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.     

Tolerance of barley seed germination to cold- and drought-stress expressed as seed vigour

In barley, high seed vigour is a precondition for rapid and homogenous field emergence and good malting quality. Seed vigour was defined as germination percentage under stress conditions (10°C, drought stress ‐ 2 Bars) in 7‐8 barley varieties grown in 7‐8 locations in the Czech Republic over 7 years. Three of the 7 years were not suitable for high seed quality, probably because of unsuitable weather, as average seed vigour reached only 61, 77 and 86%, respectively. In the remaining 4 years, the average vigour exceeded 94%. The impact of variety on seed vigour was higher in the ‘bad years’ and the impact of location was higher in the other years. Varieties with higher vigour from all locations in the ‘bad years’ were identified. Lower vigour was related to the high occurrence of fungi (indicated by ergosterol assays) and to lower field emergence rates of seed samples. The results support the possibility of selecting for improvement of barley seed vigour, which is related to tolerance to various conditions during emergence and to homogenous malting.     

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.     

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.     

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.     

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).     

Mapping quantitative trait loci for salt tolerance at germination and the seedling stage in barley (Hordeum vulgare L.)

Quantitative trait loci (QTLs) controlling salt tolerance at germination and the seedling stage in barley (Hordeum vulgare L.) were identified by interval mapping analysis using marker information from two doubled haploid (DH) populations derived from the crosses, Steptoe/Morex and Harrington/TR306. Interval mapping analysis revealed that the QTLs for salt tolerance at germination in the DH lines of Steptoe/Morex were located on chromosomes 4 (4H), 6(6H), and 7(5H), and in the DH lines of Harrington/TR306 on chromosomes 5(1H) and 7(5H). In both DH populations, the most effective QTLs were found at different loci on chromosome 7(5H). Genetic linkage between salt tolerance at germination and abscisic acid (ABA) response was found from QTL mapping. The QTLs for the most effective ABA response at germination were located very close to those for salt tolerance on chromosome 7 (5H) in both crosses. The QTLs for salt tolerance at the seedling stage were located on chromosomes 2(2H), 5(1H), 6(6H), and 7(5H) in the DH lines of Steptoe/Morex, and on chromosome 7(5H) in the DH lines of Harrington/TR 306. Their positions were different from those of QTLs controlling salt tolerance at germination, indicating that salt tolerance at germination and at the seedling stage were controlled by different loci. This revised version was published online in July 2006 with corrections to the Cover Date.     

A study of grain dormancy and viability in spring barley

In field experiments carried out between 1989 and 1991, spring barley was sampled to observe changes in the grain viability and dormancy during grain filling. The weather during the grain filling periods of 1989 and 1990 was unusually warm and dry, and the crop in 1989 was affected by drought. Some grains were capable of germination shortly after anthesis, but were dormant. Dormancy declined with increasing maturity, but was prolonged by low temperatures and reduced by warm weather. Differences were found in the dormancy characteristics of the two cultivars tested. Apart from grains at the extremities of the ear, which were sometimes found to be small and with low viability and greater dormancy, though not significantly so, no consistent differences were found between the viability and dormancy of grains in different positions in the ear.     

Inheritance of nitrogen and phosphorus utilization efficiency in spring barley at the vegetative growth stages under high and low nutrition

Inheritance of the nitrogen and phosphorus utilization efficiencies (NUE and PUE, respectively) and that of the tolerance (T) to limited NP nutrition was investigated in spring barley crosses at the vegetative growth stages. Plants were grown in sand-vermiculite cultures under high and low NP nutrition. In a diallel set (Ps and F₂s), both general combining ability (GCA) and specific combining ability (SCA) effects were significant for the variation in NUE and PUE, while the variation in T was mainly associated with GCA effects. The contribution of nonadditive genes for the utilization efficiencies was found to increase under nutrient shortages. Overdominance of genes was detected. The characters exhibited low heritabilities (0.10-0.42). Generation means analysis in two cross-combinations revealed significant effects of epistatic gene interactions. It was assumed that the involvement of both the dominance effects and epistatic interactions would not facilitate selection efforts to improve the characters in spring barley at its vegetative growth. Such selection should be performed among families of later generations. The genotype-nutrition interactions observed suggest that selection under diverse nutrition rates would be necessary for the more precise evaluation of barley efficiency under less favourable soil fertility.     

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 variation for dry matter and nitrogen accumulation and translocation in two-rowed spring barley II. Nitrogen translocation

An understanding of the partition between pre-anthesis and post-anthesis N uptake and their contribution to total grain N and protein content in spring two-rowed barley (Hordeum vulgare spp. distichum L.) is important to achieve additional breeding progress for both fodder and malting barley. N translocation from the vegetative tissues at anthesis to the kernel, N translocation efficiency, and N harvest index (NHI) were studied in field experiments during 4 years (1995–1998). Plants were harvested at anthesis and maturity and divided into leaf+culm, chaff and grain. Significant cultivar differences in N translocation, N translocation efficiency and NHI were determined. Across cultivars, the highest N translocation was in a favorable year (93 kg ha⁻¹) and the lowest in a year with poor growing conditions (40 kg ha⁻¹). Cultivar differences in N translocation were related to dry matter and pre-anthesis N accumulation (R²>0.70). N translocation efficiency varied more among the cultivars (0.27–0.66) than years (0.47–0.52). Post-anthesis N uptake was negatively correlated (P<0.01) with N translocation. NHI ranged among the cultivars from 0.49 to 0.73 and among the years from 0.57 to 0.74. The cultivars Arapiles, Schooner, Cantala, Kaskade and Pek stored in the grain more than 70% and Hiproly less than 50% of above-ground N at maturity. Translocated N participated with 85, 56, 42, and 61% in grain N in 1995, 1996, 1997, and 1998, respectively. The ratio of translocated N to grain N could be an indicator of growing conditions; a higher ratio indicates good growing conditions over the entire growth period, a lower ratio indicates poor conditions during pre-anthesis, and a medium ratio indicates some temperature and water deviations from the long-term average. Straw N concentration was in significant positive (P<0.01) correlation with N translocation and translocation efficiency. Straw N concentration adequately represents N efficiency utilization for synthesis of grain protein, and because it saves time and money compared to N harvest index determination, it can be used for the testing of breeding materials for the development of new barley cultivars.     

Diversity and geographical distribution of seed lipoxygenase‐1 thermostability types in barley

To understand the diversity in the thermostability of the seed lipoxygenase‐1 (LOX‐1), 1040 cultivars of worldwide barley (Hordeum vulgare ssp. vulgare) genetic resources were investigated. The relative thermostability of LOX‐1 (LOX‐RTS) in these lines showed a bimodal frequency distribution and these lines were categorized into the high and low thermostability types (H‐type and L‐type, respectively). The H‐type lines predominated in the wild progenitor, ssp. spontaneum. The geographical distribution of these types in the cultivars was surveyed. The frequencies of the H‐ and L‐types were almost equal to one another in southwestern Asia. The occurrence of the H‐type predominated in eastern Asia and Africa, whereas in Europe and Turkey, the L‐type did. The predominance of the L‐type in Europe and Turkey can be understood through the hypothesis that the Fertile Crescent domestication contributed the majority of diversity in Europe. The uneven geographical distribution of the LOX‐1 thermostability types in the cultivars may reflect a polyphyletic origin of barley.     

Genetic analysis of tissue culture traits in barley cv. 'Lenins'

The barley cultivar ‘Lenins’ was found to be a genotype showing high shoot regeneration ability in cultures derived from immature embryos. Five cultivars different from ‘Lenins’ in shoot regeneration ability were reciprocally crossed with ‘Lenins’ and the inheritance of tissue culture traits was investigated. F₂ plants showed continuous distributions in callus growth and percentage of shoot regeneration, suggesting that these traits were controlled by polygenes. The F₂ population, derived from a cross between ‘Lenins’ and ‘6721′, showed a monogenic segregation for the number of regenerated shoots, and the segregation ratio fitted 1:2:1. Tissue culture traits of ‘Lenins’ were controlled by several genes, whereas the number of regenerated shoots related to the efficiency of shoot regeneration is controlled by one major gene.     

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.     

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.     

Genetic analysis of heading date and other agronomic characters in barley (Hordeum vulgare L.)

Genetic analyses of heading date, tiller number, plant height, grain yield, kernel weight, and plump and thin kernels were made in three six-rowed barley crosses (Hordeum vulgare L.) involving four cultivars. Six populations, P₁ , P₂ , F₁ , F₂ , BC₁ , and BC₂ , from each cross were grown and evaluated at Fargo and Prosper, North Dakota, 1982. Parental means within crosses generally were different except for tiller number. Comparison of generation means suggested that late heading was dominant to early, high kernel weight was dominant to low, and kernel plumpness was influenced by additive gene action. The relationship between yield and heading date was not consistent among crosses and positive r values were quite low. It should be possible to select early maturing, high yielding segregates with plump kernels. Heterosis over the mid-parent was quite similar among crosses for heading date, but there was no heterosis over the high parent. Inbreeding depression was fairly constant for heading date, but was less consistent for yield. The lack of uniformity for estimates of inbreeding depression can be related to environmental variation and to its influence on type of gene action. The ratio of additive to dominance variance was inconsistent among crosses for heading date and yield. These data suggest selection for these characters should be delayed past the F 2 generation. Broad sense heritabilities for heading date ranged from 42 to 86%. Values obtained for grain yield were more consistent among broad sense than narrow sense estimates. Genetic advance estimates were low due to lack of additive variance. This revised version was published online in July 2006 with corrections to the Cover Date.     

Predicting breeding values of spring barley accessions by using the singular value decomposition of genetic similarities

As pedigree information among parental lines is often incomplete, selection response can be enhanced if a matrix containing genetic similarities is used in the mixed model equations (MME) to predict breeding values. However, a low number of molecular markers may cause this matrix to be singular. This study was conducted to determine if breeding values are still accurate when a singular value decomposition of genetic similarities was performed setting negative singular values to zero. Two‐year data for four traits of 152 spring barley accessions, which were analyzed by 23 SSR markers, were used in the prediction of breeding values. In general, similar values of Akaike Information Criteria and of overall standard error of difference and a Spearman rank correlation coefficient between breeding values ranging from 0.92 to 0.99 were obtained independent of whether the prediction was based on a singular value decomposition of genetic similarities or not. In conclusion, the singular value decomposition of genetic similarities appears to be a suitable method in the case of singular matrices.     

Genetic variation for dry matter and nitrogen accumulation and translocation in two-rowed spring barley: I. Dry matter translocation

A 4-year field study was carried out to determine dry matter and nitrogen accumulation until anthesis and at grain filling period and dry matter translocation and utilization in grain filling of barley. Twenty two-rowed spring barley (Hordeum vulgare ssp. distichum L.) cultivars originated from different countries (Yugoslavia, Germany, Australia, the Czeck Republic, Netherlands, France and USA) were grown during 1995–1998 on a non-calcareous chernozem soil near Novi Sad (45° 20′N, 15° 51′E, 86 m asl). Dry matter and nitrogen accumulation depended on the cultivar and year. In a year with favorable weather conditions, 58% of dry matter was accumulated during pre-anthesis, while in a year with less favorable weather the amount was 48%. In the favorable year 91% and in unfavorable year 65% of nitrogen was accumulated until anthesis. The results indicated that the greater amount of dry matter and nitrogen accumulated before anthesis. Dry matter translocation efficiency depended on the cultivar and ranged from 3 to 16.4%, while the contribution of pre-anthesis assimilates to kernel varied from 4 to 24.2%. Cultivars that have been developed for the growing conditions of the area where the experimental site was located, i.e. adapted ones, did not use pre-anthesis dry matter for grain filling. High positive correlations (P<0.01) were found between biomass at anthesis and biological yield, dry matter translocation efficiency, contribution of translocated dry matter to grain yield, and total plant nitrogen at maturity. Accumulated nitrogen at anthesis was positively correlated (P<0.01) with growing degree–days until anthesis, dry matter at anthesis and dry matter translocation parameters. Heritability for the investigated characters was rather high, over 0.60.     

Screening for resistance in the primary and secondary gene pool of barley against the root-lesion nematode Pratylenchus neglectus

Root-lesion nematodes of the genus Pratylenchus are significant pests in crop cultivation throughout many parts of the world. A study was initiated to determine the resistance of Hordeum vulgare and H. vulgare ssp. spontaneum (wild barley) against one major representative of the genus Pratylenchus , P. neglectus . A glasshouse test was first established. Barley seedlings were grown in 20 cm³ tubes filled with sand. Each plant was inoculated with 400 P. neglectus juveniles. After 12 weeks of cultivation nematodes were isolated from roots and sand using a misting chamber. The nematodes were counted under a microscope. A representative collection of 565 barley and wild barley accessions was tested in this way. The average number of nematodes per accession ranged from 350 to 12 000. In a verification experiment, 35 accessions with low and high infection rates were tested. This experiment identified a number of accessions with low infection rates. The perspectives for future breeding of barley cultivars resistant to root-lesion nematodes are discussed.