Analysis of QTLs for yield components, agronomic traits, and disease resistance in an advanced backcross population of spring barley.

Hordeum vulgare subsp. spontaneum, the wild progenitor of barley, is a potential source of useful genetic variation for barley breeding programs. The objective of this study was to map quantitative trait loci (QTLs) in an advanced backcross population of barley. A total of 207 BC3 lines were developed using the 2-rowed German spring cultivar Hordeum vulgare subsp. vulgare 'Brenda' as a recurrent parent and the H. vulgare subsp. spontaneum accession HS584 as a donor parent. The lines were genotyped by 108 simple-sequence repeat (SSR) markers and evaluated in field tests for the measurement of grain yield and its components, such as ear length, spikelet number per spike, grain number per spike, spike number, and 1000-grain mass, as well as heading date and plant height. A total of 100 QTLs were detected. Ten QTLs with increasing effects were found for ear length, spikelet number, and grain number per spike. Three QTLs contributed by HS584 were found to significantly decrease days to heading across all years at 2 locations. In addition, 2 QTLs from HS584 on chromosomes 2H and 3H were associated with resistance to leaf rust. Based on genotypic data obtained from this population, 55 introgression lines carrying 1 or 2 donor segments were selected to develop a set of doubled-haploid lines, which will be used to reconfirm and investigate the effects of 100 QTLs for future genetic studies.     

优异大麦种质资源的抗性鉴定和评价

1998～1999年,将1997年筛选出的191个农艺性状表现优良的大麦品种进行大麦黄花叶病、大麦白粉病、大麦赤霉病、大麦条纹病及耐盐等抗性的鉴定评价,结果表明,抗及高抗大麦黄花叶病品种60份,抗大麦赤霉病品种59份,高抗大麦条纹病品种89份,抗大麦白粉病品种19份,芽期耐盐品种17份,苗期耐盐品种2份.     

Linkage disequilibrium mapping of yield and yield stability in modern spring barley cultivars

Associations between markers and complex quantitative traits were investigated in a collection of 146 modern two-row spring barley cultivars, representing the current commercial germ plasm in Europe. Using 236 AFLP markers, associations between markers were found for markers as far apart as 10 cM. Subsequently, for the 146 cultivars the complex traits mean yield, adaptability (Finlay-Wilkinson slope), and stability (deviations from regression) were estimated from the analysis of variety trial data. Regression of those traits on individual marker data disclosed marker-trait associations for mean yield and yield stability. Support for identified associations was obtained from association profiles, i.e., from plots of P-values against chromosome positions. In addition, many of the associated markers were located in regions where earlier QTL were found for yield and yield components. To study the oligogenic genetic base of the traits in more detail, multiple linear regression of the traits on markers was carried out, using stepwise selection. By this procedure, 18-20 markers that accounted for 40-58% of the variation were selected. Our results indicate that association mapping approaches can be a viable alternative to classical QTL approaches based on crosses between inbred lines, especially for complex traits with costly measurements.     

Molecular marker-assisted selection for enhanced yield in malting barley

Brewers are reluctant to change malting barley (Hordeum vulgare ssp. vulgare L.) cultivars due to concerns of altered flavor and brewing procedures. The U.S. Pacific Northwest is capable of producing high yielding, high quality malting barley but lacks adapted cultivars with desirable malting characteristics. Our goal was to develop high yielding near isogenic lines that maintain traditional malting quality characteristics by transferring quantitative trait loci (QTL) associated with yield, via molecular marker-assisted backcrossing, from the high yielding cv. Baronesse to the North American two-row malting barley industry standard cv. Harrington. For transfer, we targeted Baronesse chromosome 2HL and 3HL fragments presumed to contain QTL that affect yield. Analysis of genotype and yield data suggests that QTL reside at two regions, one on 2HL (ABG461C-MWG699) and one on 3HL (MWG571A-MWG961). Genotype and yield data indicate that additional Baronesse genome regions are probably involved, but need to be more precisely defined. Based on yield trials conducted over 22 environments and malting analyses from 6 environments, we selected one isogenic line (00-170) that has consistently produced yields equal to Baronesse while maintaining a Harrington-like malting quality profile. We conclude there is sufficient data to warrant experiments testing whether the 2HL and 3HL Baronesse QTL would be effective in increasing the yield of other low yielding barley cultivars.     

Variation in monoculture and in mixture for grain yield and other characters in spring barley

Fourteen north-west European spring barley cultivars were grown alone or in binary mixtures sampled according to a partial diallel scheme. On the basis of the association between cultivars in mixture and monoculture, three groups of characters were distinguished. Group A characters, plant height, ear weight/tiller, grain yield/tiller, number of grains/tiller and 1000-grain weight, showed strong positive associations between performance in monoculture and mixture. Group B characters, number of tillers/plant and harvest index showed incomplete positive associations, while for group C characters, dry matter/plant, ear weight/plant, grain yield/plant and number of grains/plant, associations were weak or non-existent. Compound characters in group C showed less genetic variation in monoculture and lower general competitive effects in mixture than component characters in groups A or B. These results clearly indicate that while selection for grain yield and other characters on a per plant basis (group C characters) is confounded by intergenotypic competition, characters such as the yield components number of grains/tiller and 1000-grain weight (group A characters) are hardly affected in this range of cultivars. Selection for opposing group A characters may start in the F₂ generation, while any selection for group B and C characters should be delayed until later generations. The merits of indirect selection for yield using visual assessment of yield components are discussed. Separate analyses obtained by the inclusion of spring wheat cv. Timmo in monoculture and in the set of mixtures indicated that the use of spring wheat plants to minimise intergenotypic competition ranges from superfluous (group A characters) to useless (group C characters). A large degree of mixture advantage and the lack of complementary dominance and suppression between competitor and associate was attributed to the relatively low density of plants in the experiment which, though suitable for single plant selection, is not typical of normal seed rates for cropping.     

Molecular breeding for grain yield in barley: an evaluation of QTL effects in a spring barley cross

 We report results from a breeding strategy designed to accumulate favorable QTL alleles for grain yield identified in the SteptoeבMorex’ (SM) barley germplasm. Two map lines (SM73 and SM145) from the original mapping population were selected based on their marker genotype and QTL structure. When crossed, these lines would be expected to produce progeny with most favorable QTL alleles. One hundred doubled haploid (DH) lines from the F₁ hybrid of this cross were genotyped with ten RFLP markers and one morphological marker defining grain yield to monitor QTL segregation. A subset of 24 lines representing various combinations of putatively favorable and unfavorable QTL alleles, together with Steptoe, ‘Morex’, SM73, and SM145, were phenotyped for grain yield in five environments. Multiple regression procedures were used to explore phenotype and genotype relationships. Most target QTLs showed significant effects. However, significance and magnitude of QTL effects and favorable QTL allele phase varied across environments. All target QTLs showed significant QTL-by-environment interaction (QTL×E), and the QTL on chromosome 2 expressed alternative favorable QTL alleles in different environments. Digenic epistatic effects were also detected between some QTL loci. For traits such as grain yield, marker-assisted selection efforts may be better targeted at determining optimum combinations of QTL alleles rather than pyramiding alleles detected in a reference mapping population. Received: 2 June 1998 / Accepted: 17 September 1998     

Effects of net blotch on growth and yield of spring barley

The effect of net blotch on the growth and yield of cv. Beatrice spring barley was examined in a greenhouse experiment. Separate inoculations at growth stages 21 and 34 reduced green leaf area, root weight, leaf sheath and stem weight and tiller number. The early inoculated plants, which responded and recovered more rapidly than later treated ones, suffered a loss in grain yield and this was related to the amount of disease, the loss in green leaf area and the reduction in unit leaf rate.     

Evidence of natural selection for disease resistance in Composite Cross Five (CCV) of barley

Cambridge Composite Cross Five (CCV) of barley was studied utilising hordeins, restriction fragment length polymorphisms (RFLPs) and reaction to powdery mildew with a view to understanding the genetic changes occurring in the population. Changes in the frequency of individual hordein patterns as well as pattern combinations showed directional trends in successive generations in three parallel populations maintained as discrete populations since 1977 in Cambridge. Certain hordein pattern combinations were more common in the resistance classes and there was a strong association between hordein patterns and mildew reaction. RFLP analysis revealed that 80% of a random sample taken from generation F₂₄ of Population I had the same restriction pattern as that of the cultivar Algerian, which was one of the original 30 parental lines of CCV. This cultivar is the source of the Mla1 allele in barley improvement programmes in Europe. We argue, based on supporting evidence from hordein analysis and tests of reaction to selected mildew isolates of known virulence isolates together with UK virulence surveys, that selection for Mla1 in Cambridge has been the predominant evolutionary force in CCV in Cambridge.     

Effects of aerial pollutants on the growth and yield of spring barley

Open-topped chambers were used to determine the effect of field concentrations of aerial pollutants on the growth and yield of spring barley. Experiments were conducted in the Marston Valley, Bedfordshire, where sulphur dioxide and fluoride are the major pollutants. The charcoal filtered chambers enabled pollutant concentrations to be reduced by 60–70%. Cleaning the air increased straw and grain yields. The filtration was non-selective and did not identify the injurious agent. The chambers were found to accelerate the crop's development by 7–8 days and reduce yield by suppressing tillering.     

Breeding winter barley for hardiness and disease resistance

This crop, valuable for pasturage and feed grain, is largely confined to the southern and eastern States, and northward extension of its range is promised by the prospects of success in breeding cold-resistant varieties.     

The use of canonical analysis for selection within a population of spring barley

A selection method is described involving the application of canonical analysis to data for a number of measured traits in a composite population of spring barley grown in two seasons. Lines were selected which fell within the Euclidean space defined by the range, for a number of standard cultivars, on each significant canonical variate. Alternatively, limits were fixed for an acceptable range for each trait and transformed to canonical variates. Lines were selected which fell nearest to the centroid of this acceptable space. The method might be used as an alternative to the selection index method where difficulty is often encountered in assigning appropriate economic weightings to the traits measured.     

Evaluation of chlorophyll fluorescence in different densities of spring barley

Over the period 2008–2009, field experiments were carried out at the Lithuanian Institute of Agriculture on an Endocalcari-Epihypogleyic Cambisol (CMg-p-w-can). The study was aimed to establish the effects of spring barley crop stand density on the chlorophyll fluorescence and to estimate the relationship between grain yield and chlorophyll fluorescence value. The tests involved three spring barley varieties—Aura DS, Barke and Gustav. The three different stand densities were formed with seed rates of two, four and six million viable seeds per hectare. Our research findings indicated that increasing spring barley stand density did not exert any negative influence on major photosynthetic processes and did not result in any significant qualitative changes in light harvesting system. A significant varietal (factor B) influence was established on minimum fluorescence (F _o) and maximum fluorescence (F _m) values after short dark adaptation indicators (F _fact. = 19.66** and F _fact. = 9.33**, respectively). Growth stage (factor C) significantly influenced all fluorescence indicators—Fo, Fm and quantum efficiency of PSII after short dark adaptation (F _v/F _m)—in the five cases of the six tested. A significant effect of variety and growth stage interaction (B × C) was determined for 66.7% of the tested cases. The article presents a correlation between the grain yield of spring barley and fluorescence parameters. In most cases, the correlation was strong and significant. The interaction between the fluorescence parameters for individual varieties was responsible for 53.8–76.2% of grain yield data variation, which averaged over all varieties, amounted to 21.9–46.1%.     

Detection of quantitative trait loci for agronomic,yield, grain and disease characters in spring barley (Hordeum vulgare L.)

Quantitative trait loci (QTLs) have been revealed for characters in a segregating population from a spring barley cross between genotypes adapted to North-West Europe. Transgressive segregation was found for all the characters, which was confirmed by the regular detection of positive and negative QTLs from both parents. A QTL for all the agronomic, yield and grain characters measured except thousand grain weight was found in the region of the denso dwarfing gene locus. There were considerable differences between the location of QTLs found in the present study and those found in previous studies of North American germ plasm, revealing the diversity between the two gene pools. Thirty-one QTLs were detected in more than one environment for the 13 characters studied, although many more were detected in just one environment. Whilst biometrical analyses suggested the presence of epistasis in the genetic control of some characters, there was little evidence of interactions between the QTLs apart from those associated with yield. QTLs of large effect sometimes masked the presence of QTLs of smaller effect.     

Genome architecture and diverged selection shaping pattern of genomic differentiation in wild barley

Divergent selection of populations in contrasting environments leads to functional genomic divergence. However, the genomic architecture underlying heterogeneous genomic differentiation remains poorly understood. Here, we de novo assembled two high-quality wild barley (Hordeum spontaneum K. Koch) genomes and examined genomic differentiation and gene expression patterns under abiotic stress in two populations. These two populations had a shared ancestry and originated in close geographic proximity but experienced different selective pressures due to their contrasting micro-environments. We identified structural variants that may have played significant roles in affecting genes potentially associated with well-differentiated phenotypes such as flowering time and drought response between two wild barley genomes. Among them, a 29-bp insertion into the promoter region formed a cis-regulatory element in the HvWRKY45 gene, which may contribute to enhanced tolerance to drought. A single SNP mutation in the promoter region may influence HvCO5 expression and be putatively linked to local flowering time adaptation. We also revealed significant genomic differentiation between the two populations with ongoing gene flow. Our results indicate that SNPs and small SVs link to genetic differentiation at the gene level through local adaptation and are maintained through divergent selection. In contrast, large chromosome inversions may have shaped the heterogeneous pattern of genomic differentiation along the chromosomes by suppressing chromosome recombination and gene flow. Our research offers novel insights into the genomic basis underlying local adaptation and provides valuable resources for the genetic improvement of cultivated barley.     

Field resistance of spring barley cultivars to powdery mildew in Lithuania

During vegetative period 2004–2005 powdery mildew (Erysiphe graminis DC. f. sp. hordei Em. Marchal) field resistance of spring barley cultivars was investigated at the Lithuanian Institute of Agriculture. The spring barley genotypes tested were Lithuania-registered cultivars, cultivars from genetic resources collection, and the new cultivars used for initial breeding. In total, 23 resistance genes were present in the 84 cultivars studied. Among mono-genes only mlo and 1-B-53 showed very high resistance. Slight powdery mildew necroses (up to 3 scores) formed on cultivars possessing these genes. The maximal powdery mildew (PM) severity reached a score of 8.5 and the area under disease progress curve (AUDPC) a value of 1216.8. The cultivars ‘Primus’, ‘Astoria’, ‘Power’, ‘Harrington’ and ‘Scarlett’ were the most resistant among the non mlo cultivars. Severity of PM on ‘Primus’ reached a score of 3.5 (3.0 of PM necrosis) in average, the other cultivars were diseased from 4.5 (3.0) to 5.0 (2.0). The AUDPC values for these cultivars except ‘Scarlett’ were the lowest (85.0–145.3) among the other cultivars. The highest contrast in development of the other leaf diseases was between highly resistant and susceptible to PM cultivar groups. The fast development of PM depressed development of the other diseases 4.7 times.     

Analysis of QTLs for yield, yield components, and malting quality in a BC3-DH population of spring barley

Advanced backcross (AB)-quantitative trait locus (QTL) analysis has been successfully applied for detecting and transferring QTLs from unadapted germplasm into elite breeding lines in various plant species. Here, we describe the application of a modified AB breeding scheme to spring barley. A BC₃-doubled haploid (DH) population consisting of 181 lines derived from the German spring barley cultivar Brenda (Hordeum vulgare subsp. vulgare) as the recurrent parent and the wild species line HS213 (H. vulgare subsp. spontaneum) as the donor line was evaluated for yield and its components as well as malting quality traits. A set of 60 microsatellite markers was used to genotype the population, and phenotypic data were collected at two locations in Germany in continuous years. Altogether, 25 significant QTLs were detected by single-marker regression analysis and interval mapping. Most positive QTLs originated from the recurrent parent Brenda. A QTL, Qhd2.1, on chromosome 2HS from Brenda explained 18.3% and 20.7% of the phenotypic variation for yield and heading date, respectively. Due to the small percentage of donor-parent genome of 6.25%, the BC₃-DH lines could be directly used for the extraction of near-isogenic lines (NILs) for Qhd2.1. Consequently, it was possible to determine the precise location of the locus hd2.1 within a region of 6.5 cM, using an F₂ population consisting of 234 individuals developed from a cross between an NIL containing a defined donor segment at this locus and Brenda. The location of this QTL was consistent with the presence of a major photoperiod response gene, Ppd-H1, previously reported in this region, which is associated with pleiotropic effects on yield components. In summary, the analysis of a BC₃-DH population in barley provides a compromise between the analysis of QTLs by means of an AB scheme and the generation of defined substitution lines. Several lines carrying defined different donor segments for only one single chromosome or trait in the genetic background of Brenda could be selected for further genetic studies.     

In vitro selection of barley and wheat for resistance against Helminthosporium sativum

Summary Calli derived from immature embryos of barley and wheat genotypes were screened for their resistance to purified culture filtrate produced by the fungus Helminthosporium sativum P.K. and B. Two selection methods were used: a continuous method in which four cycles of selection were performed one after another on toxic medium and a discontinuous method in which a pause on non-toxic medium was given after the second or third cycle of selection. The latter was superior as it allowed the calli to regain their regeneration ability. About 3,000 calli from two barley genotypes and 2,000 from two wheat genotypes were used for selection. The selection with the pathotoxins resulted in 6% to 17% surviving calli. Toxin tolerant callus lines of barley were characterised by protein isozymes. Zymograms showed one more isozyme than with the unselected sensitive callus. Barley and wheat plants have been regenerated from callus lines surviving the toxin treatment and in vivo testing against pathogen revealed that the majority of these plants were less sensitive.     

The effect of combining scald resistance genes on disease levels, yield and quality traits in barley

Pairwise combinations of genes for resistance to scald in barley were developed using linked isozyme markers to test whether such combinations conferred improved resistance to the pathogen, Rhynchosporium secalis. The resistance genes originally derived from Hordeum vulgare ssp. spontaneum. The combinations were bred into an essentially similar genetic background because the scald-susceptible, Australian barley cultivar Clipper was the recurrent backcross parent in their ancestry. In field tests of the recombinants over 2 years, disease levels were lower in three of six doubly resistant lines than in backcross lines carrying a single resistance gene, which in turn were less diseased than either Clipper or recombinants that lacked the marked resistance genes. All resistant lines significantly outyielded Clipper but did not themselves differ significantly. Lines resistant to scald had significantly higher grain size and grain weight. Gains for malt yield of about 1 % were detected in the higher disease environment. Resistance was not accompanied by any obvious cost in terms of yield or quality. Protection against scald is therefore a significant requirement for new malting barley cultivars in scald-prone areas.     

The efficiency of visual assessment of grain yield and its components in spring barley rows

The efficiency of visual assessment for grain yield and its components in spring barley rows was examined using a number of assessors. In 20 out of 28 combinations of assessor and character, only 20 or less lines were needed to be retained from the set of 99 lines to save at least 50% of the best 10 lines. Assessments of tillers/row and to a lesser extent 1000-grain weight were generally more effective than assessments of yield/row and grain/ear. The low effectiveness of assessment of grains/ear was attributed to inadequate sampling of ears. Four out of five assessors showed bias towards assessment of tillers/row, the most easily assessed character, in their assessments of yield/row. Experienced barley workers were more successful in their assessments than others less familiar with the crop. Specially developed keys, aimed at making visual assessment more objective, generally had only small positive effects on the efficiency of assessment. Repeated assessments of characters by some assessors were consistent. It was concluded that visual assessment rather than direct measurement should be recognised as a basic tool of breeding in the early generations, using a large number of lines as a ‘safety net’ allowing the loss of some of the best lines.