Diallel analysis of net blotch resistance in doubled haploid lines of Nordic spring barleys

A half-diallel was made between five six-rowed Nordic spring barleys to study the genetics of resistance to net blotch. Twenty-five doubled-haploid (DH) lines from each cross and the parents were sown in hill plots in Finland in 1997 and 1998. The plots were artificially inoculated with Pyrenophora teres Drechs. f. teres Smedeg. and assessed for resistance to net blotch. There were statistically significant differences in resistance of the five parents to net blotch. General combining ability (GCA) of the parents and specific combining ability (SCA) effects in the progeny were statistically significant in both years, but GCA effects predominated. Evidence for additive epistasis was minimal. Progeny of a particular cross were less resistant to net blotch than the better parent. The most resistant progeny were derived from the cross between the two most resistant parents, Pohto and WW7977, and resistance was governed by at least eleven effective factors. Narrow sense heritability estimates for resistance to net blotch were high during both years (0.84–0.99). It appears that net blotch resistance of progeny from crosses can be largely predicted from reactions of the parents. Quantitative resistance to net blotch can be further advanced by identification and incorporation of superior parents, from a screening such as reported here, into a recurrent selection breeding programme. This revised version was published online in August 2006 with corrections to the Cover Date.     

Yield of doubled haploid lines of Nordic spring barley infected with net blotch, Pyrenophora teres

Five Nordic spring barley lines (‘Rolfi’, ‘Arve’, ‘Botnia’, ‘Pohto’ and WW7977) and doubled haploid (DH) populations from a half diallel of crosses between them, were sown in the field in Finland over 2 years and were artificially infected with Pyrenophora teres, the causal agent of net blotch. The purpose of the experiments was to determine the extent of yield loss under net blotch infection in a range of parent barleys and DH populations differing in symptom expression. Analysis of foliar damage symptoms, yield and aerial biomass data indicated that, in both years, there were statistically significant differences among parents and crosses, but the relationships between symptom expression and yield maintenance and between symptom expression and aerial biomass maintenance were stronger in 1997, when yields were higher and net blotch was less severe.     

Inheritance of resistance to Pyrenophora teres f. teres in spring barley

The inheritance of seedling resistance to a Swedish isolate of Pyrenophora teres f. teres was investigated in four resistance sources of spring barley. Accessions CI 2330, CI 5791, CI 5822 and CI 9779 were used as resistance sources, and the cultivar ‘Alexis’ was used as a susceptible parent in different crosses. From the disease reaction in the F₁, F₂ and F₃ generations it was concluded that the resistance was governed by the same two complementary genes in CI 5791, CI 822 and CI 9776. One of these genes was present in CI 2330. The first three cultivars were highly resistant to the isolate used in this investigation. These results, when combined with earlier studies, suggest that CI 5791, CI 5822 and CI 9776 may be of great value as sources of resistance to barley net blotch. Spearman's rank correlation between the disease reaction of F₂ plants and their F₃ progeny was highly significant (r = 0.75; P ≥ 0.001) It is suggested that selection in the F₂ generation is effective. In a backcross breeding scheme, single plant reactions in F₁ or F₂ need to be confirmed in later generations.     

Virulence studies of Swedish net blotch isolates (Drechslera teres) and identification of resistant barley lines

Six Swedish and one Canadian single spore isolate of Drechslera teres f. teres were used to screen 109 barley lines for disease resistance and to select a differential set of barley lines for use in assessing pathogen virulence. A large variation for net blotch resistance was found among the 109 barley lines which were classified into four groups, those showing: 1) only resistant reactions; 2) differential reactions; 3) only intermediate reactions and 4) only susceptible reactions. The European commercial varieties included, showed susceptibility to all Swedish isolates, but a few were resistant to the Canadian isolate. The 18- member differential set separated 25 Swedish and two Canadian isolates of D. teres into 14 pathotypes, three of which made up 59% of the isolates. Only one barley differential (CI 9776) was resistant to all net form isolates. Host selection on the pathogen seems to be present as all six isolates obtained from cv. Golf belonged to the same pathotype and 4 of 5 isolates from cv. Karin shared the same virulence pattern. The net form of net blotch (D. teres f. teres) predominated in the sampled regions and only one of 26 Swedish isolates was of the spot form (D. teres f. maculata). This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

Approaches for field assessment of resistance to leaf pathogens in spring barley varieties

The resistance of spring barley varieties to powdery mildew, leaf rust, leaf scald and net blotch was characterized by using results from inoculated small‐plot nurseries and larger survey plots subject to natural infection. The experiments were conducted in different environments. Both trial types often yielded complementary results with respect to the ranking of varieties suggesting that a recommended variety characterization should include both naturally infected survey‐type trials and nursery trials in which the most relevant pathogen isolates and/or isolate mixtures or populations are used for inoculation. Average and median values of the diseased leaf area of a variety were highly correlated with each other and with the ‘genotype main effect’ determined by joint regression analysis, whereas maximum diseased leaf area was poorly correlated with them. Statistics based on absolute disease severity values were highly correlated with the corresponding statistics derived from relative values. It is suggested that one should use at least two parameters to characterize the disease resistance of a variety, a parameter indicating the overall resistance level and a parameter indicating the potential susceptibility and/or resistance instability of a variety. For practical purposes, the genotype median and maximum, respectively, may represent these, although statistically more appropriate parameters do exist.     

Mapping seedling resistance to net form of net blotch (Pyrenophora teres f. teres) in barley using detached leaf assay

To develop molecular markers against Pyrenophora teres f. teres in barley, a detached leaf assay was conducted on two DH populations with a set of 11 single conidial lines (SCLs). Out of these, three showed different reactions in the DH population ‘Uschi × HHOR3073’ and two in ‘(P x V) × HHOR9484’. For SCL ‘QLB’, a 1r : 1s (χ² = 2.78) segregation was observed in the population ‘Uschi × HHOR3073’. In contrast to this, a continuous variation was observed for the SCL ‘WvB’ and ‘d8_4’ in the DH population ‘Uschi × HHOR3073’ and for ‘AR’ and ‘net1840’ in ‘(P × V) × HHOR9484’. With respect to resistance to the SCL ‘QLB’, a single major gene was located on chromosome 7H, and for resistance against SCL ‘WvB’, two QTLs were detected on chromosome 3H and 7H, and against SCL ‘d8_4’, two loci were mapped on chromosome 3HS in ‘Uschi × HHOR3073’. In the DH population ‘(P x V) × HHOR9484’, one locus conferring resistance to the SCL ‘AR’ was located on chromosome 3H. For resistance to SCL ‘net1840’, two QTLs were mapped on chromosome 4H and 5H.     

Diversity among Finnish net blotch isolates and resistance in barley

Summary Seedlings of a differential barley (Hordeum vulgare L.) series (21 genotypes) and of six check genotypes were used in the greenhouse to assess variation in virulence among 20 single-spore isolates of the net blotch pathogen. Pyrenophora teres Drechs. f.teres Smedeg., collected from various sites in Finland. The experiment was run twice and symptom expression was recorded on the first three leaves. Analysis of second leaf symptom scores from Run 1 indicated differences in virulence between isolates, all of which were pathogenic, and differential resistance among the barleys. The virulence of P. teres isolates appeared to be conditioned by the host barley from which the isolate derived; the average virulence of isolates collected from a susceptible host was greater than that of isolates collected from a less susceptible host. Results from Run 2 were similar regarding resistance within the barleys, but variation in virulence among the P. teres isolates was not consistent with that from Run 1. CI 9819 caries duplicate genes for resistance to P. teres (Rpt1b and Rpt1c), and CI 7548 possesses Rpt3d. Both genotypes were highly resistant to all isolates. The Rpt1a gene of Tifang (CI4407) conferred resistance to all isolates in Run 2, but only to about half of the isolates in Run 1. The checks, including two of the symptomatically most resistant Nordic barley genotypes, were universally susceptible during these stringent tests. No selective pressure has been placed on Finnish isolates of P. teres through previous deployment of major resistance genes, and it is speculated that any variation in virulence among isolates is likely to be due to a combination of evolutionary forces including, natural selection, random genetic drift and gene flow.     

Radiation interception and radiation use efficiency of near-isogenic wheat lines with different height

Resistance to Pyrenophora teres Drechs. f. teres Smedeg., the net blotch pathogen, was studied in six 6-row Nordic spring barleys (Hordeum vulgare L.) in the field and in the greenhouse. The barley genotypes were: Arve, Agneta, Artturi, H6221, Pohto and WW7977. Disease progress was monitored in the field (1994 and 1995) in small artificially infected plots, sown at commercial seeding rates, and in infected hill plots (1994). Areas under the disease progress curves (AUDPC) and apparent infection rates (r) were calculated for the uppermost 3 or 4 leaves. Terminal severities (TS) were also recorded. Infection response of seedlings to a range of P. teres isolates was assessed in the greenhouse using a standard scale. In small plots in the field, Arve and Agneta were very susceptible to P. teres infection, as indicated by large values for AUDPC and TS. H6221 and WW7977 were highly resistant, while Artturi and Pohto were moderately resistant. In hill plots the situation was similar, except that Artturi and Pohto appeared less resistant than in the small plots. The relatively greater resistance of H6221 and WW7977 was reflected in seedling infection responses. According to the results of these experiments, H6221 and WW7977 possess adequate levels of quantitative resistance to P. teres to make them useful parents in future crossing programs aimed at improving net blotch resistance in Finnish spring barleys. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic improvement of agronomic and qualitative traits of spring barley

Five experiments, involving 24 genotypes, were carried out over three seasons to estimate the increase both in grain yield and other selected agronomic traits, and in potential malt extract of spring barley achieved by plant breeders during a period of over 160 years. In all experiments there was neither a chemical control of fungal diseases nor physical support of the tall, older genotypes to prevent yield losses from attack by fungi and lodging, respectively. In all experiments the most modern cultivars yielded more than the older ones. Significant breeding progress was also observed for plant height, resistance score, protein content and potential malt extract. The trend over time was generally of a curvilinear nature, showing no or only a slight increase till the 1950s and/or 1960s, and a greater rate of increase during the last 40 years of barley breeding. Using data from the official trials of cultivars and advanced breeding lines a linear increase in yield per year of 60 kg/ha since the 1950s was calculated for the low‐rainfall conditions of eastern Austria.     

Inheritance of partial resistance to spot blotch in barley

Diallel crosses (without reciprocals) were made among 10 different barley genotypes with genetic variability for spot blotch resistance. Forty‐five F₁ hybrids and their parents were assessed for their combining abilities for the disease resistance. Three experiments, two in a growth chamber on detached leaf and seedlings tests and one in the field on adult plant stages, were undertaken using a randomized complete block design with five replicates. A mixed conidial suspension of nine virulent isolates of the pathogen was used for inoculation. Statistical analysis showed genetic variability for spot blotch resistance. Results showed that the cultivar Banteng, the Ethiopian line CI‐5791 and the Syrian line 79‐SIO‐9 had partial resistance in all experiments. General combining ability was significant, with either positive or negative values. Resistant genotypes show favourable GCA‐effects, and they could therefore be successfully used for breeding purposes.     

Possibility of exploiting the Yd2 resistance to BYDV in spring barley breeding

The effects of the Yd2 gene on tolerance to barley yellow dwarf virus (BYDV) and other agronomically important characters in spring barley were evaluated in a set of randomly selected doubled haploid (DH) lines of an‘Igri’/‘Atlas 68’ cross and three crosses between CIMMYT Yd2 materials and the Czech malting barley ‘Akcent’. The cleaved amplified polymorphic site (CAPS) diagnostic marker Yd2 was used for identification of the Yd2 gene and this analysis showed high agreement with the results of field infection tests. Yd2 lines exhibited significantly lower symptom scores and lower reductions of some grain yield characters, but their resistance level was not consistent over the years. The presence of secondary stresses (high temperature/drought) in 2000 led to relatively higher sensitivity to BYDV infection, strengthened by the long life cycle of genotypes. In cases where secondary stresses were mild (in 2002), the longer life cycle significantly increased sensitivity to BYDV infection only in the absence of the Yd2 gene (in susceptible genotypes). The examination of different vegetative, grain yield and malting quality characters separately for groups of Yd2 and non‐ Yd2 lines did not show any evidence of adverse effect of the Yd2 gene on any character.     

In vitro quantification of barley reaction to leaf stripe

An in vitro technique was used to quantify the infection level of leaf stripe in barley caused by Pyrenophora graminea. This pathogen penetrates rapidly through subcrown internodes during seed germination of susceptible cultivars. Quantification was based on the percentage of the pieces of subcrown internodes that produced fungal hyphae cultured on potato dextrose agar media. The disease severity was evaluated among five cultivars with different infection levels and numerical values for each cultivar were obtained. A significant correlation coefficient (r = 0.91, P < 0.02) was found among the in vitro and field assessments. In addition, the results were highly correlated (r = 0.94, P < 0.01) among the different in vitro experiments, indicating that this testing procedure is reliable. The method presented facilitates a rapid preselection under uniform conditions which is of importance from a breeder's point of view. Significant differences (P < 0.001) were found for the length of subcrown internodes between inoculated and non‐inoculated plants with leaf stripe. Isolate SY3 was the most effective in reducing the subcrown internode length for all genotypes.     

Genetic analysis of the nitrogen and phosphorus utilization efficiencies in mature spring barley plants

Inheritance of the efficiency of nitrogen and phosphorus utilization in grain production was studied in diallel hybrids of spring barley. Effects of varied NPK fertilization and soil moisture on the genetic variance was evaluated at maturity in pot‐ and field‐grown plants. A preponderance of general combining ability effects was found for the efficiency ratios and utilization efficiencies, suggesting that the generative efficiency indices were mainly controlled by an additive genetic system. However, the relatively high narrow‐sense heritabilities (0.33‐0.81) found under optimal conditions tended to decline to 0.00‐0.36 following water and nutrient shortages. The importance of non‐additive gene effects for utilization efficiencies was demonstrated in the field under reduced nutrition. This leads to the conclusion that the involvement of additive gene action should facilitate selection efforts only under favourable environments. Selection delayed until later hybrid generations should be used when trying to improve the nutrient efficiency of barley under less favourable nutrition conditions.     

Generation Mean Analysis of Inheritance of Resistance to Pyrenophora teres in Barley

F1, F2, Fl × parentl (BC1), and Fl × parent2 (BC2) generations resulting from four crosses among seven cultivars of barley used in national and international breeding programmes were tested at the seedling stage for their resistance to a mixture of five isolates of Pyrenophora teres. Four methods were used to assess disease resistance: infection type (IT), average lesion size (ALS), number of lesions per unit leaf area (NL) and percent leaf area infected (PLAI). Gene actions were estimated by generation mean analysis on each of the four crosses and on each of the evaluation methods. Significant additive and additive × additive epistatic effects were found. Infection type and percent leaf area infected were found to be highly correlated in all four crosses. These results suggest that barley breeders could improve the level of resistance to P. teres by making appropriate crosses between highly susceptible barley cultivars.     

Genetic control over grain damage in a spring barley mapping population

A genetic map was constructed using DNA‐based markers in a barley mapping population derived from the cross ‘Tankard’בLivet’, that was developed to explore the genetic control over grain damage in spring barley cultivars. Quantitative trait loci (QTL) were located for husk skinning, gape between the lemma and palea and splitting of the fused pericarp/testa/aleurone tissues. The QTL accounted for 70% of the genetic variation in Split and 60% of the genetic variation in Gape and Skinning. The QTL were clustered on chromosomes 1H, 4H, 5H, 6H and 7H. QTL analysis indicates the possibility of transgressive segregation for grain splitting and so the breeding of lines with more extreme splitting. This is of concern to the malting industry as, without extensive phenotypic assessment, such lines could be commercialized, as was the case of Landlord, and put malting barley supplies at risk. These findings are discussed in relation to the genetic control over traits including grain length and width.     

Genotype × environment interactions in the expression of net blotch resistance in spring and winter barley varieties

The net blotch resistance of barley varieties widely grown in Denmark was studied using data obtained from naturally infected field plots and inoculated disease nurseries in multiple years and locations. Data were analysed by combining two statistical procedures, 1) joint regression (JR) and 2) additive main effect, multiplicative interactive effect analysis (AMMI). Of the total variation for net blotch disease severity, 61–81% could be explained by plant genotype (G) and environment (E) main effects. Of the remaining variation, 77–86% could be explained by significant G × E interactions that were due to genotype sensitivity to the mean environmental disease level and/or to specific reactions of individual genotypes in particular environments. G × E interactions led to a different ranking of varietal performance across environments. There was evidence for G × E interactions caused by the virulence characteristics of the initial inoculum sources. The spring barley varieties Alexis and Bartok and the winter barley varieties Jolante, Ludo, and Rafiki were identified as varieties with high levels of net blotch resistance in all environments. Their performance was little affected by G × E interactions, suggesting that they represent good sources for non-specific resistance. Combining JR and AMMI reduced the dimensionality of complex G × E problems greatly, identified systematic reaction patterns of varieties possibly pertaining to different resistance mechanisms, and described specific and non-specific resistance of varieties by means of few parameters while maintaining the possibility to reconstruct the original data with little loss of information. This is very useful for improving the evaluation of varietal resistance for breeding and disease management purposes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Turkish barley varieties as a source of resistance to net blotch

Summary From the world collection, 875 barley varieties of Turkish origin have been screened for resistance to local races of net blotch caused by Drechslera teres (Sacc.) Shoem. Six of these exhibited a high degree of resistance, which varied in expressivity when tested under a wider range of environmental conditions. In this respect these varieties rank between the consistency of resistance expressed by varieties from Ehtiopia, and the inconsistency with which resistance is expressed by varieties from Manchuria. The implications are discussed.