Reactions of Hordeum spontaneum to infection with two cultures of Puccinia hordei from Israel and United States

Summary The reactions to infection with two cultures of Puccinia hordei were determined for 292 Hordeum spontaneum (syn. H. vulgare ssp. spontaneum) accessions, collected at 16 sites which encompassed the ecological range of H. spontaneum in Israel. Culture Tel-Aviv was from Israel and culture 57.19 was from the United States. Fifty-two percent of the accessions were resistant to culture Tel-Aviv and 67% were resistant to culture 57.19. Forty-three percent of the accessions were resistant to both cultures. The average infection type (IT) of accessions within sites ranged from 2.7 to 7.5 on a 0–9 rating scale. The results showed that the presence of Ornithogalum species, the alternate hosts of P. hordei, may increase the percentage of H. spontaneoum accessions resistant to P. hordei. More accessions were resistant at sites where humidity at 1400 was higher, the annual evaporation was lower, and where the glumes were shorter. Kernel weight and annual rainfall was not correlated with resistance. A lower percentage of H. spontaneum accessions were resistant to P. hordei culture Tel-Aviv from Israel than to culture 57.19 from the United States. In a previous study a lower percentage of H. spontaneum accessions also was found to be resistant to a culture of Erysiphe graminis hordei from Israel than to cultures from other countries. Previous studies also have shown that cultures of P. hordei and E. graminis hordei from Israel have many genes for virulence on barley, and that H. spontaneum accessions from Israel have many genes for resistance to these two pathogens. Previous results and the results reported in this paper support the hypothesis of coevolution of resistant host genes and virulent pathogen genes where hosts and pathogens have coexisted for many thousand years.     

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.     

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.     

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.     

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.     

Cytoplasmic male sterility in barley. III. Maintenance of sterility and restoration of fertility in the msml cytoplasm

Summary 58 varieties, strains or mutants of barley were tested in the msml cytoplasm in Finland, location ca. 61 N. Three (5%) were found to be partial restorers, the rest being maintainers of sterility. 31 Israeli strains of wild barley (ssp. spontaneum) were tested. About one third were maintainers of sterility, the majority partial restorers, and two strains were full restorers. There is probably polymorphism of the restorer gene in Israeli wild barley. Partial restoration displays an environmental response. The physiology of partial restoration 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.     

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.     

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.     

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.     

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.     

Diversity and geographical distribution of seed lipoxygenase‐1 thermostability types in barley

To understand the diversity in the thermostability of the seed lipoxygenase‐1 (LOX‐1), 1040 cultivars of worldwide barley (Hordeum vulgare ssp. vulgare) genetic resources were investigated. The relative thermostability of LOX‐1 (LOX‐RTS) in these lines showed a bimodal frequency distribution and these lines were categorized into the high and low thermostability types (H‐type and L‐type, respectively). The H‐type lines predominated in the wild progenitor, ssp. spontaneum. The geographical distribution of these types in the cultivars was surveyed. The frequencies of the H‐ and L‐types were almost equal to one another in southwestern Asia. The occurrence of the H‐type predominated in eastern Asia and Africa, whereas in Europe and Turkey, the L‐type did. The predominance of the L‐type in Europe and Turkey can be understood through the hypothesis that the Fertile Crescent domestication contributed the majority of diversity in Europe. The uneven geographical distribution of the LOX‐1 thermostability types in the cultivars may reflect a polyphyletic origin of barley.     

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.     

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     

Exotic barley germplasms in breeding for resistance to soil-borne viruses

Summary Soil-borne mosaic inducing viruses, i.e., barley mild mosaic virus (BaMMV), barley yellow mosaic virus (BaYMV), and BaYMV-2, cause one of the most important diseases of winter barley in Western Europe. Since resistance of all commercial European barley cultivars is due to a single recessive gene (ym4) which is not effective against BaYMV-2, exotic barley germplasms (Hordeum vulgare L., H. spontaneum Koch) were screened for resistance to the different viruses and analyzed for genetic diversity concerning BaMMV resistance. In these studies it turned out that resistance to BaMMV is entirely inherited recessively and that a high degree of genetic diversity concerning resistance is present within the barley gene pool at least to BaMMV. Therefore, exotic barley germplasms are a very useful source for the incorporation of different resistance genes into barley breeding lines, thereby enabling the breeder to create cultivars adapted to cultivation in the growing area of fields infested by soil-borne viruses. Furthermore, in order to obtain more information on these germplasms they were evaluated for agronomic traits and isozyme, RFLP and RAPD analyses were carried out on these varieties to detect markers linked to the respective resistance genes and to obtain information on the genetic similarity between yellow mosaic resistant barley accessions derived from different parts of the world. Actual results of these studies are briefly reviewed.     

Resistance to powdery mildew in wild emmer (Triticum dicoccoides Körn.)

Summary Wild emmer from 73 collection sites, including 107 accessions from Israel, two from Lebanon and one from Turkey, were evaluated for resistance to powdery mildew in field nurseries in Israel and the Netherlands.The wild emmer entries displayed a diversity of responses to powdery mildew infection, ranging from high resistance to complete susceptibility. Most entries were resistant in at least one of the nurseries; several entries proved to be resistant in all the tests.Comparing the reactions of 47 wild emmer accessions tested in six nurseries, 11 markedly different patterns were discerned, indicating the probable presence of several different resistance genes.Genes for resistance to powdery mildew appear to be very common in wild emmer indigenous to Israel. Resistance was found in accessions from most collection sites, in all the geographic regions represented in the collection.The common occurrence of resistance and the apparent diversity of genotypes makes wild emmer a rich gene-pool for resistance to powdery mildew. Since genes for resistance to wheat pathogens can be quite readily transferred to cultivated wheat, wild emmer may be utilized as a valuable source of powdery mildew resistance in wheat breeding.     

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.     

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.