Evaluating resistance of barley breeding lines to scald in field plots

Summary A 0–4 scoring system to quantify scald (Rhynchosporium secalis) infection is suggested. Scores 1, 2, 3 and 4 allocated to represent 1/4, 1/2, 3/4 and 4/4 of the crop canopy scalded are easy to comprehend and intermediate scores e.g. 0.5, 1.5, 2.5 and 3.5 give it the breadth of a quantitative scale. Scores on a large number of lines showed a high degree of repeatability and were found to be highly correlated with the log transformed values of the actual leaf area damage. Although it was suggested that predictions of leaf area damage at scores 3–4 should be applied with caution, broad generalization of the scores in discriminating the amount of disease were shown to be soundly based and offered plant breeders a tool to standardize the evaluation of scald resistance in field plots on a large scale with this quick and reliable scoring system.     

Genes for resistance in barley to Finnish isolates of Rhynchosporium secalis

Summary Twenty Finnish isolates of Rhynchosporium secalis (Oud.) J.J. Davis, the causal agent of scald, were taken from infected barley (Hordeum vulgare L.) plants and inoculated on to seedlings of a differential series of barley containing a range of major genes for resistance to the fungus, as well as on to six Nordic 6-row spring barleys and three winter ryes (Secale cereale L.). These fungal isolates derived from four sites and three host varieties. Disease development was monitored on two leaves of seedlings in the greenhouse employing a standard scale, and on adult plants in the field by assessing the diseased area on the three uppermost leaves. A comparison was also made between the pathogenicity and virulence of ten Finnish and ten Canadian R. secalis isolates. The Finnish isolates varied in virulence, but with the exception of Algerian (CI 1179) seedlings and adult La Mesita (CI 7565) all seedlings and adult plants of the entire differential series were resistant to all isolates. Canadian isolates were, on average, less virulent than Finnish isolates. All the Nordic checks were susceptible to all Finnish and seven Canadian isolates, but differences in the degree of susceptibility were evident. Isolates of R. secalis from barley were non-pathogenic on rye, isolates from Elymus repens L. were non-pathogenic on barley and rye, and isolates from rye were only pathogenic on rye. Finnish R. secalis isolates contain no redundant pathogenic diversity. The differential series represents a useful, but as yet untapped, source of resistance to R. secalis for Finnish barley breeders.     

The genetic basis of scald resistance in western Canadian barley cultivars

Summary The genetic basis of resistance to scald (Rhynchosporium secalis) within barley breeding populations is poorly understood. The design of effective genetically based resistance strategies is predicated on knowledge of the identity of the resistance genes carried by potential parents. The resistance exhibited by a broad selection of western Canadian barley lines was investigated by evaluating their reactions to five R. secalis isolates. Results were compared to the resistance exhibited by previously characterized lines. This comparison, combined with pedigree analysis indicated that there are two different resistance genes present inwwestern Canadian cultivars. These genes were shown to be independent through analysis of a segregating population derived from a cross between Falcon and CDC Silky. This evidence, along with observed linkage of the gene in CDC Silky with an allele specific amplicon developed for a Rhynchosporium secalis resistance locus on chromosome 3, provides evidence that the gene in Falcon is the Rh2 gene derived from Atlas, and the gene (s) in CDC Silky is located within the Rh/Rh3/Rh4 cluster and is similar to the Rh gene in Hudson.     

Development of SCAR markers linked to a scald resistance gene derived from wild barley

The F₂ progeny of a third backcross(BC₃) line, BC line 240, derived from a Turkish accession of wild barley (Hordeum vulgare ssp. spontaneum),segregated for resistance to scald (Rhynchosporium secalis) in a manner indicating the presence of a single dominant resistance gene. Two SCAR marker slinked to this resistance were developed from AFLP markers. Screens of disomic and ditelosomic wheat-barley addition lines with the SCAR markers demonstrated that the scald resistance gene is located in the centromeric region of barley chromosome 3H,a region previously reported to contain a major scald resistance locus, Rrs1. Markers that flank the Rrs1 locus were used to screen the wild barley-derivedBC₃F₂ population. These markers also flank the wild barley-derived scald resistance, indicating that it maps to the same locus as Rrs1; it may beallelic, or a separate gene within a complex locus. However, BC line 240 does not respond to treatment with the Rhynchosporium secalis avirulence factorNIP1 in the same way as the Rrs1-carrying cultivar Atlas46. This suggests that the scald resistance gene derived from wild barley confers a different specificity of response to theRrs1 allele in Atlas46.In order to increase the durability of scald resistance in the field, we suggest that at least two scald resistances should be combined into barley cultivars before release. The scald resistance gene described here will be of value in the Australian environment, and the several markers linked to it will facilitate pyramiding. This revised version was published online in July 2006 with corrections to the Cover Date.     

Race differentiation and search for sources of resistance to Rhynchosporium secalis in barley in Italy

Summary Barley in Italy has recently been seriously affected by Rhynchosporium secalis. The pathogenic variation of the fungus was studied and 17 races were differentiated on 13 barley cultivars carrying most of the currently known genes for resistance. RC 1, the most virulent and most frequent race, was virulent on 10 out of the 13 differentials and the remaining races proved to be less virulent variants of RC 1. Atlas (C.I. 4118), Atlas 46 (C.I. 7323) and Osiris (C.I. 1622) were the only three differentials resistant to all the analyzed single-spore isolates.Differential cultivars previously assumed to have identical resistance factors did not react in the same way to all the Italian races, thereby revealing either undisclosed differences in the genes described or the presence of additional unidentified ones.Our findings were compared with previous data about virulence of scald populations from different countries, on the basis of tests with common differentials: fundamental differences were found between the Italian population and those of other countries with regard to virulence patterns.The susceptible reactions to race RC 1 of most barley cultivars grown in Italy indicate the urgent need for resistance genes to be incorporated in the cultivated material. Seventy-one barley accessions, known as sources of resistance in different parts of the world, were screened for their behaviour to races RC 1 and RC 13. Twenty-two appeared resistant to both of them.     

Genes for scald resistance from wild barley (Hordeum vulgare ssp spontaneum) and their linkage to isozyme markers

Summary Accessions of Hordeum vulgare ssp. spontaneum, the wild progenitor of barley, collected in Israel (70), Iran (15) and Turkey (6) were screened for seedling response to four isolates of Rhynchosporium secalis, the pathogen causing leaf scald in barley. Resistance was very common in the collection (77%) particularly among accessions from the more mesic sites (90%). The genetics of this resistance were investigated in fifteen backcross (BC₃) lines that contained an isozyme variant from H.v. ssp. spontaneum in a H.v. ssp. vulgare (cv. Clipper) background and were resistant to scald. Segregation in the BC₃F₂ families conformed with a single dominant resistance gene in 9 of the 15 lines. Scald resistance and the isozyme marker were closely linked in three of the BC₃-lines, loosely linked in four and unlinked in the remaining eight. Scald resistance genes were identified on barley chromosomes 1, 3, 4 and 6. Crosses between several of the scald resistant BC-lines together with the linkage data indicated that at least five genetically independent resistances are available for combining together for deployment in barley. The linkage of scald resistance in several BC₃-lines to the isozyme locus Acp2 is of special interest as this locus is highly polymorphic in wild barley.     

Genetic analysis of resistance to barley scald (Rhynchosporium secalis) in the Ethiopian line `Abyssinian' (CI668)

A doubled haploid barley (Hordeum vulgare L.) population from a cross between the cultivar `Ingrid' and the Ethiopian landrace `Abyssinian' was mapped by AFLP, RFLP, SSR and STS markers and tested for resistance to isolates`4004', `2', `16-6', `17', `22' and `WRS 1872' of Rhynchosporium secalis (Oudem.) J.J. Davis, the causal agent of leaf scald. Resistance tests were conducted on parents, DH-lines, a near-isogenic line of `Abyssinian' (NIL) into `Ingrid', and an F₂ population descended from the same F₁ plants as the DHs. The DH population segregated for at least two major R. secalis resistance QTL. All isolates tested identified a major QTL on chromosome 3 (3H) associated with R. secalis resistance, in a 4 cM support interval between the co-segregating markers Bmac0209/Falc666 and MWG680. The QTL was linked with the markers Falc666 (2.3 cM), YLM/ylp (0.3 cM), MWG680 (1.7 cM), cttaca2 (2.5 cM) and agtc17 (9.8 cM). The second QTL was located on chromosome 1 (7H).However, this QTL was only detected by one isolate and was located in an interval of 16 cM in the distal part of the chromosome. At this QTL the allele for improved scald resistance originated from the parent `Ingrid'. There were a number of minor QTL on chromosomes 2 (2H), 4 (4H) and 6 (6H) that were not repeatable either across replications or analysis methods. The importance of checking QTL-models by cross-validation is stressed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Pathogenic variation among isolates of Rhynchosporium secalis from cultivated barley growing in Victoria,Australia

Summary Three hundred and nineteen Rhynchosporium secalis isolates from cultivated barley were divided into five groups on the basis of their virulence on 15 differential barley varieties. Pathogenic variation was also demonstrated for isolates from different scald lesions within the same crop and amongst different spores from the same lesions.     

Effectiveness of selection for plant characters of barley in simulated segregating rows

Summary Single plants of 12 barley (Hordeum vulgare L. and H. distichum L.) cultivars were randomized to form simulated segregating (mixed) rows. There were 16 mixed rows 15 cm apart, each row containing 24 plants 5 cm apart. In addition, every third row was a control row of one of the cultivars, Bonanza. Each plant was harvested separately.The cultivars differed significantly for all seven characters measured. These seven characters fell into 3 performance groups. Group I characters, seeds/head, 1000 kernel weight, days to ripe and height, showed a higher variance in the mixed rows than in the control rows, had high average heritabilities, and were consistent in performance from one row to another when measured by correlation amongst the cultivars. In Group II, yield/plant and seeds/plant, variance, heritabilities and performance consistency were all low or non-significant. Group III, heads/plant, fell between Groups I and II. These results suggested that single plants with the desired genotype could be successfully selected for characters in Group I only.The frequency with which the most desirable genotype would actually have been chosen with 12.5% selection intensity generally confirmed the above conclusions but indicated that the degree of differences among genotypes was also important in selecting the most desirable genotypes.Contribution No. NRG 77–9 from the Beaverlodge Research Station.     

Pathogenic variation among isolates of Rhynchosporium secalis from barley grass growing in South Eastern Australia

Summary The pathogenicity of 182 single spore isolates of Rhynchosporium secalis from Hordeum leporinum and 94 isolates from H. vulgare collected from throughout southeastern Australia was tested on 15 barley varieties, each having different combinations of resistance genes. Forty five percent of the barley grass isolates were pathogenic on 5 or more varieties but only 6% of the cultivated barley isolates attacked this range of varieties. On the basis of reaction type 20 different pathogenicity groups were recognised, with barley grass isolates being classified into 19 and the cultivated barley isolates into 5, four of which were the same as the barley grass isolates. Numerical analysis of data on leaf area damage inferred 33 groups, 24 of which were unique to barley grass isolates, two to cultivated barley isolates and 9 common to both groups. There was as much variation in pathogenicity among single spore isolates from the same lesion as between isolates from different lesions collected from the same or different locations.     

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 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.     

Variation in Ethiopian barley landrace populations for resistance to barley leaf scald and netblotch

One-hundred and eighty landrace populations and six-hundred single-head plants selected from 60 promising populations were evaluated for resistance to scald and netblotch at three locations in Ethiopia. Each accession was tested with and without the application of 50% of the recommended rate of fertilizer at planting. Plants were rated for disease attack two to four times during the season. Both diseases were enhanced by the application of fertilizer and were more severe at the testing sites of Holetta and Bekoji than at Sheno. The difference in disease resistance among and within populations was considerable. Moreover, populations from Arsi and Bale tend to be more susceptible to scald but more resistant to netblotch than populations from other regions. Populations collected from higher altitudes were more resistant to scald, but susceptible to netblotch, than were populations from lower altitudes. The paper illustrates approaches to the identification of valuable genotypes from landrace populations that can be incorporated into a breeding programme for the development of improved varieties with resistance to the principal diseases of barley in Ethiopia.     

The transfer of a gene conferring resistance to scald (Rhynchosporium secalis) from Hordeum bulbosum into H. vulgare chromosome 4HS

Scald is a serious foliar disease that infects barley (Hordeum vulgare L.) causing reduced yields and adversely affecting quality. A means to combat the disease is to breed cultivars that possess genetic resistance. However, all known resistance alleles have so far originated from within the primary genepool of barley. This reliance on H. vulgare and H. vulgare ssp. spontaneum as resistance sources may encourage virulent forms of the pathogen to become established. To broaden the genetic base of cultivated barley and provide novel resistances to many diseases we have used a species from the secondary genepool of barley, H. bulbosum, in a resistance‐breeding programme. In this study we describe the development and trialling of a scald‐resistant recombinant line derived from a hybrid between H. vulgare and H. bulbosum. The scald resistance is simply inherited and located on the short arm of barley chromosome 4 (4HS).     

Simultaneous selection of segregating two-row winter barley material in England and Italy

Summary Independent but simultaneous selection by two breeding teams of the same segregating two-row winter barley material at Cambridge, England and Fiorenzuola, Italy was carried out over the years 1978–1982 with the objective of comparing the individual selection of the two breeding teams and assessing the adaptation of the surviving selections. Visual selection in the generations F₃ to F₅ did not result in any clear cut compartmentalisation of the material and both teams were as likely to select from families previously selected only by the other team as from families they had themselves previously selected. Lines with specific adaptation to each site as well as lines well adapted to both sites were identified from yield trials carried out in F₅ and F₆. The former showed only a very small yield advantage over lines with general adaptation to both sites.     

Yield potential and drought tolerance of segregating populations of barley in contrasting environments

Summary Using the traditional approach (selection for grain yield) it has been found that F₃ families derived from F₂'s selected under unfavourable conditions were more vigorous in the early stages of growth, taller, earlier in heading and with larger yields than F₃ families derived from F₂'s selected under favourable conditions. A high and negative correlation coefficient was found between the drought susceptibility index and grain yield at the driest site, whereas at the wettest site the correlation coefficients were lower and in some cases positive, indicating the existence of traits which are desirable under drought and undesirable under favourable conditions, or vice versa.Expected responses to selection for grain yield using different selection criteria indicated that selection under stress conditions is expected to be more efficient than selection under favourable conditions when dry areas is the target environment.Expected responses to selection for grain yield using different selection criteria indicated that selection under stress conditions is expected to be more efficient than selection under favourable conditions when dry areas is the target environment.Part of this work was supported by Opec Fund for International Development.     

Pathogenicity of the Rhynchosporium secalis population in the Western Cape province of South Africa

The virulence spectra of 50 Rhynchosporium secalis isolates from a population in the Western Cape province of South Africa were determined, and 21 races were detected when evaluated against 17 differential cultivars. The virulence spectrum of the R. secalis population shows considerable variation, and carries unnecessary virulence genes which is quite unexpected, since chiefly susceptible barley cultivars are grown in the south Western Cape. The two most prevalent races, namely races 4 and 7 had three and four virulence genes respectively. Both race 4 and 7 were virulent on the most susceptible cultivars, West China, Steudelli, C.I.8618 and C.I.2226. Considering the resistance genes reported for the cultivars Atlas 46, Turk, and C.I.3515 which showed no susceptible cultivar-pathogen interaction, it would appear that the Rh-Rh3-Rh4 complex is primarily involved in conferring resistance to the local R. secalis isolates This revised version was published online in July 2006 with corrections to the Cover Date.     

Accumulating polygenes for partial resistance in barley to barley leaf rust,Puccinia hordei. II. Field evaluation

Summary Eight lines from the cross between Vada and Cebada Capa with long to very long latent periods and four barley cultivars representing the known range of partial resistance to barley leaf rust, caused by Puccinia hordei, were evaluated in the field for partial resistance and in the greenhouse for the latent period (LP) in the young flag leaf.Each of the 12 entries was sown (15-4-1983) on a plot of 1.0 m². There were four replicates. To reduce interplot interference the plots were separated from each other by 4.0 m of spring rye. The number of urediosori per tiller was evaluated at 27-6, 4-7, 12-7 and ten days after heading. The LP was measured on 10 to 15 plants per entry in 1982 and on 10 plants in 1983.The levels of partial resistance varied greatly. The difference in number of sori per tiller between the most susceptible cultivar, Akka, and the most resistant cultivar, Vada, was about 50 times. Between Akka and the most resistant line this was approximately 5000 times. The LP's varied similarly. Vada had a LP 64% longer than that of Akka, the LP of line 26-6-11 was 15% longer. The range of partial resistance has been extended more than twofold.The correlation coefficients between LP and the level of barley leaf rust, expressed in transformed scale units, varied from -0.99 for the first sampling date to -0.97 for the third sampling date. Sampling the same development stage, ten days after heading, did not improve the r-value (r=–0.98). The LP evaluated in the young flag leaf is shown to be a very reliable criterion for partial resistance in the barley-Puccinia hordei pathosystem.Earliness tends to be associated with susceptibility. The correlation of days to heading with LP was 0.63, and with the level of barley leaf rust in the field 0.64.     

Characterization of resistance genes against scald (Rhynchosporium secalis (Oudem.) J.J. Davis) in barley (Hordeum vulgare L.) lines from central Norway, by means of genetic markers and pathotype tests

Rhynchosporium secalis is a serious pathogen of barley (Hordeum vulgare L.) in central Norway. A breeding effort was initiated in 1977 to introduce resistance from different sources into adapted genotypes, and the first cultivar from the program was recently released. However, little is known about the resistance genes introgressed in this cultivar or in advanced breeding lines. An effort was made to address this issue through a set of isolates and available molecular markers. Fourteen breeding lines and their resistance donors were investigated by evaluating their reactions to 11 R. secalis isolates. Bulked segregant analysis was used to identify molecular markers linked to resistance genes in 12 of the breeding lines. The isolates were found to be of less discriminating value than the markers. Useful information has been obtained as to the nature of several of the resistance genes introgressed. Eight of the 12 breeding lines contained introgressed genes that were located at the `complex Rh' locus on chromosome 3H and hence may not easily be pyramided into the same genotype. Previous information about the nature of the resistance in `Jet' is questioned. Neither of the resistance genes Rh or Rh2 seems to have been incorporated into Norwegian breeding material.     

Variation for resistance to Puccinia hordei in Ethiopian barley landraces

Summary Up to 100 single plant derived lines of 18 Ethiopian barley landraces were evaluated for infection type in the seedling and adult plant stage, and for latent period in the adult plant stage only. A low infection type indicates the presence of race-specific resistance genes of the hypersensitive type, while the latent period is the major component of the polygenic, partial resistance.In the seedling stage 1721 of these single plant derived landrace lines were assessed for infection type against two barley leaf rust races. In the adult plant stage 1227 from these 1721 lines were evaluated for infection type against one race. In the seedling stage 2 (against race 1-2-1) and 29 against race A) out of the 1721 lines showed an infection type lower than 6–7 on the 0 to 9 scale. In the adult plant stage none of the 1227 lines had an infection type lower than 6–7 against race 1-2-1.The variation between and within the landraces for latent period in the adult plant stage was large. Some landraces such as landrace 212845 showed a highly significant and longer mean latent period than most other landraces. Virtually all plants in all landraces carry at least some partial resistance.The near-absence of race-specific, major, resistance genes and the high frequency of moderate levels of partial resistance indicates that the durability of leaf rust resistance in Ethiopian barley landraces is due to the latter type of resistance, and that the multiline principle does not operate.