Variation for resistance to head blight caused by Fusarium graminearum in wild emmer (Triticum dicoccoides) originating from Israel

Head blight of wheat (FHB, scab) caused by Fusarium spp. has been associated with yield and quality losses in many wheat-growing regions. In tetraploid wheat sources of resistance are scarce. In the search for novel sources of resistance, 151 Triticum dicoccoides genotypes, originating from 16 habitats in Israel and one habitat in Turkey together with several control genotypes, were evaluated for reaction to fungal spread (Type II resistance) in replicated greenhouse experiments. Significant genetic diversity was found among the tested genotypes, the broad sense heritability for Type II FHB resistance was 0.71. Most of the tetraploid accessions were highly susceptible, only a few showed moderate resistance. Among the eight T. dicoccoides lines with the lowest relative infection rates, five originated from the Mt. Gerizim population, and three from the Mt. Hermon population. None of the T. dicoccoides lines reached the level of resistance present in the common wheat cultivar Sumai3. This revised version was published online in July 2006 with corrections to the Cover Date.     

A valuable source of yellow rust resistance in Israeli populations of wild emmer,Triticum dicoccoides Koern.

Summary An extensive collection of wild emmer was screened for yellow rust resistance in a joint project by research institutes in Israel and The Netherlands. In inoculation experiments performed both in seedling stage and at maturity, the wild emmer accessions displayed a diversity of responses to yellow rust infection, ranging from immunity to complete susceptibility.SelectionTriticum dicoccoides var.aaronsohni G-25 was most promising proving highly resistant to all 21 races and field races used. Selection G-7, which was also very resistant in the trials, differed slightly from G-25 in infection types produced on inoculation with one of the isolates; this may be an indication that the genetic factors controlling resistance in these two selections are not identical.It is concluded that the diversified populations of wild emmer indigenous to Israel possess genes which confer high resistance to a very wide spectrum of yellow rust races, including all common races in Western Europe and the Middle East. As resistance factors can be transferred easily fromT. dicoccoides to durum and common wheat, this source may be utilized in producing varieties of cultivated wheat with improved yellow rust resistance.Joint contribution from the Volcani Institute of Agricultural Research, Bet Dagan, Israel, 1969 Series, No. 1594-E, and the Institute of Phytopathological Research, Wageningen, The Netherlands.     

Genetic resources of wild cereals in Israel and vicinity. II. Phenotypic variation within and between populations of wild barley,Hordeum spontaneum

Summary Populations of wild barley, Hordeum spontaneum, in Israel, originating from diverse habitats, and tested earlier for allozyme (Nevo et al., 1979a) and disease resistance polymorphisms (Moseman et al., 1984; Nevo et al., 1984a), were compared and contrasted for performance in agronomically important phenotypic traits. The traits compared involved 10 variables comparing germination, earliness, numerical and weight variables of biomass and yield. The field experiments were conducted in 1980, 1981 and 1982 in two relatively standardized and contrasting environments: mesic (Mount Carmel, Haifa) and xeric Avedat farm, and Sede Boqer (both in the northen Negev desert). The experimental design involved 26 population quadrangles at Avedat farm (1980), and rows of randomized plants of 11 populations in both Haifa and Sede Boqer (1981 and 1982).The results indicate that the characters studied are partly genetically determined. Striking genetic variation was found between and within populations in each site, whereas remarkable environmental variation including genetic environmental interaction was found between the mesic and the xeric sites, as well as between populations and years. We conclude that natural populations of wild barley in Israel vary not only in genetic polymorphisms of allozymes and disease resistance, but also in quantitative traits of agronomie importance. These traits are economically significant and should be conserved and utilized in barley crop improvement.     

Genetic resources of wild cereals in Israel and vicinity. I. Phenotypic variation within and between populations of wild wheat,Triticum dicoccoides

Summary Populations of wild emmer wheat, Triticum dicoccoides, in Israel, originating from diverse habitats, and tested earlier for allozyme (Nevo et al., 1982) and disease resistance polymorphisms (Moseman et al., 1983a, 1983b; Nevo et al., 1984a, 1984b), were compared and contrasted for performance in agronomically important phenotypic traits. The traits compared involved 10 variables comparing germination, earliness, biomass and yield variables. The field experiments were conducted in 1980, 1981 and 1982 in two relatively standardized and contrasting environments: mesic (Mount Carmel, Haifa) and xeric (Acedat Farm, and Sede Boqer, in the northern Negev desert). The experimental design involved 12 population quadrangles at Avedat Farm in 1980, and rows of randomized genotypes of five populations in both Haifa and Sede Boqer in 1981 and 1982.The results indicate that the characters studied are partly genetically determined. Striking genetic variation was found between and, at least in some characters, also within populations in each site, whereas remarkable environmental variation including genetic-environmental interaction was found between the mesic and the xeric sites. We conclude that natural populations of wild emmer wheat in Israel vary not only in genetic polymorphisms of allozymes and disease resistance, but also in quantitative traits of agronomic importance. These traits are economically significant and should be conserved and utilized in wheat crop improvement.     

A new powdery mildew resistance gene: Introgression from wild emmer into common wheat and RFLP-based mapping

An Israeli accession (TTD140) of wild emmer, Triticum turgidum var. dicoccoides, was found resistant to several races of powdery mildew. Inoculation of the chromosome-arm substitution lines (CASLs) of TTD140, in the background of the Israeli common wheat cultivar ‘Bethlehem’ (BL), with five isolates of powdery mildew revealed that only the line carrying the short arm of chromosome 2B of wild emmer (CASL 2BS) exhibited complete resistance to four of the five isolates. To map and tag the powdery mildew resistance gene, 41 recombinant substitution lines, derived from a cross between BL and CASL 2BS, were used to construct a linkage map at the gene region. The map, which encompasses 69.5 cM of the distal region of chromosome arm 2BS, contains six RFLP markers, a morphological marker (glaucousness inhibitor, W1 ^I), and the powdery mildew resistance gene. Segregation ratios for resistance in F₂ of BL × CASL 2BS and in the recombinant lines, combined with the susceptability of F₁ progeny to all tested isolates, indicate that resistance is controlled by a single recessive allele. This alleleco-segregated with a polymorphic locus detected by the DNA marker Xwg516, 49.4 cM from the terminal marker Xcdo456. The new powdery mildew resistance gene was designated Pm26. This revised version was published online in July 2006 with corrections to the Cover Date.     

Postulation of resistance genes to yellow rust in wild emmer wheat derivatives and advanced wheat lines from Nepal

Summary Seedlings of 38 wild emmer derivatives, and a total of 53 advanced wheat varieties/lines introduced from the International Maize and Wheat Improvement Centre (CIMMYT) or other sources, Nepalese breeding lines and local cultivars were inoculated with 18 different yellow rust isolates to postulate yellow rust resistance genes (Yr). Many wild emmer wheat derivatives used were resistant to all isolates indicating the presence of undescribed genes. Some derivatives carried Yr9, Yr6 and/or YrSU. Genes Yr1, Yr2, Yr6, Yr7, Yr8, Yr15, YrSU and YrA+ are no longer effective in Nepal; Yr4, Yr5, Yr9, Yr10, YrSP and YrSD are still effective; the effectiveness of Yr3 remains unclear. This study shows that stripe rust resistance in seedling stage of most Nepalese cultivars and advanced materials is based on Yr9 with combinations of Yr2, Yr6, Yr7, and YrA+, of which only Yr9 is still effective in Nepal. In many countries Yr9 has lost its effectiveness. Therefore the introduction of new Yr-genes from wild emmer wheat in Nepalese cultivars is highly important.     

Molecular characterization of a novel powdery mildew resistance gene Pm30 in wheat originating from wild emmer

Powdery mildew caused by Erysiphe graminis f. sp. tritici is one of the most important wheat diseases in many regions of theworld. A powdery mildew resistance gene, originating from wild emmerwheat (Triticum dicoccoides) accession `C20', from Rosh Pinna, Israel,was successfully transferred to hexaploid wheat through crossing andbackcrossing. Genetic analysis indicated that a single dominant genecontrols the powdery mildew resistance at the seedling stage. SegregatingBC₁F₂ progenies of the cross 87-1/C20//2*8866 wereused for bulked segregant analysis (BSA). The PCR approach was used togenerate polymorphic DNA fragments between the resistant and susceptibleDNA pools by use of 10-mer random primers, STS primers, and wheatmicrosatellite primers. Three markers, Xgwm159/430,Xgwm159/460, and Xgwm159/500, were found to be linked tothe resistance gene. After evaluating the polymorphic markers in twosegregating populations, the distance between the markers and the mildewresistance gene was estimated to be 5–6 cM. By means of ChineseSpring nullisomic-tetrasomics and ditelosomics, the polymorphic markersand the resistance gene were assigned to chromosome arm 5BS and werephysically mapped on the gene rich regions of fragment length (FL) 0.41–0.43 by Chinese Spring deletion lines. As no powdery mildew resistancegene has been reported on chromosome arm 5BS, the mildew resistancegene originating from C20 should be a new gene and is designated Pm30.     

The introduction into bread wheat of a major gene for resistance to powdery mildew from wild emmer wheat

Summary A new source of resistance to wheat powdery mildew caused by Erysiphe graminis has been transferred to hexaploid bread wheat, Triticum aestivum, from the wild tetraploid wheat, Triticum dicoccoides. The donor was crossed to bread wheat and the pentaploid progeny was then self-pollinated. Plants having a near stable hexaploid chromosome complement were selected in the F₃ progeny and topcrossing and backcrossing of these to a second wheat cultivar to improve the phenotype was undertaken. Monosomic analysis of early backcross lines showed the transferred gene to be located on chromosome 4A. The gene has been designated Pm16.     

Resistance of wild emmer wheat to stem rust: Ecological,pathological and allozyme associations

Summary Seedling resistance to wheat stem rust was determined in populations of wild emmer wheat, Triticum dicoccoides, and characterized by means of ecological factors and allozyme genotypes. Reactions to wheat stem rust were studied in 102 single plant accessions of T. dicoccoides from ten populations by inoculation with Puccinia graminis tritici race 14, isolate GSR-739. Six populations displayed different degrees of response polymorphism with reactions ranging from high resistance to complete susceptibility, whereas four populations contained only susceptible plants. In some of the accession, unexplained intrasib variation in resistance and intraplant variation of infection-types were found. Resistance to stem rust was negatively correlated with two ecological factors, altitude and number of Sharav (hot-dry) days which are unfavorable to disease development. Variation in stem rust response was shown to exist in ecogeographic regions where climatic variables enhanced the development of the fungus, conceivably maintained by natural selection. Likewise, allozyme genotypes, single or in multiple loci combinations, appeared to be associated with resistance or susceptibility to rust. Such association need to be verified by genetic studies in order to become established as useful markers.     

Grain yield and grain protein percentage of common wheat lines with wild emmer chromosome-arm substitutions

Wild emmer wheat, Triticum turgidum subsp. dicoccoides, (2n = 4× = 28; genome BBAA), the progenitor of domesticated wheat, is genetically closely related to durum and common wheat. This wild taxon has characteristics that would be valuable if transferred to domesticated wheat. A series of chromosome-arm substitution lines (CASLs) of wild emmer wheat were produced in the background of an Israeli common wheat cultivar. These CASLs were evaluated in a pot experiment and in field trials in Israel and California for their grain yield (GY) and its components and for grain protein percentage. In addition, the extent of genetic interactions (epistatic effects) between “wild” and “domesticated” alleles, within and between homoeologous groups 1 and 7 as expressed in grain and protein yields and other quantitative traits, were determined. The research has shown that wild emmer wheat harbors genetic variability for quantitative traits and that the “wild” genes interact among themselves in a non-additive way in the common genetic background. Several chromosome arms improve GY and protein percentage in common wheat, but their effects will be presumably enhanced when combination of genes from several “wild” arms are integrated into a single “domesticated” genotype. Hence, the interaction between these genes and those in the recipient common wheat must be accounted for when higher yield or protein content is desired. The results of this study indicate on the potential of this material for breeding and genetic analysis, and support the idea of pyramiding genes from a wild species.     

Identification and genetic mapping of a powdery mildew resistance gene in wild emmer (<Emphasis Type="Italic">Triticum dicoccoides</Emphasis>) accession IW72 from Israel

Powdery mildew, caused by Blumeria graminis f. sp. tritici (Bgt), is a devastating disease of wheat (Triticum aestivum) in China and worldwide, causing severe yield losses annually. Wild emmer (T. dicoccoides) accession IW72 collected from Israel is resistant to powdery mildew at the seedling and adult stages. Genetic analysis indicated that the resistance was controlled by a single dominant gene, temporarily designated MlIW72. The F₂ population and F₃ families derived from a hybrid between IW72 and susceptible durum wheat line Mo75 were used for molecular mapping of the resistance gene. MlIW72 was linked with SSR loci Xgwm344, Xcfa2040, Xcfa2240, Xcfa2257 and Xwmc525 on the long arm of chromosome 7A. In addition, two STS markers, MAG2185 (derived from RFLP marker PSR680) and MAG1759 (developed from EST CD452874), were mapped close to MlIW72. All these markers were physically located in the terminal bin 0.86–1.00 of 7AL. The chromosome location and genetic mapping results suggested that the powdery mildew resistance gene identified in wild emmer accession IW72 might be a new allele at the Pm1 locus or a new locus closely linked to Pm1.     

Phenotypic and genotypic analysis of a commercial cultivar and wild populations of Anemone coronaria

Seven wild populations of Anemonecoronaria were assessed for 11 phenotypic traits, most of them having economic value for the flower industry. The wild populations were sampled to represent the diversity in habitats, climates, rock and soil types, terrains, and elevations in Israel. AFLP analysis was carried out on 12 individuals from each of six out of the seven wild populations and for six individuals from the commercial cultivar ‘Mona-Lisa’. It was found that the Dorot population, which is located in the area bordering the semi-arid zone at the very end of the species distribution, exhibits extreme and different phenotypes with relatively low variability compared withthe other wild populations. The other six wild populations, that grow in more favorable geographic and climatic conditions exhibit phenotypes with larger plants, larger numbers of flowers and less dissected leaves. These populations were less uniform than that of Dorot. Genetic characterization by AFLP markers revealed a total of 165 bands. The wild populations exhibit wide variation within-population, with about 80% polymorphic bands and average gene diversity between pairs of about 30%. The Dorot population has the lowest genetic variation and the Megido population the highest. Thus, the phenotypic variation reflects the genetic variation. The cultivar ‘Mona-Lisa’, as expected, has much lower genetic variation. The Dorot population and the ‘Mona-Lisa’ cultivar were found to have the largest genetic distances from the other wild populations, and the highest genetic variation between themselves. Phenetic analysis yielded a dendrogram describing the genetic relatedness of these populations. This revised version was published online in August 2006 with corrections to the Cover Date.     

Environmental, phenotypic and genetic variation of wild barley (Hordeum spontaneum) from Israel

Wild relatives of crop plants offer an attractive gene pool for cultivar improvement. We evaluated genetic and phenotypic variation for a set of 72 Israeli accessions of wild barley from 21 populations. These populations were grouped further into four ecotypes. In addition, environmental variables describing the local conditions for the populations were used to infer the environmental divergence. Genetic, phenotypic and environmental distances were estimated from the data and UPGMA dendrograms constructed. The results showed that genetic variation was larger between populations than within them, whereas for phenotypic measurements variation was larger within populations than between them. No significant correlation was found between genetic and phenotypic similarities, or between phenotypic and environmental similarities, whereas a weak correlation between genetic and environmental similarities was detected. Twenty-three AFLP-markers were identified to be ecotype specific. Chromosomal location was known for five of these markers. Four of the five ecotype specific markers were correlated with both phenotypic traits and environmental variables.     

Cytogenetic and genetic relationships between populations of Aegilops ventricosa Tausch

Summary Five samples of Aegilops ventricosa (2n=4x=28, genome formula DDNN) from different geographical origins, were crossed in a diallelic scheme. Metaphase I chromosome pairing of the hybrids, accounting for all the possible genetic combinations, was analysed. Only bivalents were formed in some hybrids, while multivalents were scored in other ones. Seed storage proteins, gliadins and albumins, were also analysed by means of polyacrylamide gel electrophoresis. Based on the presence of multivalents in hybrids, and on the differences in seed protein profiles, the samples could be grouped into two clusters. Meiosis was regular in hybrids obtained within samples of the same group, while multivalents were present in hybrids involving a sample of one group and one of the other. The evolutionary trends in Ae. ventricosa are discussed.     

Crop yield and grain quality of emmer populations grown in central Italy, as affected by nitrogen fertilization

The increasing demand for traditional and natural foods has renewed the interest in hulled wheat species. Among these, Triticum dicoccum Schübler has survived in Italy only in a few hilly and mountainous areas of central and southern Italy. As a rule, emmer is cultivated in marginal areas with organic farming procedures that use very low N inputs, since this wild species is characterised by low yield, long and weak culms that easily lodge under windy conditions. Unfortunately, there is a lack of information on crop productivity and N effect for this crop. Research was carried out to study the effect of cultivation year (2004 and 2005), plant origin (Garfagnana, Leonessa and Molise) and N dressing (N₀, N₃₀, N₆₀ and N₉₀ kg ha⁻¹) on crop yield and grain quality of emmer grown in south-central Italy. Fertilizer was split at seeding, tillering and stem elongation. Tested parameters were highly influenced either by crop origin or N application (biomass, hulled and threshed grain yield, spikes m⁻², spikelets per spike, kernel weight, plant height, lodging, kernel ash and proteins). Several parameters augmented as fertilization rate increased (hulled and unhulled grain yield, biomass accumulation, spikes m⁻², kernels m⁻², protein content); 1000-kernel weight showed an opposite trend and in some cases no differences were noticed among fertilized treatments (plant height and spikelets per spike). Molise was the most productive population, closely followed by Garfagnana. The present research rejected some common belief that emmer has to be grown without N dressing, and crop undergoes lodging in marginal mountainous areas. Besides, grain yield of N₉₀ and N₆₀ treated emmer was only 6 and 18% lower compared to the five most important durum wheat varieties cultivated in the same area, suggesting emmer as a possible alternative crop to durum wheat in marginal areas of Mediterranean-type agro-ecosystems.     

A genetic diversity analysis of wild Lespedeza populations based on morphological characters, allozyme and RAPD methods

Genetic variability within and among 14 Chinese natural populations of Lespedeza Michx. Used mainly for wildlife food and cover and for erosion control was evaluated on the basis of morphological characteristics, allozyme markers and random amplified polymorphic DNAs (RAPDs). All methods disclosed a high degree of allelic heterogeneity among samples, also revealing a high degree of gene diversity. Genetic variation among populations was higher for RAPD than for enzyme loci. Correlation between morphological, allozyme and RAPD distance matrices were calculated by Mantel’ tests. There was a little disparity between the three level's distances on population relationships. However, there is a weak significant correlation (P < 0.001) between morphology and allozyme distances as well as morphology and RAPD distances, and the test of this correlation is very high.     

Genetic characterization of wild barley populations (Hordeum vulgare ssp. spontaneum) from Kazakhstan based on genome wide SNP analysis

The wild ancestral form of barley, Hordeum vulgare ssp. spontaneum, is a valuable source for gene enrichment of cultivated barley. The purpose of this work was to study the area of distribution as well as the extent and structure of genetic variation of wild barley populations grown in Kazakhstan. It was found that distribution of wild barley populations in Kazakhstan is restricted to the most southern province. A genome wide single nucleotide polymorphism (SNP) analysis was performed in order to study the level of the genetic diversity in 96 accessions representing 14 wild barley populations from Kazakhstan and 25 accessions from the Middle East which is the center of diversity of this subspecies. The oligonucleotide pooled assay was used to genotype 384 SNPs distributed throughout the genome. In total 233 polymorphic SNPs were selected for further statistical analysis. The level of genetic diversity of wild barley populations from Kazakhstan was predictably narrower (He = 0.19 ± 0.01) in comparison with wild barley samples from the Middle East (He = 0.29 ± 0.01). The results suggested that H. vulgare ssp. spontaneum populations in Kazakhstan probably represent a recent spread of a limited number of plants from the primary distribution area and might be well adapted to winter low temperature.     

Cytogenetic analysis of translocations in cotton

In order to create cotton translocation lines, 191 translocations were recovered as heterozygotes following combined treatment of seeds by colchicine and γ‐rays, irradiation of seeds by thermal neutrons and c‐irradiation of pollen. Cytogenetic analysis showed that chromosome translocations involving two chromosomes arose more often (180) than those involving three chromosomes (11). The heterozygous translocations were characterized by different formation and frequencies of the multivalents at metaphase I of meiosis. Only 104 translocations were characterized by a multivalent frequency of more than 0.25 per cell. Most of the translocations (156) exhibited a high meiotic index, but 11 were characterized by a decreasing meiotic index and by an increase in the percentage of tetrads with micronuclei. The pollen fertility of translocations differed significantly and varied from high fertility to pollen sterility. The high frequency of abortive pollen grains was typical for about one half of translocations studied. The translocations were made homozygous and as a result, 12 new reciprocal homozygous translocation lines were obtained in cotton.     

Cytogenetic studies and the occurrence of triploidy in the wild potato species Solanum commersonii Dun.

Summary The South American wild potato species Solanum commersonii Dun. exists as two subspecies, subsp. commersonii and subsp. malmeanum (Bitter) Hawkes and Hjerting, and as both diploid and triploid cytotypes. The two subspecies were completely cross fertile but there were reciprocal differences in the number of seeds set per berry and the mean seed weight. Crosses between diploids and colchicine-induced autotetraploids yielded only a few seeds per berry, but 16 of the 18 germinated seeds were triploid or nearly so. Such a high proportion of triploids is unusual for such crosses in potatoes. Some triploids has a high percentage of stainable pollen. The origin of naturally occurring triploids is discussed in the context of 2n gametes, a triploid block, and endosperm balance numbers.