Net blotch resistance in wild species of Hordeum

Resistance to net blotch was evaluated in 175 Hordeum vulgare subsp. spontaneum (H. spontaneum) accessions and 149 accessions of thirteen species or subspecies of wild Hordeum. Most H. spontaneum accessions showed resistance to each of the four Pyrenophora teres f. teres (P. teres) isolates tested. However, H. spontaneum accessions showed different resistance reactions, depending upon their origin. In particular, some accessions from Afghanistan and Russia showed a high level of resistance, and accessions from Morocco were susceptible. Among the four P. teres isolates, the virulence spectra on the H. spontaneum accessions were more different between isolates from different countries than between those from the same country. Hordeum spontaneum accessions susceptible to the Canadian isolate WRS102, but resistant to the other three isolates were found in Iraq suggesting the geographical differentiation of resistance genes in H. spontaneum. All accessions of the other wild Hordeum species, especially some accessions of H. marinum subsp. gussoneanum, showed high levels of resistance. These resistance genes may be useful candidates for incorporation into cultivated barley. This revised version was published online in July 2006 with corrections to the Cover Date.     

Molecular characterization and sequence polymorphism of the alcohol dehydrogenase 1 gene in Hordeum vulgare L.

Partial sequences (appr. 900bp) spanning from exon 2 to 4 of the gene encoding alcohol dehydrogenase 1 were obtained from four varieties of cultivated barley, Hordeum vulgare ssp. vulgare, and from 21 accessions of its wild progenitor, H. vulgare ssp. spontaneum. Sequence diversity was low and restricted to the non-coding intron sequences. In the 648–660bp long intron 3 both a microsatellite locus and a major duplication (153/156bp) were found. Sequence diversity in H. vulgare ssp. spontaneum was slightly higher than in cultivated barley. In general the Adh1 gene displays only limited polymorphism and is not appropriate for seeking correlations between molecular and eco-geographical data. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic variation in barley of crossability with wheat and its quantitative trait loci analysis

To study genetic variation in crossability, 80 barley accessions of diverse geographic origin consisting of 50 wild barleys (H. vulgare ssp. spontaneum or ssp. agriocrithon) and 30 cultivated barleys (H. vulgare ssp. vulgare) were crossed as the male parent with a highly crossable wheat variety, Shinchunaga. Crossabilities, expressed as the percentage of pollinated florets giving embryo-containing caryopses, ranged from 0% to 68.6%. Barley accessions from East Asia had generally a low crossability, while barley accessions from other regions exhibited a wider range of crossability including highly crossable genotypes. No significant difference in mean crossability was found between wild and cultivated barleys. To estimate the number and location of barley genes controlling the crossability, doubled haploid lines derived from the cross between the barley varieties Steptoe and Morex were crossed as the male parent with wheat. Quantitative trait loci (QTL) analysis using molecular markers identified four QTL. These were mapped to the centromeric regions of chromosomes 2H, 3H and 5H and the short arm of chromosome 7H. The QTL on chromosomes 3H and 5H had larger effects than those on chromosomes 2H and 7H. The four QTL collectively explained 35.4% of the total variance under a multiple QTL model. Relationships of the QTL identified in the present study with previously reported crossability genes of barley and wheat are discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Inheritance of gramine content in barley

The inheritance of gramine indole alkaloid was studied in F1 and F2 generations derived from crosses between wild ( Hordeum vulgare subsp. spontaneum) and cultivated ( H. vulgare subsp. vulgare) barley at adult stage in the field. The means of gramine content were very low in F1s and F2s in April and May. The ratio of progeny containing gramine to those without gramine (near zero or undetectable) fit a mono-genic or di-genic model in F2 generation. However, the content of gramine might be controlled by minor genes in the gramine-containing plants. The direction of dominance was toward lower gramine content. The averages of dominance ( h/d) in F1 and F2 ranged from partial to complete. Broad-sense heritabilities were high (0.70 to 0.77). The usefulness of hybrids for breeding aphid resistance and possibility increasing palatability of grazing pasture for sheep and cattle is discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Detection of QTLs for salt tolerance in Asian barley (Hordeum vulgare L.) by association analysis with SNP markers

Two hundred ninety-six Asian barley (Hordeum vulgare L.) accessions were assessed to detect QTLs underlying salt tolerance by association analysis using a 384 single nucleotide polymorphism (SNP) marker system. The experiment was laid out at the seedling stage in a hydroponic solution under control and 250 mM NaCl solution with three replications of four plants each. Salt tolerance was assessed by leaf injury score (LIS) and salt tolerance indices (STIs) of the number of leaves (NL), shoot length (SL), root length (RL), shoot dry weight (SDW) and root dry weight (RDW). LIS was scored from 1 to 5 according to the severity of necrosis and chlorosis observed on leaves. There was a wide variation in salt tolerance among Asian barley accessions. LIS and STI (SDW) were the most suitable traits for screening salt tolerance. Association was estimated between markers and traits to detect QTLs for LIS and STI (SDW). Seven significant QTLs were located on chromosomes 1H (2 QTLs), 2H (2 QTLs), 3H (1 QTL), 4H (1 QTL) and 5H (1 QTL). Five QTLs were associated with LIS and 2 QTLs with STI (SDW). Two QTLs associated with LIS were newly identified on chromosomes 3H and 4H.     

Genetic Resources for Salt Tolerance in the Wild Progenitors of Wheat (Triticum dicoccoides) and Barley (Hordeum spontaneum) in Israel

Salt tolerance was tested in the progenitors of cultivated cereals, wild barley (Hordeum spontaneum) and wild emmer wheat (Triticum dicoccoides) from Israel. Plants from five selected populations of H. spontaneum from the Mediterranean Coastal Plain and northern Negev desert, were grown on 250 and 350 mM of NaCl. Likewise, five populations of T. dicoccoides from the eastern Samaria steppes, Mt. Hermon and Mt. Carmel, were grown on 175 and 250 mM of NaCl. Here we report on superior genotypes of H. spontaneum, ripening at 350 mM NaCl (= 60 % sea water), and of T. dicoccoides ripening at 250 mM (— 40 % sea water). We are proceeding now with both genetical and physiological studies aimed at chromosomally-locating salt tolerant genes and unravelling the mechanism(s) of salt resistance in these wild cereals.     

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.     

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.     

The Genetic Basis of Variation for Salt Tolerance in Sorghum bicolor (L.) Moench Seedlings

The genetic basis of salt (NaCll tolerance at the seedling stage was examined in eight accessions of Sorghum bicolor (L.) Moench which differed in relative NaCl tolerance. The eight accessions were crossed in all possible combinations. The Nad tolerances of the resulting 56 F: hybrids and their eight parents were assessed after two weeks growth in solution cultures containing 0, 103, and 150mM NaCl. Indices of salt tolerance (relative tolerance) were used for analysis using the diallel method. Both additive and dominance effects appeared to be important for the expression of variation under low and high salinity levels, but the effect of genes with dominance properties was more pronounced. Estimates of narrow sense hentabilities for sail tolerance were appreciable, suggesting that rapid improvement in salinity tolerance is possible in this species using high selection pressures in the F₂ population.     

Effects of Epichloë endophyte infection on growth,physiological properties and seed germination of wild barley under saline conditions

Wild barley (Hordeum brevisubulatum) is a grass that inhabits alkalized meadows in northern China. An asexual Epichloë bromicola endophyte was detected in seeds and leaf sheaths in all wild barley samples from Gansu Province, China. In this research, we determined the effects of the E. bromicola endophyte on growth, physiological properties and seed germination of wild barley under salt stress through a set of experiments. Our results demonstrate that endophyte-infected (E+) plants produced more tillers, higher biomass and yield, higher chlorophyll content and superoxide dismutase activity than endophyte-free (E−) plants under high salt stress. Seed germination parameters of E+ biotype were significantly higher than those of E− plants when NaCl concentration reached 200 and 300 mM. Our results demonstrate that E. bromicola endophytes increased tolerance to salt stress in wild barley by increasing seed germination and growth, and altering plant physiology.     

Evidence for common genetic mechanisms controlling the tolerance of sudden salt stress in the tribe Triticeae

Previous studies in several Triticeae species have suggested that salt tolerance is a polygenic trait, but that genes on some chromosomes confer better tolerance to salt stress than others. This suggests an intriguing possibility that there may be a similar basis for salt tolerance in the species of the tribe Triticeae. In this study, chromosomal control of the tolerance to sudden salt stress, measured as the mean rate of leaf elongation in solution cultures with a single increment of 200 mM NaCl, was investigated in the genomes of cultivated barley (Hordeum vulgare L.), rye (Secale cereale L.), and Dasypyrum villosum (L.) Can-dargy by using disomic addition lines of individual pairs of chromosomes or chromosome arms of each of the three species in the ‘Chinese Spring’ wheat genetic background. It was observed that the chromosomes of homoeologous groups 3, 4, and 5 in barley, 5 and 7 in rye, and 4 and 6 in D. villosum carry loci with significant positive effects on salt tolerance. Increased doses of chromosomes of group 2, however, reduce or do not increase the tolerance to salt stress. These results are in agreement with a previous study of the tolerance of this salt stress regime in wheat and wheatgrass Lophopyrum elongatum. A ranking analysis of the chromosomal effects within each genome of the five Triticeae species investigated in this and previous studies revealed that the chromosomes of homoeologous groups 3 and 5 consistently confer large positive effects on the tolerance of sudden salt stress, while the chromosomes of homoeologous group 2 in increased dose have no or negative effects on the tolerance. This strongly suggests that species of the tribe Triticeae share some common genetic mechanisms of tolerance of sudden salt stress. The findings in this study give credence to the proposal that wild relatives can be exploited in the development of wheat cultivars with greater tolerance to salt stress.     

Variation in developmental patterns of wild barley (Hordeum spontaneum L.) and cultivated barley (H. vulgare L.)

Summary Six wild barley (Hordeum spontaneum) accessions, from a diverse range of habitats, and two spring-cultivated barleys, were examined for variation in durations of development phases. The durations of the leaf initiation and spikelet initiation phases were longer and spikelet growth phases shorter, in wild than in cultivated barley. Across all wild and cultivated barleys the rate and duration of spikelet initiation were negatively correlated, but neither was related to the number of spikelets per spike. The number of spikelets was positively correlated with the number of leaves and the ratio of the number of spikelets to the number of leaves declined with increasing time to anthesis, indicating that each successive leaf was associated with a diminishing increase in the number of spikelets. The duration of culm elongation and final culm length were shorter in accessions of cultivated barley compared with wild barley. This paper also discusses the feasibility for increasing the number of spikelets per spike through breeding for genetic changes in lengths of pre-anthesis phases of development.Abbreviations ANOVA Analysis of variance - HV Hordeum vulgare - CE Culm elongation - DR Double ridge - HS Hordeum spontaneu - ° Cd Degree days     

Genotypic variation in polyphenol content of barley grain

The polyphenol content in pearl barley, which is highly correlated to a browning reaction after heat treatment, was investigated using 1,347 cultivated barley varieties (H. vulgare) and two wild accessions (H. vulgare subsp. spontaneum) collected from different areas of the world. The polyphenol content in the cultivated barley shows a wide variation ranging from 0.19 to 0.75 mg/g with a nearly normal frequency distribution. The polyphenol content in the hulless varieties from Japan and Korea was low. On the other hand, the polyphenol content in wild barley was about two times higher than the average value recorded in cultivated barley. Based on HPLC analysis, five lowest-polyphenol content local varieties do not represent proanthocyanidin-free mutants. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genetic diversity and population structure of wild and cultivated barley from West Asia and North Africa

Population structure and relationships within and among 185 accessions of wild (Hordeum vulgare ssp. spontaneum) and cultivated barley (H. v. ssp. vulgare) from five countries in the West Asia and North Africa (WANA) region were studied using 36 simple sequence repeat (SSR) markers. The accessions were divided into subspecies/origin (S/O)‐groups and marker results were analysed in relation to genetic diversity and genetic structure. Wild barley from WANA was found to be highly diverse. The landraces from different countries of the Near East showed genetic diversity that was nearly as high as the wild barley from the same country. Further analyses showed that wild barley from Palestine/Israel represented the group with the highest diversity and the most complex structure. However, this group was distantly related to the cultivated barley in WANA, while the wild barley from the rest of the WANA region was closely related to the cultivated WANA barley. The high diversity and the close relationship to the wild barley make the WANA landraces an interesting genetic resource for both conservation and exploitation.     

Growth and Development of Wild Oat and Dual-Purpose Barley Seedlings

Wild oat ( Avena fatua L.) and dual-purpose barley ( Hordeum spontaneum L.) are native species that grow in many parts of Jordan. We compared seedling growth and development of wild oat and dual-purpose barley in pots under field conditions. The seedling fresh root volume (cm³), root oven-dry weight (g), shoot oven-dry weight (g) and dry root weight to dry shoot weight ratio of both species were evaluated at ages 20, 40, 60, 80 and 100 days after emergence (DAE). Seedlings of the two species at 20 and 40 DAE had similar fresh root volumes and weights. At 60 DAE, the root volumes of barley and oat increased to 2- and 3-fold the volumes at 40 DAE, respectively. By 80 DAE, root volume had increased to 13.6 and 10.2 cm³ for wild oat and dual-purpose barley, respectively. At 100 DAE, the root weight of dual-purpose barley significantly (P ≤ 0.01) decreased to 1.1 g, whereas wild oat root weight did not change significantly. Wild oat showed a slightly higher root/shoot ratio at 20 DAE than dual-purpose barley (0.88 for wild oat compared to 0.70 for dual-purpose barley). At 60 and 80 DAE, both species showed a reduction in root/shoot ratio, and this reduction increased at 100 DAE (0.46 and 0.33 for wild oat and dual-purpose barley, respectively). Our data indicate that the growth and development of the two grass species follow similar patterns.     

Geographical differentiation and diallel analysis of seed dormancy in barley

Seed dormancy is one of the most important parameters affecting the malting process and pre-harvest sprouting in barley (Hordeum vulgare L.). Variation of seed dormancy in 4365 cultivated and 177 wild barley (ssp. spontaneum) accessions derived from different regions of the world was investigated in Okayama University, Kurashiki, Japan. Seed dormancy of each accession was estimated from their germination percentages at 0, 5, 10 and 15 weeks post-harvest after-ripening periods. All of the wild barley accessions showed less than 10% germination at 0 week after-ripening period. Level of seed dormancy in 4365 cultivated barley accessions showed a clear geographical differentiation. Seventy seven percent of Ethiopian accessions showed high germination percentages, while 86% of Japanese, Turkish and North African accessions showed low germination percentages at 0 week after-ripening period. A half diallel cross using eleven barley accessions with different level of dormancy revealed that seed dormancy was predominately controlled by additive gene effects. These results suggest that large genetic diversity for seed dormancy in barley is explained as different levels of additive accumulation of genetic factors. Barley varieties showing appropriate dormancy could be developed by crossing among barley germplasm accessions used in the present study.     

福建省野生大豆耐盐种质的筛选

盐害是一种重要的非生物胁迫之一,限制植物的生长并降低作物的产量和品质,野生大豆是栽培大豆的近缘野生种,含有诸多优异的基因。为评价野生大豆的耐盐性,试验采用漂浮育苗方法,以NaCl致死浓度为评价指标,对244 份野生大豆苗期的耐盐性进行鉴定。结果表明野生大豆种质致死浓度最低为200 mmol/L,最高为300 mmol/L,并从中筛选出致死浓度为290 mmol/L 耐盐种质14 份,占供试材料的6%,致死浓度为300 mmol/L 的耐盐种质2 份,占供试材料的1%;致死浓度为200 mmol/L 的盐敏感种质15 份,占供试材料的6%;致死浓度为270 mmol/L的材料最多,共计36 份,占供试材料的15%,其次是致死浓度为220 mmol/L 的材料32 份,致死浓度为280 mmol/L 的材料29 份。综上表明野生大豆种质资源耐盐性差异较大,含有丰富的遗传变异,有望筛选出优异的耐盐种质,用于耐盐QTL的定位和耐盐机理的研究。     

Intra-specific Variation for Salt Tolerance in Linseed (Linum usitatissimum L.)

Thirty-six accessions of linseed ( Linum usitatissimum L.) were screened for salt tolerance at the seedling stage after 2 weeks growth in solution culture salinized with 150mol m⁻³ NaCL Considerable variation in salt tolerance was observed in this set of germplasm. Two salt-tolerant accessions, T-5(69-S9)B and LS-2 and two salt-sensitive accessions P-70 and Aver-Rer-Seic, selected at the seedling stage, were grown to maturity in sand culture salinized with 0 (control), 70, 140, and 210 mol m⁻³ NaCl. Both the salt-tolerant accessions exceeded the two salt-sensitive accessions in biomass production, seed yield, and yield components, but the accessions did not differ significantly in seed oil content in the salt treatments. The tolerant accessions accumulated greater amounts of Na⁺ in the shoots and less in the roots compared with the salt-sensitive accessions. The shoot K⁺, K: Na ratios, and K⁺ versus Na⁺ selectivity were lower in the salt-tolerant accessions compared with the salt-sensitive accessions; the reverse was true for their roots. Contributions of Na⁺ and Ca²⁺ to osmotic adjustment were much higher in the two salt-tolerant accessions than in the salt-sensitive ones.
It has been established that there is a great deal of variation for salt tolerance in linseed. The degree of salt tolerance of this crop does not vary during ontogeny. The physiological basis of salt tolerance of this species appears to be associated with the inclusion of Na⁺ in the shoots.     

大豆出苗期耐盐性鉴定方法建立及耐盐种质筛选

土壤盐渍化是影响农业生产的重要问题,筛选耐盐大豆资源对于大豆主产区盐渍化土壤的利用具有重要意义。以中黄35、中黄39、Williams82、铁丰8号、Peking和NY27-38为供试材料,以蛭石为培养基质,设0、100和150 mmol L?1 NaCl 3个处理,进行出苗期耐盐性鉴定,分析与生长相关的6个指标,旨在明确大豆出苗期耐盐性鉴定指标和评价方法。结果表明, 150 mmol L?1NaCl处理显著降低大豆的成苗率、株高、地上部鲜重、根鲜重、地上部干重和根干重,并且不同材料间差异显著。基于幼苗生长发育状况的耐盐指数方法与耐盐系数方法对6份种质耐盐性评价结果显著相关。耐盐指数法对植株无损坏、可省略种植对照,节约人力和物力,提高种质鉴定的效率。因此,以150 mmol L?1 NaCl作为出苗期耐盐鉴定浓度,以耐盐指数作为大豆出苗期耐盐鉴定评价指标,鉴定27份大豆资源,获得出苗期高度耐盐大豆(1级) 3份、耐盐大豆(2级) 7份,其中4份苗期也高度耐盐(1级),分别为运豆101、郑1311、皖宿1015和铁丰8号。本研究建立了一种以蛭石为基质,利用150 mmol L?1 NaCl处理,以耐盐指数作为评价指标的大豆出苗期耐盐性鉴定评价的简便方法,并筛选出4份出苗期和苗期均耐盐的大豆,对耐盐大豆种质资源的高效鉴定和耐盐大豆新品种培育具有重要意义。     

The resistance to leaf rust and powdery mildew of recombinant lines of barley (Hordeum vulgare L.) derived from H. vulgare × H. bulbosum crosses

A set of 23 recombinant lines (RLs) of barley ( Hordeum vulgare L.) derived from H. vulgare  ×  H. bulbosum L. crosses was inoculated with barley leaf rust ( Puccinia hordei ) and powdery mildew ( Blumeria graminis f.sp. hordei ) at the seedling stage to identify their levels and mechanisms of resistance. Eight RLs were studied further in glasshouse and field tests. All three barley parents ('Emir', 'Golden Promise' and 'Vada') were highly susceptible to powdery mildew and leaf rust isolates. Several RLs showed partial resistance expressed as high relative latency periods and low relative infection frequencies against leaf rust. This high level of partial resistance was due to a very high level of early aborting colonies without host cell necrosis. Several RLs showed hypersensitive resistance to some or all isolates. For powdery mildew, one RL was completely resistant to the CC1 isolate and had a hypersensitive resistance to the CO-02 isolate. Three RLs derived from 'Emir' were completely resistant to both powdery mildew isolates, and three more RLs tested in the field had higher levels of partial resistance than their parents. The results indicate that H. bulbosum contains major and minor gene(s) for resistance to leaf rust and powdery mildew that can be transferred to cultivated barley.