance. A set of 16 winter barley accessions originating from four gene banks was studied where each accession was represented by five single plant progenies. For resistance tests, 56 isolates of the pathogen were used. A new resistance with a proposed designation of Lu was found in all 16 selected accessions. Apart from Lu, eight well‐known Ml genes (a6, a8, a12, g, h, Lo, ra and Ru2) were postulated. Two accessions of cv. 'Borwina' originating from different gene banks were found to differ in their set of resistance genes.     

Powdery-mildew-resistance genes Mla29 and Mla32 in H. spontaneum derived winter-barley lines

Allelism to the highly polymorphic Mia locus was demonstrated for the powdery-mildew resistance of two Hordeum spontaneum derived winter–barley lines, ‘110-4 × Sonja’ and ‘142–29 × Dura’, by testing the F2 progeny of crosses between these lines and the winter-barley cv. ‘Triton’ (Mlal3) with two appropriate isolates. The results were confirmed by RFLP analysis, using the probe MWG 1H036, which is very closely linked to the Mia locus. The designations Mla29 and Mla32 are proposed for the genes identified in the two lines.     

Hordeum bulbosum-a new source of disease resistance-transfer of resistance to leaf rust and mosaic viruses from H. bulbosum into winter barley

An outline is given of results for the transfer of new resistances against leaf rust and barley mosaic viruses from Hordeum bulbosum into winter barley. Progenies from backcrosses of barley cultivars with H. bulbosum hybrids were tested both in conventional breeding trials and in additional tests under controlled conditions. Resistance to both pathogens proved to be stable and of good heritability, with differences occurring which depended on the combinations used. Lines with resistance to all leaf rust and mosaic viruses tested, or to either one, were selected. Both resistances segregated independently.     

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.     

Conservation of powdery mildew resistance genes in three composite cross populations of barley

Summary Barley powdery mildew was used as a model to evaluate the potential of barley composite cross populations for conservation of disease resistance. The objective was to determine if increases in resistance to powdery mildew could be detected over periods of time in composite cross populations developed in California, where the disease might have had a selective influence on the populations, and the same populations grown in Montana, where no selective influence of powdery mildew was expected. Four isolates of Erysiphe graminis f. sp. hordei were used to monitor the frequencies of plants with specific mildew resistances through early, intermediate and late generations of three composite cross populations (CCII, CCV, CCXII) grown at Davis, California, and Bozeman and Moccasin, Montana. Changes in frequencies of plants resistant to the four isolates were observed between generations in all populations from the three locations. Trends in the frequencies of resistance are discussed in relation to selection pressure applied by E. graminis. It is suggested that associations with gene complexes other than resistance to E. graminis might help to explain the increased resistance observed in these studies.This research was funded in part by U.S. Agency for International Development Contract No. AID/DSAN-C-0024. The authors are grateful to Dr A. L. Kahler for seed of the composite cross populations and to Dr J. G. Moseman for the powdery mildew cultures.Cooperative investigations of the Agricultural Research Service, U.S. Department of Agriculture; the Department of Plant Pathology, Montana State University; and The Montana Agricultural Experiment Station. Journal Series Paper No. 1381 of the Montana Agricultural Experiment Station.     

Identification of growth stage dependent expression of partial resistance of barley to powdery mildew

Summary Genotypic differences for growth stage dependent expression of partial resistance to barley powdery mildew have been identified on the basis of two components of resistance in the glasshouse and the pattern of epidemic development in the field.Differences for infection frequency and proportion of sporulating colonies were highly significant between the 16 genotypes investigated at four stages of plant development in the glasshouse. Both resistance components were significantly correlated mutually (r=0.73 to 0.86) and with the infection level on leaves developed at comparable growth stages in the field (r=0.52 to 0.73). The infection level of seedlings in the glasshouse was significantly correlated with the infection level of the leaves of the first until the third node in the field (r=0.70 to 0.73). Adult plant resistance was predominantly expressed at the uppermost leaf. A clear genotype × growth stage interaction was apparent for infection level; some genotypes showed partial resistance predominantly at the seedling stage and others predominantly at the adult plant stage.Abbreviations IF Infection Frequency, number of colonies per cm² leaf area, glasshouse experiment - IL Infection Level, number of colonies per cm² leaf area, field experiment - PSC Proportion Sporulating Colonies from total number of colonies, glasshouse experiment     

Molecular mapping of major genes and quantitative trait loci determining flowering time in response to photoperiod in barley

Two major genes (eam8 and eam10) and two quantitative trait loci (QTL) determining flowering time in barley were associated with restriction fragment length polymorphism markers. The loci eam8 and eam10 were found to map in regions of chromosomes 1HL and 3HL, respectively, already estimated from previous classical linkage analyses. While investigating doubled haploid lines of a spring habit barley mapping population, two QTL for flowering time were detected on chromosomes 1HL and 7HS, respectively, when the material was grown under long photoperiod conditions. When growing the same lines under short photoperiod, no QTL were discernible. Allelic and homoeologous relationships with flowering time loci described earlier in barley and other Triticeae species are discussed.     

EcoTILLING for the identification of allelic variation in the powdery mildew resistance genes mlo and Mla of barley

In this investigation the successful implementation of a CEL I‐based mutation detection technique for the discovery and detection of DNA polymorphism in the genes mlo and Mla of barley is described. The technique is called EcoTILLING, which is a high‐throughput method to detect and discover new point mutations and small insertions/deletions in DNA. That the method not only reveals polymorphism between different alleles but can also be used as a powerful genetic marker is demonstrated. The genes mlo and Mla are involved in the defence of barley against the fungal pathogen powdery mildew. The powdery mildew resistance gene mlo is a single copy gene, whereas multiple alleles exist at the Mla locus. With EcoTILLING it was possible to identify point mutations and deletions in each of the 11 mlo mutants tested. For Mla 25 natural barley variants were tested, and although the identification was complex due to the presence of highly similar paralogues of Mla, most of the recently identified alleles from Hordeum vulgare ssp. spontaneum were identified. This method offers the possibility to combine different mlo alleles with different Mla alleles from wild barley to obtain cultivars with more durable resistance.