Cryopreservation of Embryogenic Callus Cultures from Barley (Hordeum vulgare L.)

Embryogenic callus cultures of barley (Hordeum vulgare L.) were frozen in the presence of a cryoprotector and subsequentjy stored in liquid nitrogen. After thawing, a high percentage of cultures resumed growth after a lag-period of 2–4 weeks. Plant regeneration was achieved at a frequency comparable to that observed in control cultures.     

Spring Cereals for Forage and Grain Production in a Cool Maritime Climate

Newfoundland's climate is marginal for agricultural production. The availability of locally grown cereal grain and high‐quality forage are major limitations to successful animal agriculture in this region. Here, our overall objective was to compare several spring cereal species for both annual forage and grain production in Newfoundland's cool Maritime climate. Several varieties of barley (Hordeum vulgare L.), wheat (Triticum aesitivum L.), oats (Avena sativa L.) and pea (Pisum sativum L.)–cereal mixtures for forage yield and quality, as well as grain yield and maturity, were compared in field trials on the east and west coasts in both 1999 and 2000. Barley headed earliest, yielded greatest forage dry matter, had lowest forage protein and acid detergent fibre (ADF) percentages, and had neutral detergent fibre (NDF) mean values greater than those of pea–cereal mixtures, but less than those of oats and wheat. Forage harvested from pea–cereal mixtures was similar to that of barley for yield, ADF and NDF, while P and protein percentage were much greater. Barley matured 10–15 days earlier than both wheat and oats. In general terms, all three spring cereals exhibited similar grain yield potential. Oats tillered less, but compensated by producing more kernels spike^?1. Days to maturity for cereal grains in western Newfoundland were roughly similar to those reported for the Maritime provinces of Canada. Yield and maturity results for both forage and grain production suggest that eastern Newfoundland is a unique agro‐ecoregion in North America, and agronomic recommendations specific to other regions may not be applicable in this region.     

The Effect of Carbohydrate Composition and Concentration on Anther Culture Response in Barley (Hordeum vulgare L.)

Culture medium composition is critical for the successful induction of microspore division in vitro. The present experiments have focused on a relatively neglected area of cell and tissue culture research, namely the carbohydrate component used in the medium. Three spring barley genotypes were cultured on a medium which was modified by replacing sucrose with the following carbohydrates (6 % w/v): maltose, fructose, malt extract, galactose and a glucose (3 % w/v)/fructose (3 % w/v) mixture. Both maltose and malt extract were superior to sucrose in their capacity to induce green plantlet differentiation from microspores. The concentrations of both sucrose and maltose were also varied. Overall the response of anthers on maltose based media was higher than on sucrose based media. Furthermore, a concentration of maltose m the range 6—12 % w/v produced a higher frequency of green plants than a low concentration (1—3 % w/v). The effect of maltose based media on germplasm of direct relevance to barley breeders was also tested. The cultivar ‘Blenheim’ was shown to be very responsive and this genetic factor was transmitted to the F₁ hybrid. The frequency of haploid to diploid regenerants was not consistent over genotypes, but in general there were more haploid than diploid regenerants. The implications of these results for barley breeding are discussed.     

大麦基因组GRAS基因家族的全基因组鉴定与表达分析

GRAS基因家族是一类仅存在于植物并广泛参与其生长发育调控的转录因子,根据其序列结构和系统发育树分化特征,GRAS转录因子包含PATI,DELLA,HAM,SCR,SHR等多个亚家族成员.本研究利用大麦最新的基因组数据库,采用生物信息学的方法筛选鉴定出41条GRAS基因序列,其中有34条序列具有完整的GRAS家族蛋白特...     

The Effect of Seeding Rate and Nitrogen Fertilization on Barley Yield and Yield Components in a Cool Maritime Climate

Soil and climatic conditions in Newfoundland are on the margins of agricultural capability, and almost all feed grain is imported. The overall objective of this work was to develop guidelines for the production of barley in Newfoundland, with the goal of establishing modern cropping recommendations. We conducted a 4-year study near St. John's to examine the effect of seeding rate and topdress ammonium nitrate (N) fertilization rate on Chapais six-row barley ( Hordeum vulgare L.) yield components and grain yield. Increasing seeding rate from 200 to 380 seeds m^–2 did not alter grain yield in any year. Increasing topdress fertilization from 0 to 60 kg N ha^–1 increased spike density m^–2 at harvest, resulting in linear increases in grain yield in all years. Highest N rates had greatest lodging in two years. Based on our results, agronomic recommendations for eastern Newfoundland now include barley seeding rates of 250 ± 50 seeds m^–2, with topdress applications up to at least 30 kg N ha^–1.     

An Evaluation of Hordeum vulgare L. ×Psathyrostachys fragilis (Boiss.) Nevski Crosses for Doubled Haploid Barley Production

Investigations were carried out to assess the suitability of the intergeneric cross Hordeum vulgare×Psathyrostachys fragilis for haploid barley production. H. vulgare cvs. ‘Emir’ and ‘Vada’ were each pollinated with P. fragilis P.I. 343192 and plants regenerated from embryos cultured on a modified B5 medium. Seed sets on ‘Vada’ were significantly lower than on ‘Emir’, and all the planes from ‘Vada’×P. fragile remained hybrid. Several of these flowered but there was little pairing between the parental chromosomes. Most of the plants from ‘Emir’×P. fragilis died, as seedlings but 3 plants developed into haploid barley. Because of the practical limitations of pollen availability from P. fragilis and the inconsistencies in haploid plant formation, it is unlikely that the cross will prove as valuable as that between H. vulgare×H. bulbosum for a doubled haploid barley programme.     

Studies on Low Crossabilities Encountered with the Hordeum bulbosum Method for Haploid Production of Barley, Hordeum vulgare L.

Crosses were made between four spring barley (Hordeum vulgare L.) cultivars and five F₁, hybrids with one genotype of Hordeum bulbosum L. in two locations to investigate further previous low crossabilities which had been found in the barley cultivar ‘Apex’ with H. bulbosum. Data at all the main steps of the H. bulbosum technique were recorded and statistically analyzed. Significant differences between barley genotypes were demonstrated for all characters. It was confirmed that ‘Apex’ has poor crossability with H. bulbosum. Out of the three F₁ hybrids having ‘Apex’ as one parent, two exhibited low crossability similarly to ‘Apex’ but one showed significantly higher seed setting than ‘Apex’. The effect of the location was only significant on seed setting, while genotype X location interactions were significant on seed setting, seed quality and rate of haploid plants in relation 10 florets pollinated. Another problem which has influenced the success rate of the H. bulbosum method was found in the cultivar ‘Havilla’. Although seed setting and seed quality were high for this cultivar, embryo differentiation was low. However, this latter problem was found to influence less the overall success rate than poor crossability. Mahalanobis's distances were calculated and the dendrite of the shortest distances between barley genotypes was plotted.     

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.     

Using Hemispherical Radiation Measurements to Predict Weight-Related Growth Traits in Oilseed Rape (Brassica napus L.) and Barley (Hordeum vulgare L.) Canopies

Methods for determining growth traits in the field are mostly work‐intensive and/or often erroneous. Further development of multi‐temporal techniques of growth analyses provided by fast and non‐destructive tools for monitoring canopy properties may result in a substantial gain for practical use. Among them, the LAI‐2000 Canopy Analyzer can be used to determine the canopy area index (CAI) resulting from radiation measurements. In plant breeding, single‐date measurements were usually made across a wide number of genoptypes. In contrast, multi‐temporal information on canopy status is needed in agronomy. In our study, data of the CAI based on hemispherical radiation measurements with the LAI‐2000 were used to estimate developmental courses of weight‐related growth traits. For this purpose, seasonal courses of shoot fresh matter, shoot dry matter, shoot N and water content and CAI of winter oilseed rape (Brassica napus L.) and spring barley (Hordeum vulgare L.) were investigated simultaneously in field experiments from 2002 to 2006 in northern Germany with and without nitrogen (N) fertilization. Courses of 2003 and 2004 were used to calculate relationships between CAI and growth traits. Results show that each of the weight‐related growth trait is related to the CAI. In this mechanistic approach relationships were found gathered across 2 years, two N‐treatments and several growth stages being unaffected by N‐fertilization, cultivar and experimental year. These relationships were verified using independent data sets of the years 2005 (winter oilseed rape and spring barley) and 2006 (winter oilseed rape). The results of the current study show that the use of LAI‐2000 and its rapid and non‐destructive determination of weight‐related growth traits over the entire growth period will substantially improve the determination of weight‐related growth traits in agriculture.     

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.     

Effects of plot type on detection of quantitative-trait-locus effects in barley (Hordeum vulgare L.)

Hill plots offer advantages in testing large germplasm arrays with limited area and seed. However, the relationship of trait expression between hill vs. row plots remains in question. The development of quantitative-trait-locus (QTL) analysis offers alternative methods of testing hypotheses regarding levels of coincident gene effects in divergent plots. The objective of this study was to compare hill and row plots in terms of QTL detection, number of common QTLs and plot-type-specific QTLs for various characters in barley. Assessments were based on sets of 100 F₁-derived doubled haploid progeny from two environments of hill-plot data and two sets of multiple-environment row-plot data. Common large-effect QTLs for height, heading date, and 1000-kernel weight were detected in both plot types. Fewer QTLs were detected in the hill-plot data, with only one QTL detected for grain yield. There were no hill-plot-specific QTL effects. Hill plots appear to be best suited to highly heritable characters such as height, heading date, and 1000-kernel weight. Evaluation of grain yield is best deferred until larger experimental units can be employed.     

Breeding Progress in Grain Yield and Selected Agronomic Characters of Winter Barley (Hordeum vulgare L.) over the Last Quarter of a Century

This study gives estimates of breeding progress achieved in winter barley (Hordeum vulgare L.) since the 1960′s in grain yield and the change in several morphological traits for seventeen historically important cultivars: 12 six-rowed and 5 two-rowed, grouped into four epochs according to the period of major commercialization in Italy. All genotypes were evaluated across a range of soil fertility and climatic environments. The modern six-rowed cultivars have shown a consistent increase in tillers/m² and seeds/spike (16 and 28 %, respectively) over the local populations. In the modern two-rowed cultivar ‘Igri’ the increase in tillers/m² and seed weight were, respectively, 20 and 14 % snd seeds/spike decreased by 20 % over the local populations. The modern six-rowed and two-rowed cultivars have less lodging, are earlier and shorter than the old varieties and local populations. In the last quarter of a century, the breeding progress for gram yield has been increased by 52 and 74 kg/ ha/year or 0.75 and 1.1 % per year respectively, for six-rowed and two-rowed genotypes: while the gain in biomass yield in the same period was not uniform for six-rowed cuitivars and increased by 64 kg/ha/ year or 0.46 % in two-rowed cultivars. The grain yield improvement in the modern six-rowed and two-rowed cultivars is the result of a better partitioning of the photosynthetic products into the grain.     

Resistance to the Barley Yellow Mosaic Virus Complex — Differential Genotypic Reactions and Genetics of BaMMV-Resistance of Barley (Hordeum vulgare L.)

Barley yellow mosaic disease is caused by several viruses, i.e. barley yellow mosaic virus (BaYMV), barley mild mosaic virus (BaMMV) and BaYMV-2. The reaction of different barley germplasms to the barley mosaic viruses was studied in field and greenhouse experiments. The results show a complex situation; some varieties are resistant to all the viruses, while others are resistant to one or two of them only. Crosses between different barley germplasms were earned out in order to test whether genetic diversity of resistance against mosaic viruses does exist, particularly, BaMMV. A total of 45 foreign barley varieties were crossed to German cultivars carrying the resistance gene ym4. In F₂ of 27 crosses, no segregation could be detected, leading to the conclusion that the resistance genes of the foreign parents are allelic with ym4 e.g. Ym1 (‘Mokusekko 3’) and Ym2 (‘Mihori Hadaka 3’). A total of 18 crosses segregated in F₂ indicating that foreign parents, like ‘Chikurin Ibaraki 1’, ‘Iwate Omugi 1’, and “Anson Barley”, carry resistance genes different from the gene of German cultivars, e.g. ‘Asorbia’ or ‘Franka’. By means of statistical evaluation (Chi²-test), the observed segregation ratios were analyzed in order to obtain significant information on the heredity of resistance. All the resistance genes described here as being different from the gene ym4, act recessively. Most of the exotic varieties seem to carry only one resistance gene. In a few cases, more than one gene may be present.     

不同磷水平下大麦分蘖期磷效率相关性状QTL定位分析

磷素营养与大麦品质及产量密切相关,磷高效遗传机制和品种改良是近年的研究热点之一。本研究利用由大麦栽培品种Baudin和种质材料CN4079杂交构建的重组自交系(RIL)群体,低磷胁迫(0.02 mmol L~(-1) KH_2PO_4)与正常供磷(0.2 mmol L~(-1) KH_2PO_4)条件下,对地上部和地下部磷素利用效率、磷素吸收效率和干重,以及分蘖数相关的QTL定位,并预测相关位点基因。表型鉴定结果表明,各性状在RIL群体中表现连续变异,并存在超亲分离。两种磷水平下,共检测到16个QTL,分布在2H、3H和5H染色体上,表型贡献率14.1%~28.5%。3H染色体上含有3个磷素利用效率位点,其增效等位基因均来源于Baudin,其中Qspue.sau-3H.1和Qrpue.sau-3H与控制磷素吸收效率的Qspae.sau-3H和Qrpae.sau-3H处于同一区段,而Qspue.sau-3H.2与控制分蘖数的位点Qtn.sau-3H处于同一区段。5H染色体上含有3个磷素吸收效率位点,其中Qspae.sau-5H.2和Qrpae.sau-5H的增效等位基因来自CN4079,且与控制磷素利用效率的Qspue.sau-5H和Qrpue.sau-5H,以及控制干重的Qsdw.sau-5H和Qrdw.sau-5H处于同一区段。在磷效率相关的4个区段中,除Qspue.sau-3H.1所处区间仅含有磷酸代谢与磷脂代谢相关基因外,其他区间均包含磷酸盐转运蛋白基因、磷酸代谢与磷脂代谢相关基因。     

Cloning, genetic and physical mapping of resistance gene analogs in barley (Hordeum vulgare L.)

The majority of verified plant disease resistance genes isolated to date belong to the NBS‐LRR class, encoding proteins with a predicted nucleotide binding site (NBS) and a leucine‐rich repeat (LRR) region. Using degenerate primers, designed from the conserved motifs of the NBS region in tobacco N and Arabidopsis RPS2 genes, we isolated 190 resistance gene analogs (RGA) clones from barley genomic DNA. A total of 13 single‐ and low‐copy RGAs were genetically mapped onto chromosomes 1H–7H (except 5H) using three barley double haploid (DH) mapping populations: Steptoe × Morex, Harrington × TR306 and LUGC × Bowman. Sequence analysis of the RGAs showed that they are members of a diverse group. As a result of BLAST searches, one RGA proved unique as it did not detect any significant hit. Another RGA is putatively functional, because it detected several barley expressed sequence tag (EST) matches. To physically map the RGAs, 13 sequences were used to screen a 6.3 × cv. ‘Morex’ bacterial artificial chromosome (BAC) library. After fingerprint analysis, eight contigs were constructed incorporating 62 BAC clones. These BAC contigs are of great value for positional cloning of disease resistance genes, because they span the regions where various barley R genes have been genetically mapped.     

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.     

Adaptation of barley (Hordeum vulgare L.) to harsh Mediterranean environments

Summary Differences in development of the apex may be the reason for cultivar differences in adaptation of barley to terminal drought in Mediterranean environments. The present study was conducted to identify apical development patterns of barley adapted to terminal drought stressed Mediterranean environments and to determine plant characteristics which can be used as criteria to select for an adapted development. Thirty-five two-rowed barley (Hordeum vulgare L.) entries were grown at two sites in northern Syria (Tel Hadya and Breda) in 1988/89. Four apical development patterns were observed: a slow or fast vegetative development, depending on the vernalization requirement, combined with a slow or fast generative development, depending on the daylength response of the crop. Early heading was related to fast generative development. Leaf appearance rates on the main shoot were constant during a major part of the pre-anthesis period, but significant differences were observed among development patterns. Genotypic differences in main shoot tiller number were associated with differences in the onset of tiller appearance and not with differences in tiller appearance rate or final leaf number on the main shoot. Since vernalization requirements and daylength responses are largely independent of terminal drought stress, selection for an adapted phenology can be done in favourable environments. Morphological traits related to these responses (winter growth habit, cold tolerance, plant colour, growth vigour, heading date) can be used as criteria for selection for adaptation to low-rainfall Mediterranean environments.     

Effect of powdery mildew (Erysiphe graminis f.sp. hordei) on photosynthesis and grain yield of partially resistant genotypes of spring barley (Hordeum vulgare L.)

The effect of powdery mildew on the photosynthesis and grain yield of partially resistant spring barley was investigated. The effect of powdery mildew on the CO₂ exchange rate (CER) of inoculated and non-inoculated leaves was measured in several glasshouse experiments. Grain yield reduction by powdery mildew was determined in three field experiments. The CER of the inoculated leaves was reduced by infection but that of the non-inoculated leaves was not significantly different from that of the corresponding leaves of the control plants. The reduction in CER of the different genotypes was closely related to the percentage leaf area infected with powdery mildew. The correlation between degree of mildew infection and grain yield reduction varied between r = 0.84 (significant at P = 0.01) and r = 0.41 (not significant). Differences in greenness of the crop were correlated better with grain yield reduction (r = 0.66, P = 0.01) than powdery-mildew infection itself. Low percentages of powdery-mildew infection did not lead to significant reduction in grain yield. Partial resistance to powdery mildew could thus be a valuable attribute of future barley varieties.     

Use of Isozyme-Marked Segments from Wild Barley (Hordeum spontaneum) in Barley Breeding

Fifty-six accessions of Hordeum spontaneum from various Israeli habitats were hybridized with, and their progeny backcrossed three times to H. vulgare cv. ‘Clipper’. Selected chromosome segments marked by one of 24 isozyme loci were transferred to form 84 backcross lines which were nearly isogenic with ‘Clipper’. Each line was made homozygous for a single isozyme-marked segment. These lines were evaluated in preliminary and advanced field trials. The yield, malt extract percentage and grain size of the lines approached or equalled, and sometimes significantly surpassed, that of ‘Clipper’ and more recent commercial cultivars. Three isozyme-marked segments were combined in pairs and the three resultant doubly-marked lines were compared with the reconstituted parental, singly-marked lines and with the ‘Clipper’ recombinant from the same F²-array. The additive effects of single segments on yield or gram size were not significant, whereas half of the six interactions were significant, but in the unfavorable direction. It was concluded that near-isogenic lines can identify segments of wild barley germplasm that are useful for improving yield, but that pairs of these from different habitats may interact unfavourably.