Variation in response to infection with Erysiphe graminis hordei and Puccinia hordei in some wild barley populations in a centre of diversity

Summary The Near East Fertile Crescent extending from Iran to Israel is the centre of origin of cultivated barley and a region of great genetic diversity in wild barley, Hordeum spontaneum C. Koch (syn. H. vulgare ssp. spontaneum (C. Koch Thell)). Wild barley accessions collected from different parts of this region were evaluated for their reactions to infection with three isolates of Erysiphe graminis hordei and two of Puccinia hordei. One culture of each pathogen was isolated in Israel and the others, either in Japan or the United States. Out of a total of 330 wild barley accessions collected from 14 sites in Iran, Turkey and Syria, only 18.8% were resistant to the Israeli culture, and 14.8% were resistant to a composite of the Japanese and American cultures of E. graminis hordei. Out of 105 accessions collected from six sites in Iran and Turkey, none was found to be resistant to the Israeli culture and 34.3% were resistant to the American culture of P. hordei. Considerable variation was observed both within and among sites for reactions to infection with different cultures of each of the two pathogens. The results of this study were compared with those of an earlier study involving wild barley accessions from Israel to illustrate the relative importance of different subregions in the Near East Fertile Crescent as sources of new genes for resistance to E. graminis hordei and P. hordei. Implications of these studies for in situ conservation of genetic diversity in wild barley are discussed.     

Resistance of Triticum dicoccoides to infection with Erysiphe graminis tritici

Summary The reactions of 233 Triticum dicoccoides acessions, collected at 10 sites in Israel and elsewhere, to infection with cultures of Erysiphe graminis tritici, were determined. The reactions indicated that the number of sources of resistance to E. graminis tritici which can be obtained from T. dicoccoides plants growing wild in Israel and elsewhere is almost unlimited. One hundred and fourteen or 49% of the accessions were resistant, and 137 or 59% of the accessions were resistant or moderately resistant to infection with four cultures of E. graminis tritici which possess the virulence genes corresponding to most of the identified resistance genes in wheat. Accessions collected at sites with marginal habitats where T. dicoccoides grows poorly and has lower grain weight, were more susceptible than were accessions collected at sites with an optimal habitat for growth of T. dicoccoides. The results agreed with those in a previous study with Hordeum spontaneum, and indicate that to obtain H. spontaneum or T. dicoccoides accessions with the highest level of resistance to the powdery mildew pathogens, plants should be collected at sites in ecological and geographic regions where those two species occupy optimum habitats and are exposed to the powdery mildew pathogens.     

Identification of Resistance Genes Against Powdery Mildew in Four Accessions of Hordeum Vulgare SSP. Spontaneum

Summary Four newly detected accessions of wild barley (Hordeum vulgare ssp. spontaneum) resistant to powdery mildew caused by Blumeria graminis f. sp. hordei were studied with the aim of finding the number of genes/loci conferring the resistance of individual accessions, the type of inheritance of the genes and their relationships to the Mla locus. F₂ populations after crosses between the winter variety ‘Tiffany’ and four wild barley accessions and use of microsatellite DNA markers were focused on the identification of individual resistance genes/loci by means of their chromosomal locations. In PI466495, one locus conferring powdery mildew resistance was identified in highly significant linkage with the marker Bmac0213. This location is consistent with the known locus Mla on chromosome 1HS. In the other three accessions the resistance was determined by two independent loci. In PI466197, PI466297 and PI466461, one locus was identified on chromosome 1HS and three new loci were revealed on chromosomes 2HS (highly significant linkage with Bmac0134), 7HS (highly significant linkage with Bmag0021) and 7HL (significant linkage with EBmac0755). Our prospective aim is identification of further linked DNA markers and the exact location of the resistance genes on the barley chromosomes.     

Sources of Resistance to Powdery Mildew in Barley Lines Derived from Hordeum spontaneum Collected in Israel

Fourty two barley lines direved from the F₇ of crosses between barley cultivars and different accessions of Hordeum spontaneum collected in Israel and 30 lines or varieties with known genes for resistance to powdery mildew were included m this study. Eleven European and three Israeli powdery mildew cultures, possessing virulence genes corresponding to known resistance genes, were used to make comparisons between the varieties with known resistance genes and H. spontaneum derived lines. The reaction pattern of 39 H. spontaneum derived lines was clearly different from the reaction pattern o; any of the known genes for mildew resistance included in this study. Only two cases were observed in which the reaction pattern of H. spontaneum derived lines agreed with reaction patterns of known genes for mildew resistance viz. Ml-a9 and Ml-p. Trie Mildew resistance of one line apparently traces back to uncontrolled outcrossing with a Ml-a.6+Ml-g resistant cultivar. Since the majority of the 42 host genotypes tested showed distinctive variation in resistant reaction types against different mildew cultures, this study docs not support the assumption that differences in resistant infection types against distinct mildew cultures are sufficient to indicate the presence of supplementary genes for resistance in a given genotype of the host. The results justify the conclusion that the natural population of H. spontaneum in Israel forms a large gene pool for mildew resistance which is not yet used m cultivated barley.     

Resistance to Rhopalosiphum padi (Homoptera: Aphididae) in Hordeum vulgare subsp. spontaneum and in hybrids with H. vulgare subsp. vulgare

Summary Accessions of Hordeum vulgare subsp. spontaneum, the progenitor of cultivated barley, were screened in field and glasshouse trials for resistance to the aphid Rhopalosiphum padi. A few selected lines were furthermore hybridized with a modern barley variety and the resulting populations evaluated. High levels of resistance were found among some of the spontaneum accessions resulting in lower aphid growth rates (maximum reduction 57%). Segregation patterns among siblings in F2 populations were continuous, indicating the presence of several genes with possibly additive effects. The usefulness of H. vulgare subsp. spontaneum for breeding aphid resistant barley is discussed.     

Genes for scald resistance from wild barley (Hordeum vulgare ssp spontaneum) and their linkage to isozyme markers

Summary Accessions of Hordeum vulgare ssp. spontaneum, the wild progenitor of barley, collected in Israel (70), Iran (15) and Turkey (6) were screened for seedling response to four isolates of Rhynchosporium secalis, the pathogen causing leaf scald in barley. Resistance was very common in the collection (77%) particularly among accessions from the more mesic sites (90%). The genetics of this resistance were investigated in fifteen backcross (BC₃) lines that contained an isozyme variant from H.v. ssp. spontaneum in a H.v. ssp. vulgare (cv. Clipper) background and were resistant to scald. Segregation in the BC₃F₂ families conformed with a single dominant resistance gene in 9 of the 15 lines. Scald resistance and the isozyme marker were closely linked in three of the BC₃-lines, loosely linked in four and unlinked in the remaining eight. Scald resistance genes were identified on barley chromosomes 1, 3, 4 and 6. Crosses between several of the scald resistant BC-lines together with the linkage data indicated that at least five genetically independent resistances are available for combining together for deployment in barley. The linkage of scald resistance in several BC₃-lines to the isozyme locus Acp2 is of special interest as this locus is highly polymorphic in wild barley.     

A new virulent strain of Puccinia hordei

Summary Four cultures of Puccinia hordei isolated from the alternate Ornithogalum hosts in Israel were virulent on seedlings of barley cultivars carrying resistance genes Pa, Pa2, Pa2 + Pa5, Pa2 + Pa6, Pa3, Pa4, and Pa7. Cultures with such a spectrum of virulence have never been reported. The cultures remained stable through 20 successive inoculation and isolation cycles. One of the cultures, T-40SS, retained its specific virulence after selfing on the alternate host. The evolution of virulent races of obligate parasites in the centers of origin of their hosts is discussed.     

Avoidance of leaf rust fungi in wild relatives of cultivated cereals

Summary Avoidance of rust fungi that was based on poor appressorium induction was previously found in Hordeum chilense. In the present study 95 accessions of Triticeae were screened for avoidance of Puccinia hordei. The percentage of appressorium formation per germinated spore ranged from 6 to 90%. On none of the 41 accessions of Aegilops, Agropyron, Elymus, Secale, Thinopyrum or Triticum studied was the rate of appressorium formation lower than 25%. Lower rates of appressorium formation were, however, found on accessions of wild barley species Hordeum brachyantherum, H. marinum, H. parodii and H. secalinum. Its implications in cereal breeding are discussed.     

The origin of six-rowed ‘wild’ barley from the western Himalaya

Summary Plants were grown from seed of two-rowed wild barley, Hordeum spontaneum, and six-rowed brittlerachised barley. H. agriocrithon, collected in Ladakh, north-western India. Whereas the H. spontaneum remained true to type, segregation for morphological characters was observed in progeny rows grown from heads of H. agriocrithon plants. The H. agriocrithon heads also showed segregation for a biochemical character, the polypeptide pattern of the endosperm storage protein fraction (hordein). The H. agriocrithon seed therefore originated from natural hybridization between cultivated H. vulgare and weedy H. spontaneum. Crosses of H. vulgare and H. spontaneum gave progeny which resembled H. agriocrithon and showed similar hordein polypeptide segregation patterns. The results indicate that six-rowed brittle-rach ised barleys from the Himalayas have a similar origin to forms found in the Middle East, and that H. agriocrithon does not play a direct role in the evolution of barley.     

Variation in utilization efficiency and tolerance to reduced water and nitrogen supply among wild and cultivated barleys

Broad genotypic variation in the response to low soilmoisture and reduced nitrogen supply was found amongthe wild Hordeum spontaneum accessions and thelandraces and modern cultivars of H. vulgare ofdifferent geographic origin. Measurements at the endof vegetative growth in plants grown in soil culturesrevealed genotypically specific responses to the usedenvironmental factors. Cultivars and breeding linesfrom Syria and Ethiopian landraces combined bothdrought resistance and tolerance to low nitrogen. TheSyrian barleys were also distinguished by a highnitrogen utilization efficiency (NUE) under low Nnutrition. European cultivars indicated a pooradaptation to N shortage, but some of them wereresistant to soil drought. No stress resistant barleyswere found among the wild accessions and Sardinianlandraces. Genotypic differences in the relativevalues of NUE and water use efficiency were associatedwith low N-tolerance. Some Syrian selections,Ethiopian landraces and the modern German cv. Maresiwere found to be most drought resistant. Maintenanceof a relatively high photosynthetic activity of theuppermost leaves was associated with droughtresistance. As far as concerning with the vegetativegrowth phase, the modern Syrian germplasm andEthiopian landraces may be recommended as donors ofadaptative characters for local barley breeding.     

Genetic analysis of powdery-mildew (Erysiphe graminis f.sp. hordei) resistance derived from wild barley (Hordeum vulgare ssp. spontaneum)

The inheritance of resistance to powdery mildew was investigated in 20 accessions of Hordeum spontaneum and in 20 F₄ lines derived from crosses between the variety ‘Aramir’ and 13 accessions of H. spontaneum. Two resistance genes were detected in 17 accessions, and three resistance genes in one accession. In two accessions, only one resistance gene was present. The 20 breeding lines showed a large variation in infection type and infection level. The genetic relationship between the resistance genes detected was investigated in the seven most resistant F₄ lines. These F₄ lines were divided into three groups which carried different resistance genes. In two lines, the detected resistance gene was shown to be race-specific.     

Exotic barley germplasm: variation and effects on agronomic traits in complex crosses

Summary Utilization of exotic germplasm offers an approach to broaden genetic variability in breeding populations. This study was conducted in order to 1) compare germplasm of exotic origin with adapted Swedish barleys with respect to genetic differences and 2) to evaluate how exotic material affected agronomic performance in complex crosses. Allozyme studies showed the following Nei's gene diversities among parents: 0.13 (adapted parents), 0.16 (landraces) and 0.25 (H. spontaneum). Cluster analysis indicated that parental groups were genetically divergent. Earliness, straw length, number of ears per plant and thousand kernel weight (TKW) were studied. Variation in agronomic traits showed the following pattern: landraces > H. spontaneum > adapted lines. The best sources for earliness were adapted parents and landraces. Mean straw length was greatest in H. spontaneum lines. Number of ears per plant was quite similar in all groups. The highest TKW was among landraces and adapted parents. Hybrids from the complex crossing programme exceeded parents in earliness and TKW. An index composed from the four traits showed the most favorable frequency distributions for adapted parents and hybrids. Both genetic and agronomic studies indicate that new valuable variation from exotic germplasm may be introduced into barley breeding material.     

Variation in vegetative parameters related to the nitrogen economy of wild barley,Hordeum spontaneum,in Israel

Summary Hordeum spontaneum, the wild progenitor of cultivated barley, has previously been examined in various studies as a germplasm resource in breeding for grain protein content and related nutritional traits.The nitrogen content and dry weight of leaf and stem (stem plus sheath) at anthesis, and the final grain size and grain protein content were measured in 33 H. spontaneum and two H. vulgare genotypes. H. spontaneum was generally higher in nitrogen content of leaves and stems, but lower in dry weight at anthesis. Consistent with previous reports, the H. spontaneum genotypes were considerably higher in grain protein than the cultivars. There was wide variation between and within populations of H. spontaneum suggesting that for breeding purposes lines combining high vegetative nitrogen content, dry weight and grain protein content can be selected.Incumbent of the Seagram Chair for Plant Science     

Transgressive segregation in inter and intraspecific crosses of barley

Summary Six populations of F₂-derived lines of barley and their parents were evaluated for heading date, plant height, grain yield, bundle weight, and harvest index in a replicated experiment in the field. These data were used to estimate the minimum number of effective factor pairs segregating for each trait, the number of favorable factors contributed by each parent in a cross, and the frequencies and magnitudes of transgressive segregates.Heading date, plant height, and harvest index were controlled by three to four effective factor pairs, whereas grain yield and bundle weight were controlled by five or more.All three H. spontaneum strains used in our study contributed one or more useful genes for each of the traits, grain yield, heading date, plant height, bundle weight, and harvest index. Therefore, it seems that H. spontaneum can be a useful source of favorable genes for quantitative traits, especially for grain yield, which could be incorporated into barley varieties readily by backcrossing. Transgressive segregates for grain yield in the interspecific crosses may provide the basic materials for improving the productivity of cultivated barley varieties.Journal Paper No. J-9760 of the Iowa Agric. and Home Econ. Exp. Stn., Ames, IA 50011. Project 2227. Supported in part by a grant from the Science and Education Administration-Cooperative Research, United States Department of Agriculture.     

Two-rowed and six-rowed wild barley from the western Himalaya

Summary Plants of seeds of Hordeum spontaneum and H. agriocrithon were collected from eight villages in Ladakh, north western India. This is the furthest east that H. spontaneum has been reported. It is the most westerly occurrence and largest find of H. agriocrithon in the Tibetan and Himalayan region. It is possible that H. agriocrithon occurs in Ladakh, not as a hybrid swarm resulting from a cross between H. spontaneum and sin-rowed cultivated barley, but as a pure population.     

The importance of abortive stoma penetration in the partial resistance and nonhost reaction of adult barley and wheat plants to leaf rust fungi

Summary Early stages of the infection process of Puccinia hordei isolate 1.2.1 and of a P. recondita f.sp. tritici isolate were studied on adult plants of four barley lines and one wheat line. Two of the barley lines are extremely susceptible to P. hordei, the other two have a very high level of partial resistance.A histological study based on a trypan blue staining indicated that stoma penetration by P. hordei isolate 1.2.1 was equally successful on the susceptible as on the partially resistant adult barley plants. Abortion of substomatal vesicles was rare in all lines. These results do not support a hypothesis that mechanisms of partial resistance in adult plants differ from those in seedlings by a substantial abortive stoma penetration.Also in the nonhost combinations wheat-P. hordei and barley-P. recondita f.sp. tritici inhibition of stoma penetration and of substomatal vesicle development appears to play a biologically insignificant role in adult plants.The proportion of stoma penetration on the leaf sheaths of two of the barley lines was as high as on the leaf blades of the flag leaf and the leaf below the flag leaf. There was no evidence for stomatal exclusion as a crucial factor in the relatively low infectibility of leaf sheaths to leaf-blade specialized rust species.     

Analysis of the yield potential and yield components of F1 and F2 hybrids of crosses between wheat (Triticum aestivum L.) and spelt (Triticum spelta L.)

Summary Crosses of wild barley (Hordeum vulgare ssp. spontaneum and Hordeum vulgare ssp. agriocrithon) with Hordeum vulgare ssp. vulgare were used to select high yielding grain types under dryland Mediterranean conditions. No special difficulties were faced in making the crosses, in eliminating the brittle rachis genes from the grain types or in selecting 6-rowed types in crosses between 2-rowed wild barley and 6-rowed ssp. vulgare varieties. Brittle rachis genotypes, present in the segregating populations were used in developing self-reseeding permanent pastures for dry areas. The best selections were tested in seven trials during 1989–92 and some of them outyielded their parents and also the best improved check variety by 13–22%. Indications for transgressive segregation were obtained for grain yield, straw yield, total biological yield, harvest index and volume weight. The crude protein content of some of the selections was significantly higher than that of the checks. For breeding programs aiming at large seeds, special ssp. spontaneum lines should be used as parents. High grain yield was positively correlated with high straw yield, total biological yield, earliness in heading date, high harvest index and negatively with volume weight. It was concluded that unexploited useful genes, even when not directly observed in wild barley, could be transfered easily into high yielding genotypes by breeding.     

Spectrum of resistance conferred by ML-O powdery mildew resistance genes in barley

Summary Ten barley mutants and five Ethiopian barley lines representing 11 independently arisen powdery mildew resistance genes in the ml-o locus were tested at the seedling stage to cultures of the powdery mildew fungus from Europe, Israel, USA. Canada, and Japan. They were resistant with infection type 0/(4) in all tests. They were also resistant to field populations of the pathogen when scored in disease nurseries at more than 78 locations in 29 countries in Europe, the Near East, North and South America. New Zealand, and Japan. This indicates that the 11 genes confer the same, world-wide spectrum of powdery mildew resistance. They have no effect on several other barley diseases such as stripe rust and leaf rust.Part of the research reported here was carried out under IAEA Research Agreement No 1043 and Research Contract No 139-74-1 BIO DK with the European Atomic Energy Community.     

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     

New molecular markers linked to qualitative and quantitative powdery mildew and scald resistance genes in barley for dry areas

A partial genetic linkage map was constructed on 71 doubled-haploid lines derived from a cross between the barley lines Tadmor and WI2291 with 181 molecular markers. The segregating population was used to detect markers linked to the gene Mlg conferring resistance to powdery mildew (Erysiphe graminis f. sp. hordei) and to genes for quantitative resistance to scald (Rhynchosporium secalis). The gene Mlg on chromosome 4H was flanked by two AFLP markers at a distance of 2.0 and 2.4 cM, respectively. QTLs for resistance to scald were detected on chromosomes 2H and 3H. This association of molecular markers with qualitative and quantitative disease resistance loci represents a valuable starting-point for marker-assisted selection. This revised version was published online in July 2006 with corrections to the Cover Date.