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.     

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.     

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.     

Reaction of European spring barley varieties to a population of the net blotch fungus

About 320 European spring barley varieties and about 40 spring barley lines were tested in the field for reaction to a local population of the spot form of the net blotch fungus Pyrenophora teres forma maculata with the imperfect stage Drechslera teres. The European barley varieties showed a wide range of disease reactions, from nearly resistant to very susceptible. At least three groups of genetically related barley varieties could be distinguished: (0)‘Agneta’ and ‘Clermont’, (2)‘Nordal’ and ‘Arve’, and (3)‘Tellus’,‘Pamina’, ‘Albert’ and ‘Birka’. The parentages of these three groups of barley are mutually distantly related and may thus constitute three different sources of resistance to the pathogen population used. The range of disease reaction is similar in varieties from different European countries but it differs among different breeding stations. The relatively high level of resistance is fairly uniformly distributed in contemporary barley varieties all over most of Western Europe. Over time, from about 1830‐1982, the average level of resistance varied around 4.0 (scale value) (from 2.8 to 4.8). However, since about 1940 the variation in disease reaction has become much wider (from 108 to 6.9) for unknown reason(s).     

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.     

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.     

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.     

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.     

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.     

QTL mapping of net blotch resistance genes in a doubled-haploid population of six-rowed barley

Net blotch, caused by Pyrenophora teres f. teres, is a damaging foliar disease of barley worldwide. It is important to identify resistance germplasm and study their genetics. 'Chevron', a six-rowed barley used as a parent for the production of a doubled haploid (DH) population for mapping of Fusarium head blight (FHB) resistance, was also found to be resistant to net blotch. To map the resistance genes, the population was evaluated for resistance at the seedling stage in a greenhouse. The resistance data showed a two-peak distribution. Through linkage mapping, one resistance gene, tentatively called Rpt, was located on chromosome 6HS flanked by Xksua3b-Xwg719d, which was also detected by QTL mapping. This QTL explained 64% of the phenotypic variance for the resistance in this DH population. In addition, a minor QTL was found on chromosome 2HS defined by Xcdo786-Xabc156a. 'Chevron' and 'Stander' contributed the resistant alleles of Rpt and the 2HS QTL, respectively. Both QTLs together explained nearly 70% of the phenotypic variance. The markers for these QTLs are useful for marker-assisted selection of net blotch resistance in barley breeding.     

Salinity tolerance of wild barley Hordeum vulgare ssp. spontaneum

Salinity tolerance of 47 wild barley genotypes and six barley cultivars was evaluated under control and salinity stress (300 mM NaCl) conditions. Shoot and root dry weight (DW), plant height, membrane stability index (MSI), relative water content, survival rate, leaf malondialdehyde (MDA) and proline contents, root and leaf Na, K, Ca and K/Na ratio, and chlorophyll a fluorescence were measured. Salinity stress caused significant increase in the MDA, proline content, Na and Ca concentrations of the roots and leaves, but resulted in a decrease in the other traits. H. spontaneum genotypes were considerably less affected by the salinity than the genotypes of H. vulgare. Plant survivability was negatively correlated with the Na concentration (r =−.66) but positively correlated with the leaf K/Na ratio (r = .67) and MSI (r = .68). Tolerance mechanisms such as ion exclusion (Na) were likely to be present in the wild barley causing K/Na homeostasis as well as the much lower root and shoot Na, resulting in the higher survival rate.     

Evaluation of common wheat cultivars for tan spot resistance and chromosomal location of a resistance gene in the cultivar 'Salamouni'

A total of 50 wheat (Triticum aestivum L.) cultivars were evaluated for resistance to tan spot, using Pyrenophora tritici‐repentis race 1 and race 5 isolates. The cultivars ‘Salamouni’, ‘Red Chief’, ‘Dashen’, ‘Empire’ and ‘Armada’ were resistant to isolate ASC1a (race 1), whereas 76% of the cultivars were susceptible. Chi‐squared analysis of the F₂ segregation data of hybrids between 20 monosomic lines of the wheat cultivar ‘Chinese Spring’ and the resistant cultivar ‘Salamouni’ revealed that tan spot resistance in ‘Salamouni’ was controlled by a single recessive gene located on chromosome 3A. This gene is designated tsn4. The resistant cultivars identified in this study are recommended for use in breeding programmes to improve tan spot resistance in common wheat.     

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.     

Screening for resistance in wild Lycopersicon species to Fusarium oxysporum f.sp. lycopersici race 1 and race 2

Wild Lycopersicon accessions were screened for resistance to the Fusarium wilt disease caused by Fusarium oxysporum f.sp. lycopersici (Fol) race 1 and race 2. In total, four isolates of each race were used. Among 17 accessions of six Lycopersicon species tested, a wide genetic variation for wilt resistance was observed. Most accessions were highly susceptible, some showed intermediate resistance, but one accession of L. cheesmanii (G1.1615 = PI 266375) and two accessions of L. chilense (G1.1556 and G1.1558) were highly resistant to Fol races 1 and 2. The resistance in the latter three accessions equalled or was higher than the resistance determined by the known I-genes, that have been widely used in breeding programmes. These newly found resistant accessions provide breeders with more opportunities for Fusarium disease resistance and may contribute to our understanding of Fusarium disease resistance gene organisation and evolution.     

Screening for resistance against Rhopalosiphum padi (L.). 2. Hordeum species and interspecific hybrids

Summary Twenty-seven accessions of Hordeum species and interspecific hybrids were screened in a growth chamber for resistance against the aphid Rhopalosiphum padi. Resistance measurement was based on the aphids' population growth during a 16-day period. The mean number of nymphs produced per female was only about 14% of that on the control cv. Tellus. H. bogdani was the most resistant accession. As a group, interspecific F₁-hybrids with common barley (H. vulgare × H. sp.) held an intermediate position. Differences in resistance between hybrids as well as between species (H. jubatum and H. lechleri) were observed. The most resistant species were all diploids. No effects of the host plant on formation of alatae or on the within-plant distribution of aphids were found.     

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.     

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.