Inheritance of Partial Resistance to Net Blotch in Barley

Reciprocal crosses were made between 9 different barley genotypes with high genetic variability for net blotch resistance. Parents and 72 F₁ plants were used to determine the inheritance of partial resistance to net blotch. Four experiments, one in a growth chamber on seedlings and 3 others in the field on adult plants, were undertaken using a randomised complete block design. An isolate of net blotch from local cultivars was used for inoculation. Non-inoculated plants of one of the field experiments were used for the detached leaf test in petri dishes. Results show that the cultivars ‘Banteng’, and ‘Arrivate’, the Ethiopian line CI 5791, and the Syrian line 79 SIO-10, had a high partial resistance in all experiments. Diallel analysis showed high significant general and specific combining ability when maternal and reciprocal effects were not significant. As the resistance genotypes have a high additive genetic effect, they could be successfully used for breeding purposes.     

Inheritance of resistance to Rhynchosporium secalis in spring barley (Hordeum vulgare L.)

The inheritance of durable resistance of selected spring barley varieties to Rhynchosporium secalis was investigated. Data from the F₂ generation of a 4 × 4 diallel, without reciprocals and the F₄ generation of three crosses selected out of this diallel, suggest that resistance in this sample of varieties tested is complex in inheritance. Significant additive effects were detected indicating that the resistance level of barley cultivars may be improved by the hybridisation of suitable varieties. However, the genes conferring resistance seem to be concealed by the expression of one completely dominant resistance gene in our set of varieties. These results are partly in conflict with previous results on the inheritance of resistance to R. secalis in the breeder's line ‘11258/2286₁₃A’ indicating that the effectiveness of this resistance gene may be greatly influenced by the genetic background of the current population of R. secalis.     

A rapid technique for determining common root rot reaction in barley

The reaction of two barley cultivars (‘Thibaut’ and ‘Arizona’) and three lines (AECS 76,‘Tadmor’ and WI 2291) to infection by Cochliobolus sativus was evaluated by three methods. Deep seeding of inoculated seeds was performed in pots and field. Gibberellin (GA3) was applied to inoculated seeds in vitro. Susceptibility assessments were made in vitro at 2 weeks, and in pots at 7 weeks after inoculation, using a scale of 1 (very resistant) to 5 (very susceptible) according to the percentage infected area of the subcrown internodes. There were no differences between the results using the three methods.‘Arizona’ was the most susceptible, whereas ‘Thibaut’ and AECS 76 were partially resistant genotypes. The WI 2291 line and ‘Tadmor’ were moderately susceptible and moderately resistant, respectively. In vitro tests yielded the same order as tests from pot and field experiments.     

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.     

Genetic Variability for Grain Yield and Protein Content in Barley and Its Modification by Net Blotch

Protein content, yield and 1000-grain weight of healthy and net blotch-diseased plants were studied in a diallel cross programme concerning nine different varieties of barley (Hordeum vulgare L.). F₁ hybrids and their parents were grown in the field in two randomized block design experiments each with three replications (three inoculated and three non-inoculated plots). General and specific combining abilities were determined for yield, protein percentage, 1000 grain weight, protein yield, and non-protein yield. A high genetic variability for the traits studied was observed. Non-protein yield and 1000-grain weight decreased significantly in inoculated plants where protein yield was not affected. Significant general and specific combining abilities were observed for all traits. Values for general and specific combining abilities were, in some cases, significantly modified by net blotch of barley. This indicates that attention must be paid to the danger of drawing conclusions in quantitative genetics studies dealing with both diseased and healthy plants.     

In vitro quantification of the reaction of barley to common root rot

An in vitro technique was used to quantify the infection level of common root rot. This disease produces a brown to black discoloration of the subcrown internodes of barley. Quantification was based on the percentage of germinated infected pieces (1.5 mm) of subcrown internodes cultured on potato dextrose agar media. The disease severity was apparent among four different visually classified categories and numerical values for each category were applied. The results were highly correlated (r = 0.97, P < 0.01) among the different in vitro experiments, indicating that this testing procedure is repeatable. Highly significant differences (P < 0.001) were found for the length of first leaf and fresh weight between plants inoculated and uninoculated with common root rot. However, the effect of inoculation on fresh weight only differed significantly (P < 0.02) among the genotypes.     

Barley mutants with enhanced level of resistance to Swedish isolates of Bipolaris sorokiniana,casual agent of spot blotch

Spot blotch, caused by Bipolaris sorokiniana, has become an increasingly important disease on barley in Sweden. In order to enhance our knowledge of this disease, eight Swedish B. sorokiniana isolates were assessed for their virulence pathotype on 30 barley genotypes including eight lines of mutant origin, designated B. sorokiniana tolerant (bst). A leaf screen assay revealed the presence of virulence groups 1 and 7 in Sweden. Several mutant genotypes demonstrated enhanced resistance responses (P = 0.0001), of which bst1:3 had the lowest leaf scoring value to the most virulent isolate, Swe2 (virulence group 7). To evaluate the host root responses, two contrasting isolates Swe2 and Swe4 (virulence group 7 and 1) were used to inoculate a set of eight differential lines, including four promising mutant lines from the leaf screen. Mutant bst1:6 was significantly less susceptible (P = 0.001) compared with the other barley genotypes tested, independent of the two isolates used. Mutants bst1, bst1:3 and bst1:4, which displayed less or no spots, are also considered promising genotypes for further breeding attempts.     

Genetic analysis of resistance in barley to barley yellow dwarf virus

The inheritance of resistance to barley yellow dwarf virus (BYDV) was studied in the selected 24 spring and winter barley cultivars that showed a high or intermediate resistance level in 1994‐97 field infection tests. The polymerase chain reaction diagnostic markers YLM and Ylp were used to identify the resistance gene Yd2. The presence of the Yd2 gene was detected with both markers in all the resistant spring barley cultivars and lines from the CIMMYT/ICARDA BYDV nurseries. The results of field tests and genetic analyses in winter barley corresponded with marker analyses only when the Ylp marker was used. Genes non‐allelic with Yd2 were detected by genetic analyses and the Ylp marker in moderately resistant spring barley cultivars ‘Malvaz’, ‘Atribut’ and ‘Madras’, and in the winter barley cultivars ‘Perry’ and ‘Sigra’. Significant levels of resistance to BYDV were obtained by combining the resistance gene Yd2 with genes detected in moderately resistant cultivars. The utilization of analysed resistance sources in barley breeding is discussed.     

Genome-wide association studies in a barley (Hordeum vulgare) diversity set reveal a limited number of loci for resistance to spot blotch (Bipolaris sorokiniana)

Spot blotch caused by Bipolaris sorokiniana is an important disease in barley worldwide, causing considerable yield losses and reduced grain quality. In order to identify QTL conferring resistance to spot blotch, a highly diverse worldwide barley set comprising 449 accessions was phenotyped for seedling resistance with three isolates (No 31, SH 15 and SB 61) and for adult plant resistance at two locations (Russia and Australia) in two years. Genotyping with the 50 k iSelect barley SNP genotyping chip yielded 33,818 informative markers. Genome-wide association studies (GWAS) using a compressed mixed linear model, including population structure and kinship, revealed 38 significant marker-trait associations (MTA) for spot blotch resistance. The MTA corresponded to two major QTL on chromosomes 1H and 7H and a putative new minor QTL on chromosome 7H explaining between 2.79% and 13.67% of the phenotypic variance. A total of 10 and 14 high-confidence genes were identified in the respective major QTL regions, seven of which have a predicted involvement in pathogen recognition or defence.     

Genetic Analysis for Partial Resistance to an Iranian Strain of Bacterial Leaf Streak (X. campestris pv. hordei) in Barley

Seven barley genotypes with high genetic variability for resistance to bacterial leaf streak (Xanthomonas campestris pv. hordei) were crossed in diallel fashion to determine the inheritance of resistance to this disease. Two experiments were undertaken in a controlled growth chamber using a complete-block design with four replicates. Each replicate consisted of a row of 20 seedlings per parent or F₁ hybrid. An Iranian strain of bacterial leaf streak was used for inoculation of 12-day-old seedlings. Results showed that the cultivars ‘Express’ and ‘Morex’ and the Iranian pure line Iran-3a had a high partial resistance in booth experiments. Diallel analysis showed highly significant general and specific combining abilities. ‘Morex’ and Iran-3a were the best combiners for partial resistance to bacterial leaf streak in barley and could be successfully used for breeding purposes.     

Inheritance of tolerance to zinc deficiency in barley

Barley (Hordeum vulgare L.) is often grown on alkaline zinc (Zn)‐deficient soils where reductions in yield and grain quality are frequently reported. Currently, the use of Zn‐based fertilizer along with Zn‐deficiency‐tolerant genotypes is considered the most thorough approach for cropping the Zn‐deficient soils; however, developing or breeding genotypes with higher Zn efficiency requires a good understanding of the inheritance of tolerance to Zn deficiency. This study was conducted to determine genetic control of this trait in barley. Two parental cultivars ('Skiff, moderately tolerant; and ‘Forrest’, sensitive), 185 F₂ plants, and 48 F₂‐derived F₃ families from this cross were screened to determine inheritance of tolerance to Zn deficiency using a visual score of deficiency symptoms. The segregation ratios observed indicated that greater tolerance to Zn deficiency in ‘Skiff compared with ‘Forrest’ at the seedling stage is controlled by a single gene with no dominance. The results also indicate that visual scores are useful for genetic analysis of tolerance to Zn deficiency.     

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).     

Characterization of greenbug resistance in barley

The greenbug [Schizaphis graminum (Rondani)] is an extremely damaging pest of barley (Hordeum vulgare L.), particularly in the southern Great Plains of the USA. Winter barley targeted for production in this region should incorporate resistance to greenbug in the form of the resistance gene Rsg1a (in ‘Post 90′) or Rsg2b (in PI 426756). This study was conducted to fully characterize the resistance profile of these two genes against important greenbug biotypes, and to determine which of the two resistance genes is most effective in protecting barley from the greenbug. Eight barley and four wheat cultivars and germplasms were challenged with six greenbug biotypes and damage ratings were recorded for each combination. In five of the six tests, ‘Post 90’ was significantly more resistant than PI 426756 to greenbug feeding damage. Based on the results presented here, we conclude that the resistance gene, Rsg1a, in ‘Post 90’ is the better choice for use in breeding programmes and will provide better protection than Rsg2b against the greenbug.     

Genetic analysis of some chlorophyll fluorescence and productivity parameters in barley (Hordeum vulgare)

Reciprocal crosses were made between five barley genotypes of diverse origin. Parents and 20 F₁S were grown in a greenhouse in a randomized block design with three replications. Each replication consisted of one pot containing three plants. Chlorophyll fluorescence activity of the penultimate leaf (below the flag leaf) of the main stem in each plant was measured at two stages of development (six-leaf stage and anthesis). At anthesis, and after measuring fluorescence parameters, a drought stress was applied. Yield components were measured at complete maturity. Statistical analysis for chlorophyll fluorescence parameters showed high genetic variability for the traits studied at both the six-leaf stage and anthesis, but most fluorescence parameters had decreased by anthesis. General and specific combining abilities of a diallel analysis were significant for most of the characters studied. General combining ability (GCA) values were usually more important than those of specific combining ability (SCA), showing the importance of additive genetic control for the fluorescence traits. Reciprocal effects were not significant for the fluorescence traits studied. A high genetic variability was also observed for yield components. Additive gene effects for plant weight, spike number, spike weight, number of grains per plant and 1000-grain weight were demonstrated. Potential photosynthetic activity (Rfd) at anthesis was positively correlated with plant weight, spike number per plant and 1000-grain weight.     

Postulation of leaf-rust resistance genes in Czech and Slovak barley cultivars and breeding lines

Leaf‐rust resistance (Rph) genes in 61 Czech and Slovak barley cultivars and 32 breeding lines from registration trials of the Czech Republic were postulated based on their reaction to 12 isolates of Puccinia hordei with different combinations of virulence genes. Five known Rph genes (Rph2, Rph3, Rph4, Rph7, and Rph12) and one unknown Rph gene were postulated to be present in this germplasm. To corroborate this result, the pedigree of the barley accessions was analysed. Gene Rph2, as well as Rph4, originated from old European cultivars. The donor of Rph3, which has been mainly used by Czech and Slovak breeders, is ‘Ribari’ (‘Baladi 16’). Rph12 originates from barley cultivars developed in the former East Germany. Rph7 in the registered cultivar ‘Heris’ originates from ‘Forrajera’. A combination of two genes was found in 10 cultivars. Nine heterogeneous cultivars were identified; they were composed of one component with an identified Rph gene and a second component without any resistance gene. No gene for leaf rust resistance was found in 17 of the accessions tested. This study demonstrates the utility of using selected pathotypes of P. hordei for postulating Rph genes in barley.     

Molecular mapping of greenbug (Schizaphis graminum) resistance gene Rsg1 in barley

The greenbug, Schizaphis graminum (Rondani) is an extremely damaging aphid pest of barley (Hordeum vulgare L.) particularly in the southern Great Plains of the USA. The simply inherited, dominant resistance gene Rsg1 is in all greenbug‐resistant US barley cultivars. In this study, we conducted molecular mapping of Rsg1 using an F_2:3 population derived from a cross between the greenbug‐resistant Post 90*4/R015 and susceptible CI2260 inbred lines. Segregation of host responses to greenbug biotype E infestation confirmed that a single dominant gene is responsible for greenbug resistance in Post 90*4/R015. Simple sequence repeat (SSR) markers evenly distributed along the seven barley chromosomes were employed for the construction of a framework genetic map. Linkage analysis placed the Rsg1 locus in the long arm of chromosome 3H (3HL) flanked by SSR markers Bmag0877 and GBM1420 that were 35 cM apart. Polymorphic single‐nucleotide polymorphism (SNP) markers in 3HL were identified from an Illumina GoldenGate SNP assay and used for targeted mapping to locate Rsg1 to an 8.4‐cM interval. Comparative analysis identified syntenic genomic regions in Brachypodium distachyon chromosome 2, in which 37 putative genes were annotated including a NB‐LRR‐type resistance gene homologue that may be a potential candidate gene for the Rsg1 locus of barley. Results from this study offer a starting point for fine mapping and cloning of this aphid resistance gene in barley.     

Inheritance of resistance to spot blotch caused by Bipolaris sorokiniana in spring wheat

Three F₁ progenies and their families in the segregating generations (F₃, F₄, F₅ and F₆), obtained after crossing resistant × susceptible wheat genotypes were studied in the field to determine the genetics of resistance to spot blotch caused by Bipolaris sorokiniana. Spot blotch scores in the F₁ generation showed absence of dominance. Individually threshed F₂ plants were used to advance the generations. Progenies (200‐250) of resistant genotypes Acc. No. 8226, Mon/Ald, Suzhoe#8 crossed with susceptible ‘Sonalika’ were evaluated in the F₃, F₄, F₅ and F₆ generations under induced epiphytotic conditions. Based on disease score distribution in individual progeny rows, F₃ progenies were grouped into four classes: homozygous resistant, homozygous susceptible, segregating resistant and segregating susceptible. Resistance appeared to be under the control of three additive genes. The presence of three genes was also noted in the distribution of F₄ and F₅ lines. In the case of F₆ progeny rows, both quantitative and qualitative models were used to estimate the number of segregating genes based on a 2‐year trial. It appeared that resistance to spot blotch was controlled by the additive interaction of more than two genes, possibly only three.     

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.     

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.