Direct and Correlated Responses to Four Cycles of Divergent Selection for Heading Date in Barley*

In order to investigate the direct and correlated responses to selection for heading date in barley, two F₂ populations were subjected to four cycles of divergent selection. For each cross, eight populations selected for earliness (from 1E to 4E) and for lateness, (from 1L to 4L) and four unselected populations (1, 2, 3 and 4 SSD) were obtained and than compared together with parental lines. Selection caused a change greater than one day/cycle in Doth directions, and the cumulative responses were -5.4 and -5,9 days in the direction of earliness, 7.7 and 7,1 days in the opposite direction. Such results suggest that in the two base populations the heading, date is under the control of a polygenic system with prevailing additive effects and a slight directional dominance toward earliness. Early populations showed a reduction of ear moisture and an increase of plant height, ear extrusion, kernel weight and harvest index, whereas late populations showed the opposite changes. Grain yield was linearly reduced by selection for lateness, whereas no substantial change was evidenced in the opposite direction, so that 4E significantly out-yielded 4L in both crosses.     

Selection of Barley Lines for Drought Tolerance in Low-Rainfall Areas

An experiment was conducted at five locations in Jordan (Khanasri, annual rainfall 150 mm; Ramtha, 225 mm; Muwaqar, 150 mm; Rabba, 350 mm; Ghweer, 250 mm) during the 1996/1997 growing season, to evaluate the yield performance and some agronomic traits of 84 barley breeding lines and three long‐term checks (Zanbaka, Arta and Klaxon) for drought tolerance. An α‐lattice design with two replications and six incomplete blocks for each replication was used. Genotype 6 (WI2291/Tadmor) was found to be superior in grain yield, especially compared to the best check (Arta) at the two wettest locations (Rabba and Ghweer). Genotype 21 (Mo.B1337/WI2291/5/Emir/Sb//CM67/3/F8‐HB‐854‐23/121//148‐221/4/CI 08887/CI05761) out‐yielded the best check (Arta) at the driest location (Khanasri), while genotype 61 (Salmas/Arabia Aswad) produced a higher grain yield than the best check (Zanbaka) at Ramtha. The correlations amongst grain yield, biological yield, straw yield, plant height and harvest index were always significant and positive regardless of the location. The correlations amongst days to heading, days to maturity and grain yield were significant at the two driest locations only. This suggests that different phonologies are required to maximize grain yield in wet and dry environments. Also, the relationship between grain yield and the length of grain‐filling period was positive in the wettest location (Rabba), negative in the driest (Khanasri) and not significant at the two intermediate locations. These results emphasize the importance of selection in the target environment and the need to develop early‐maturing genotypes as a way of withstanding drought and high temperatures during the grain‐filling period. A high and negative correlation coefficient was found between the drought susceptibility index and grain yield at the driest site, whereas at the wettest site the correlation coefficient was lower and in some cases positive, indicating the existence of traits that are desirable under drought and undesirable under favourable conditions.     

Changes in Frequency of Plants Resistant to Barley Leaf Rust Caused by Puccinia hordei Otth. in a Barley Composite Cross Population

The response of barley Composite Cross XLI to selection for resistance to leaf rust was evaluated. Representative samples from eight cycles of recurrent selection were inoculated with an isolate of Puccinia bordei Otth. in both the seedling and adult plant stage. Disease symptoms were evaluated 14 days after inoculation. The resistance in CC XLI increased with selection in the seedling stage from 15.1 to 20.5 %, and in the adult plant stage from 20.5 to 37.5 %, respectively. The largest gains in resistance were observed in the earlier cycles of recurrent selection. Significant fluctuations in percent resistant plants occurred between the years.     

Screening by PCR for Defined DNA Sequences in Minimal Amounts of Barley Tissue

The potential use of the polymerase chain reaction (PCR) in screening plants for defined DNA sequences was demonstrated in barley (Hordeum vulgare). As templates for PCR, purified DNA as well as DNA from crude cell extracts of different barley tissues can be used. Specific synthesis of the target sequence was possible in both cases. Under optimal conditions, as little as 20 copies of the barley genome, i.e., 20 cells, were sufficient for successful amplification. This amount of DNA is equivalent to significantly less than 1 mg of fresh tissue.     

Classification of Global Environments and Cultivars of Spring Barley Based on Heading Time Interactions

Global heading time data collected by the International Center for Maize and Wheat Improvement (CIMMYT) during the International Barley Yield Trials were used to assess similarity of environments, variation among cultivars, and genotype × environment interactions. Data for 29 spring barley cultivars grown in 89 environments over three years were analyzed by cluster analysis. The deviation m days to heading of the mean of 28 photoperiod sensitive cultivars from the cultivar ‘Mona’ and the mean of ‘Mona’, homozygous recessive for the ea_k gene conferring photoperiod insensitivity and thermal stability, were used as environmental coordinates. In addition, diversity of heading time responses among genotypes was illustrated by differences among overall means and patterns of deviation for days to heading from ‘Mona’ in selected environments. Three main clusters were identified. Mexican environments were similar to warmer Mediterranean, eastern and southern African, West Asian, and Latin American environments. Heading time responses in Syria were similar to those observed m other cool Mediterranean environments. Early heading cultivars exhibited greater variation for heading response, especially in extremely warm-and equaiorial- short daylength environments, com-pared to late heading cultivars, presumably because of larger photothermal × genotype interactions. Photoperiod flux about the winter solstice appeared to be a major environmental cue for heading time in photoperiod sensitive spring barleys.     

Components of Response in Barley Anther Culture

Anther culture response with 17 widely-grown varieties and one model variety of barley was studied with one replication from field-grown donor plants and one replication from a growth-chamber. Plants were regenerated from all 18 varieties and green plants were obtained from 16 of them. On average, 1.6 green plants were obtained per 100 cultured anthers from all the material. Estimated variance components for the formation of embryos/callus from the anthers were dominated by the effects of the genotypes and interactions between plant material and environments which together accounted for 60.1 and 17.0 % of the total variation respectively, while environments were nonsignificant for this character. Plant regeneration from embryos/callus were not significantly influenced by either genotype or environments. Components of variance for green plant formation were dominated by the effects of the genotypes, accounting for 73.2 % of the total variation, and a smaller effect from environments accounting for 11.2% of the total variation. Main effects from genotypes on the percentage of green regenerants divided 7 varieties into two distinct groups, indicating that major genetic factors were involved. The genetic basis for green plant regeneration seems different from that governing embryo formation. The results are discussed with respect to the possible prediction of anther culture response for new barley hybrids, as a means for directing the use of barley anther culture towards material that responds well.     

Performance of S2 Winter Barley Progenies from Original and Improved Populations Developed via Recurrent Selection

The effect of recurrent selection procedure for improving grain yield in autogamous cereals was evaluated in a six-rowed winter barley population. Gain from selection was estimated by field testing 90 random S2 lines from each of the C₀, C₁ and C₂ populations. Response to selection for grain yield was 0.78 t/ha from C₀ to C₁ and 1.09 t/ha from C₁ to C₂. Broad-sense heritability and genotypic variance for grain yield remained high in all cycles which suggests further gain from additional selection cycles. Yield increase was due to a higher number of seeds/m2. Significant differences among mean values were observed for heading date (189 days in C₀ and 182 days in C₂), whereas no variation was seen for plant height and 1000-kernel weight. The proposed recurrent selection procedure appears effective to improve the population and to extract superior genotypes for varietal development.     

Diallel Analysis of Grain Yield in Barley Using Doubled-Haploid Lines*

Grain yields of seven barley cultivars (Hordeum vulgare L.) and 398 doubled-haploid lines derived from their diallel crosses were evaluated in hill plots at two locations in Ontario. Additive × additive epistasis and linkage disequilibrium were detected in the seven cultivars. Additive × additive genetic variance was significantly greater than zero but additive genetic variance was not significantly greater than zero. The narrow-sense heritability on the basis of line means was intermediate (0.33). It was predicted that 13.1% yield improvement could be realized in the next recurrent selection cycle using the doubled-haploid method.     

In vitro Selection for Fusaric Acid Resistant Barley Plants

Calli of two genotypes of barley,‘Dissa’and W 193, were used for selection of resistance against fusaric acid, a pathotoxin of Fusarium. Callus was initiated from 7- to 10 days old immature embryos. 1000 calli of the‘Dissa’and 500 of the W 193 genotypes were grown for 4 selection cycles on medium with 0.8 mM fusaric acid. In the first selection cycle, about 80 % of the calli were killed; after the 4 selection cycles, 8 to 11 % resistant calli were obtained and plants were regenerated. Resistant calli maintained on non-toxic medium showed retention of resistance ability after 3 months of sub-culturing. Plants could be regenerated from the surviving calli and testing by leaf bioassay revealed that many were resistant to the same toxin concentration employed for callus selection (100 %), while some were only resistant up to a concentration of 75 %.     

基于高密度Bin图谱的水稻抽穗期QTL定位

以粳稻品种02428和籼稻品种玉针香进行杂交, 按单粒传法连续自交10代, 得到包含192个株系的重组自交系(RIL)作图群体。通过对两亲本重测序及RIL群体简化基因组测序, 构建了包含2711个Bin标记的高密度遗传图谱。该图谱各染色体标记数在162~311个之间, 标记间平均物理距离为137.68 kb。将亲本及192个株系分别于4个环境下采用随机区组种植, 并记录抽穗期。使用WinQTL Cartographer 2.5软件的CIM分析方法, 进行抽穗期相关QTL检测及定位。在4个环境下定位到影响抽穗期的QTL共14个, 分布于第1、第2、第3、第7、第8、第9和第10染色体。其中, qHD2.2和qHD10.2能在3个环境中被重复检测到, 表型贡献率分别为5.14%~11.15%和5.35%~16.97%, 分别能缩短抽穗期1.66 d和1.56 d, 具有聚合育种的应用价值。通过物理位置比对, 14个QTL中有11个与前人定位在相同或邻近区域, qHD1.1、qHD2.2和qHD9.1尚未见报道。经对qHD2.2详细分析, 在其染色体区间内找到3个与抽穗期相关的注释基因LOC_Os02g46450、LOC_Os02g46710和LOC_Os02g46940, 其中LOC_Os02g46450已被克隆。测序分析发现, 这3个基因在两亲本间都存在差异, 可作为候选基因。     

Development of RFLP Markers for Barley

A PstI-based genomic library from barley DNA was screened for RFLPs in the three relatively-distant cultivars ‘Alexis’ (2-row spring type), ‘Igri’ (2-row winter type) and ‘Mammut’ (6-row winter type), digested with BamHI, EcoRI and HindIII. 50 % of the 108 DNA fragments studied represented single-copy sequences, 29 % low-copy and 21 % repetitive sequences. The DNA probes were assigned to discrete barley chromosomes with the aid of wheat/barley addition lines. 80 % of the single- and low-copy sequences hybridized with both barley and wheat DNA, whereas most repetitive sequences gave signals only with barley DNA.     

Male Sterility in Barley

The anther form and development of eight two-rowed genie male sterile barley mutants are abnormal. The male sterility in each mutant is conditioned by single non-allelic recessive genes. These mutant genes cause reductions in anther size and lumen diameter. Though in seven mutants, the male meiosis is normal until microspore liberation, the micro-spores abort in all cases after release from the PMCs. In one mutant, the microspores degenerate at the tetrad stage before release from the PMCs. In four mutants, the tapetal development and disintegration are normal, in four others they are abnormal. Despite these variations, the male sterility in all the eight mutants is complete.     

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.     

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.     

Divergent selection for sprouting resistance in spring wheat

The development of sprouting-resistant spring-wheat ( Triticum aestivum L.) cultivars is a major breeding objective in many wheat-producing regions. Sprouting resistance is thought to be associated with delayed maturity. The primary objective of this study was to measure the reciprocal effects of selection for sprouting resistance and maturity. Two experiments were conducted over a 3–4-year period in Saskatoon, Canada. In the first experiment, two populations of hard red spring wheat were subjected to divergent selection (k = 10%) for maturity. In the second experiment, six populations derived from crosses between two sprouting-resistant, late-maturing, white-grained cultivars ('AUS1293' and 'AUS1408') and three early maturing, red–grained cultivars ('Park', 'PT516' and 'Roblin'), were subjected to divergent selection (k = 10%) for sprouting resistance. Selection for earliness reduced sprouting resistance in one population but had no effect in the second. For both populations, earlier maturity was associated with higher test weight but lower grain yield. In the second experiment, selection for increased sprouting resistance was effective, with realized heritabilities averaging 0.74. Increased sprouting resistance was associated with a slight delay (1–2.5 days) in time to spike emergence in four out of six populations, but had little effect on time to maturity in most populations. There was a trend towards redder grain in the sprouting-resistant selections. The recovery of sprouting-resistant, early maturing segregants was relatively low, averaging less than 10% over the six populations. In conclusion, selection for increased sprouting resistance can result in delayed maturity, but the magnitude of that delay will vary among populations.     

Combining Ability and Heterosis for Quantitative Powdery Mildew Resistance in Barley

The estimation of combining abilities and heterosis for quantitative resistance against Erysiphe graminis f. sp. hordei of eight spring barley (Hordeum vulgare) cultivars is presented. Fur this purpose a half diallel cross and its parents were arranged in five Latin rectangles, each inoculated with a current mildew isolate. Significant general combining ability was found whereas specific combining ability was non-significant. A great pan of the general combining ability could be explained by variety effects. Significant variety heterosis was obtained too. Significant average heterosis was obtained but us effect was small. Among those selected for this study, ‘Grit’ and ‘Hora’ were the best parents for further crosses.     

Improved Culture System for Microspores of Barley to Become a Target for DNA Uptake

For the winter barley cultivar ‘Igri’, a microspore isolation and regeneration procedure is described which allows the production of such vigorous micro-spore fractions that this single-cell system can be used as a target for DNA uptake. Up to 60 % of the vigorous microspores isolated from the anther, and cultured in liquid modified MS medium, formed embryoids and/or calli. Such preparations were used for trials in DNA uptake with the plasmid pBI 221. Transformation trials were performed with polyethylene glycol as the inducing agent. With this treatment, a relative increase of fluorescence could be shown under UV light indicative of transient expression of the uidA gene.     

Responses in Agronomic Traits Associated with Three Recurrent Selection Regimes for Groat-Oil Content in Oat*

Crop strains with high values for the selected trait but unacceptable agronomic characteristics are not useful as varieties. In this study, we determined whether selection for groat-oil content in oat (Avena sativa L.) via three recurrent selection regimes caused associated changes in 10 agronomic traits. Random lines of each cycle of selection of each regime were evaluated in a replicated field experiment at two locations. Biomass, grain yield, groat yield, and test weight decreased in all three regimes. Seed weight and plant height decreased in two regimes, and groat fraction in one. Oil yield increased in two regimes and decreased in the other. Plants headed earlier in one and later in another regime over cycles of selection. No changes in harvest index occurred. Heritabilities and genetic variances were relatively high for all traits in each regime. Factor analysis of the phenotypic correlation matrices indicated that simultaneous increases in grain yield and in groat-oil content are possible, that continuous selection for groat-oil content decreases groat fraction and test weight, and that seed weight is strongly and negatively associated with groat-oil content. A new recurrent selection regime for increasing groat-oil content of oat that may overcome the negative responses of agronomic traits was proposed.     

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.