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.     

Variation for resistance to Fusarium head blight in spring barley

Fusarium head blight (FHB) is a fungal disease of barley and other cereals, causing substantial yield and quality losses, mainly due to the contamination of the harvest with mycotoxins. We aimed to evaluate genetic variation for resistance to FHB and its association with other plant characters in diverse barley germplasm in order to identify useful lines for resistance breeding. The 143 barley lines consisted of 88 current European spring barley lines and cultivars, 33 accessions from the genebank at IPK Gatersleben, and 22 lines obtained from North American institutions. We conducted artificially inoculated field experiments with Fusarium graminearum Schwabe during two seasons. FHB severity was evaluated by repeated assessment of visual symptoms. On a set of 49 lines several trichothecene mycotoxins were analyzed. Variation for FHB severity was quantitative. The lines with lowest FHB severity were 'CIho 4196' and 'PI 566203'. Also within the European spring barley collection variation for FHB severity was highly significant. There was a significant negative correlation between plant height and FHB severity (r=– 0.55). FHB severity assessed in the field and the amount of deoxynivalenol in the harvested grains were positively correlated (r= 0.87). Several lines with a useful level of FHB resistance were found or confirmed and are recommended as crossing partners.     

Genetic variation for head blight resistance in triticale caused by Fusarium graminearum isolates of different deoxynivalenol production

Summary Fusarium head blight infection causes severe yield losses and contamination of the grain with mycotoxins in triticale (× Triticosecale Wittmack) grown in temperate and semihumid areas. In a two-year experiment thirty-six genotypes were inoculated separately with two isolates of Fusarium graminearum differing fivefold in their in vitro deoxynivalenol (DON) production and the effect on various traits was studied. All traits were significantly affected by head blight. The two isolates differed considerably in their aggressiveness resulting in a mean reduction of grain weight per spike of almost 25% and 50%, respectively. Inter-annual correlation was high for average disease rating (r=0.63, P<-0.01) and low for the other traits. Therefore, disease rating, averaged from two to three records, was regarded a suitable criterion for screening purposes. The effect of isolates on genotypes was not stable over years. The mean DON content of five genotypes with diverse resistance levels was 68 mg kg^-1. In vitro DON production of the two isolates used for inoculation did not correspond to their aggressiveness and DON contamination of the grain.     

Diallel analysis of resistance to head blight caused by Fusarium culmorum in winter wheat

Summary Ten homozygous winter wheat genotypes representing different levels of resistance to Fusarium head blight were crossed in all possible combinations excluding reciprocals. Parents, F₁ and F₂ were inoculated with one pathogenic strain of Fusarium culmorum. Data for head blight, observed 21 days after first inoculation (OBS-2), and for the area under the disease progress curve, based on observations 14, 21 and 28 days after first inoculation (AUDPC), were analyzed. The contrast between parents and F₁ crosses indicated dommance effects of the resistance genes. Diallel analysis according to Griffing's Method 4, Model 1 showed significant general combining ability (GCA) effects for both F₁ and F₂; specific combining ability effects were not significant. With the exception of one genotype for which general performance for Fusarium resistance was not in agreement with its GCA, the resistance to F. culmorum was uniformly transmitted to all offspring, and the parents can be described in terms of GCA. It is suggested that in the progenies with one of the awned lines as parent, one resistance gene was linked with the gene coding for presence of awns, located on chromosome 4B. A single observation date, taken at the right time, was as effective in assessing resistance as the AUDPC.     

Expression of a Triticum turgidum var. dicoccoides source of Fusarium head blight resistance transferred to synthetic hexaploid wheat

Yield and quality reductions caused by Fusarium head blight (FHB) have spurred spring wheat (Triticum aestivum L.) breeders to identify and develop new sources of host plant resistance. Four wheat synthetic hexaploids (×Aegilotriticum sp.) were developed, each having a quantitative trait locus (QTL), Qfhs.ndsu‐3AS, providing FHB resistance from Triticum turgidum L. var. dicoccoides chromosome 3A. Synthetics were produced by hybridizing a ‘Langdon’‐T. dicoccoides‐ recombinant chromosome 3A substitution line (2n = 4x = 28, AABB with two accessions of T. tauschii (2n= 2x = 14, DD). Synthetics were inoculated and evaluated for FHB resistance in two separate greenhouse seasons. One synthetic, 01NDSWG‐5, exhibited FHB severity ratings of 36% and 32% in the separate seasons, compared with ratings of 9% and 30% for ‘Alsen’, a FHB‐resistant spring cultivar, and ratings of 70% and 96% for ‘McNeal’, a susceptible spring cultivar, respectively. Synthetic × Alsen backcross‐derived lines were produced to initiate combining different sources of FHB resistance.     

Genetic variation for resistance to Fusarium head blight in bread wheat

Summary During a four year period, a total of 258 winter and spring wheat genotypes were evaluated for resistance to head blight after inoculation with Fusarium culmorum strain IPO 39-01. It was concluded that genetic variation for resistance is very large. Spring wheat genotypes which had been reported to be resistant to head blight caused by Fusarium graminearum were also resistant to F. culmorum. The resistant germplasm was divided into three gene pools: winter wheats from Eastern Europe, spring wheats from China/Japan and spring wheats from Brazil. In 32 winter wheat genotypes in 1987, and 54 winter wheat genotypes in 1989, the percentage yield reduction depended on the square root of percentage head blight with an average regression coefficient of 6.6. Heritability estimates indicated that for selection for Fusarium head blight resistance, visually assessed head blight was a better selection criterion than yield reduction.     

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.     

Genetics of morphological traits in wild wheat,Triticum turgidum var.dicoccoides

Summary The mode of inheritance, linkage groups, and chromosomal location of 23 morphological and 4 biochemical traits were characterized in the wild tetraploid emmer wheat,Triticum turgidum var.dicoccoides. These traits were described and their mode of inheritance was determined by their segregation in four F₂ populations derived from crosses between four var.dicoccoides accessions and a tetraploiddurum cultivar. Linkage groups among the genes encoding for these traits were determined or postulated, and their chromosomal location was deduced by linkage to previously located genes. The genetic control of the following traits was characterized and is first reported here: black keel; hairy leaf sheath; hairy auricles; hairy rachilla; hairy kernel brush; obtuse flag leaf; and curved neck/peduncle. The linkage data indicated that developmentally-related genes tended to occur in clusters.     

The inheritance of resistance to head blight caused by Fusarium culmorum in winter wheat

Summary Crosses were made among ten winter wheat genotypes representing different levels of resistance to Fusarium head blight to obtain F₁ and F₂ generations. Parents, F₁ and F₂ were inoculated with one strain of Fusarium culmorum. Data on incidence of head blight 21 days after first inoculation were analyzed. Broad-sense heritabilities averaged 0.39 and ranged from 0.05 to 0.89 in the individual F₂ families. The joint-scaling test indicated that the inheritance of Fusarium head blight resistance was adequately described by the additive-dominance model, with additive gene action being the most important factor of resistance. With respect to the non-additive effects, dominance of resistance predominated over recessiveness. The number of segregating genes governing resistance in the studied populations was estimated to vary between one and six. It was demonstrated that resistance genes differed between parents and affected resistance differently.     

Temperature-sensitive genes for resistance to Puccinia striiformis in Triticum dicoccoides

Summary Twenty-six selections of wild emmer collected at different sites in Israel and three entries obtained from Turkey were studied for the possible presence of temperature-sensitive genes controlling reaction to Puccinia striiformis. In tests carried out in Israel with a local P. striiformis isolate at two temperature regimes, 16 selections showed a change in infection rating toward resistance at the higher temperature regime. In comparable tests performed in the United States with a Montana P. striiformis isolate of different virulence pattern-including 14 of the same selection—ten entries displayed a temperature-sensitive reaction. Although slightly different temperature regimes were used in Israel and the United States, the results obtained were in general agreement. The shift toward resistance observed in the seedling tests at the higher temperature-profile was also evident in the field in the mature plant stage with increasing spring temperatures.     

Response to selection in F2 generations of winter wheat for resistance to head blight caused by Fusarium culmorum

Summary In a field trial, F₃ winter wheat lines from plants selected for Fusarium head blight resistance in F₂ generations of a set of crosses, composing a 10×10 half diallel, were tested with their parental lines for resistance to Fusarium culmorum. Selection responses averaged 3.7% on the head blight percentage scale and ranged from –22.0% to 27.1%. Realized heritabilities averaged 0.23 and ranged from 0 to 0.96. Significant transgression for resistance was observed which was suggested to be genetically fixed. It was estimated that resistant parents differed in one or two resistance genes. The possibility of accumulation of resistance genes was shown. The level of head blight resistance of the parental line appeared to be a good indicator of the potential resistance level of its crosses.     

Location of genes for high grain protein percentage and other quantitative traits in wild wheat Triticum turgidum var. dicoccoides

Summary The genetic control of grain protein percentage (GPP) in the wild tetraploid wheat, Triticum turgidum var. dicoccoides, was determined by crossing four accessions of this taxonomic variety with durum cultivar Inbar, and analyzing the parents, F₁ and F₂ populations. Reciprocal crosses indicated no cytoplasmic effect on GPP. The F₂ variation was continuous in all crosses, showing no transgressive segregation. However, crosses between different accessions of var. dicoccoides showed transgressive segregation indicating the presence of different genes for high GPP in these accessions. Grain protein percentage was mostly codominant with high GPP, showing either no dominance, or a weak dominance. Heritability coefficients (broad sense) ranged from 0.30 to 0.53. Correlation coefficients between GPP and yield components were usually significantly negative, with the exception of the number of spikelets per spike, and in some crosses, grain weight.The number and chromosomal location of genes coding for high GPP were determined by the association between GPP and 27 markers (23 morphological and 4 biochemical markers). For this purpose, the genetic control of these markers, their linkage groups and chromosomal location were studied. At least four loci for high GPP that segregated in the F₂ populations are suggested: one on chromosome arm 1AS, marked by the black glume gene (Bg); one on 1BS, marked by the HMW gliadin locus Gli-B1; one on group 5, marked by the genes for beaked glume (Bkg) and toothed palea (Tp); and one on group 7, marked by the kinky neck gene (Kn). The relationship between GPP and several yield components was studied in a similar manner. In general, loci of markers that correlated positively with high GPP were not correlated with a decrease in yield components. Moreover, several loci of var. dicoccoides were associated with an increase in yield components.The utilization of markers for chromosomal location of genes coding for quantitative traits is compared to the technique of aneuploid analysis, commonly used in wheat. The significance of the above findings for breeding is discussed.     

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.     

Genetic analysis of resistance to Fusarium head blight caused by Fusarium graminearum in Chinese wheat cultivar Sumai 3 and the Japanese cultivar Saikai 165

The genetic constitution of resistance to Fusarium head blight (FHB, scab) caused by Fusarium graminearum in the Chinese wheat cultivar Sumai 3 and the Japanese cultivar Saikai 165 was investigated using doubled haploid lines (DHLs) and recombinant inbred lines (RILs). Frequency distributions of DHLs derived from two F₁ crosses, Sumai 3 (very resistant to resistant; VR-R) / Gamenya (very susceptible; VS) and Sumai 3 / Emblem (VS), fitted well to 1: 2: 1 (resistant: moderately resistant: susceptible) ratios for reaction to FHB in the field. It is suggested that the resistance of Sumai 3 is controlled by two major genes with additive effects. One of the resistance genes may be linked in repulsion to the dominant suppressor B1 for awnedness with recombination values 15.1 ± 3.3% in Sumai 3 /Gamenya and 21.4 ± 4.3% in Sumai 3 / Emblem. Saikai 165 is a Japanese resistant line derived from an F₁ Sumai 3 / Asakaze-komugi (moderately resistant; MR). The data for RILs derived from the cross Emblem / Saikai 165, indicates that three resistance genes control the resistance of Saikai 165. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic analysis of inbred lines and their crosses for resistance to head blight (Fusarium culmorum,F. graminearum) in winter rye

Summary For genetic analysis of head blight in winter rye (Secale cereale) caused by Fusarium culmorum, six homozygous inbred lines from the Petkus gene pool were crossed in all combinations to obtain 15 diallel F₁ crosses and the corresponding 15 F₂ crosses. These materials and 10 additional inbreds were artificially inoculated in a 2-year field experiment. The inbreds were also tested with F. graminearum in a separate sub-experiment.Single disease rating, average disease rating, and yield components (grain-weight per spike, 1000-grain weight, kernel number per spike) relative to the non-inoculated treatment were significantly affected by Fusarium head blight in all material groups. The relative grain weight per spike ranged from 26% to 88%. Significant genotypic and genotype x year interaction variances were found throughout. Heritabilities were highest for homogeneous inbreds (h²=0.6–0.8) and lowest for heterogeneous F₂ crosses (h²=0.4–0.6). Disease rating and relative grain-weight per spike were highly correlated for the inbreds and F₂ crosses (r0.7, P0.01), but lower for the F₁ crosses (r0.6, P0.05). Inter-annual correlation coefficients for disease ratings and relative grain-weight per spike ranged from r0.7 (inbreds) to r0.5 (F₂ crosses). The diallel analysis showed significant GCA effects only for relative 1000-grain weight in 1990, but significant SCA and SCAx year interaction variances for most traits. The resistances of 16 inbreds to F. culmorum and F. graminearum were tightly associated for all traits (r=0.96–0.97, P0.01).In conclusion, only slow progress can be expected from selecting for Fusarium head blight resistance in rye due to the limited amount of additive genetic variance and the great improtance of environmental factors.     

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.     

Plant nitrogen distribution in wild tetraploid (Triticum turgidum dicoccoides) and hexaploid wheat (Triticum aestivum)

Summary Differences were found in total nitrogen uptake and its pattern of distribution in the main tiller amongst five lines of wild tetraploid wheat (Triticum turgidum dicoccoides) and between it and two hexaploid wheats (Triticum aestivum) under low (48 ppm) and higher (240 ppm) levels of soil nitrogen.Under the low soil nitrogen level the hexaploids had higher amounts of total nitrogen in the main tiller than the dicoccoides lines, but under the higher soil nitrogen level, three of the dicoccoides lines had significantly (P<0.01) higher, and the other two lines, similar amounts as the hexaploids. The total amount of grain nitrogen in the hexaploids was significantly (P<0.01) higher than the five dicoccoides under the low nitrogen soil level but under the higher level, two of the dicoccoides lines had similar amounts as one of the hexaploids (cv. Bencubbin) but significantly (P<0.01) lower than the other (cv. Argentine IX).The efficiency of nitrogen translocation to the grain was significantly (P<0.01) lower in a primitive, compared with four cereal forms of dicoccoides under both low and high levels of soil nitrogen. The cereal forms of dicoccoides, while similar in nitrogen translocation efficiency under low soil nitrogen as the lower translocation efficiency hexaploid (cv. Bencubbin), were significantly (P<0.01) and substantially lower than it under the higher soil nitrogen level.     

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.     

Race-specificity of temperature-sensitive genes for resistance to Puccinia striiformis in Triticum dicoccoides

Summary Twenty-four entries of wild emmer possessing temperature-sensitive genes for resistance to yellow rust were studied in the seedling stage, at two temperature-profiles, with 15 pathogenic races from 11 countries in South America, Africa, Asia and Europe. It was shown that the majority of the resistance genes in these wild emmer entries were race-specific. In most of these entries a more resistant reaction was displayed at the higher temperature-profile; however in three entries a shift in reaction towards resistance was observed with certain races but towards susceptibility with some of the other races, suggesting that two different kinds of temperature-sensitive genes were involved in each of these entries. The similarity of temperature-sensitive genes occurring in wild emmer and in cultivated wheat is discussed.     

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.