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.     

Increase in grain protein percentage in high-yielding common wheat breeding lines by genes from wild tetraploid wheat

Summary Forty-one breeding lines of common wheat, derived from crosses between the Israeli cultivars Miriam and Lakhish and high-protein lines of wild tetraploid wheat, Triticum turgidum var. dicoccoides, were tested for various protein and yield parameters in field trials, under typical agronomic conditions. All lines had a higher grain protein percentage (GPP) than the leading Israeli cultivar Deganit, which was grown as a control. Grain yield (GY) ranged in the breeding lines from a low of 2.44 t/ha to as high as that of Deganit (6.95 t/ha). Despite the weak negative correlation between GPP and GY, several lines excelled both in GPP and in GY. The grain protein yield (GPY) of some of these selected breeding lines was higher than that of Deganit; e.g., 1.19 t/ha in the best line vs. 1.02 t/ha in Deganit. The 16.7% increase in GPY in this line reflected a more efficient utilization of nitrogen.     

Segregation of genetic markers among wheat doubled haploid lines derived from wheat x maize crosses

Summary Wheat doubled haploid (DH) lines were produced from the F₁ hybrid, Fukudo-komugi x Oligo Culm, through intergeneric crosses between wheat and maize. F₂ plants and 203 DH lines were analyzed for the segregation of the eight genetic markers, namely, grain proteins, grain esterases, GA-insensitivity and glume traits. The segregation in the F₂ plants fitted to the expected ratios. No deviation was observed among the DH lines, either, except for the glume pubescence. The result indicates the absence of correlation between the markers investigated and the efficiency of embryo formation in the DH lines.     

Inheritance of glume color and gluten strength in durum wheat

Summary The correlation between glume color and gluten strength and the heritability of each trait was estimated in two durum wheat crosses. Brown glume color appeared to be dominant to white in both crosses. In one cross, glume color was clearly controlled by one gene while in the other cross it appeared to be controlled by one or two genes with modifiers. The heritability of gluten strength was moderately high. The correlation of F₂ glume color and F₃ gluten strength was high (r=0.66 and 0.78) indicating that F₂ glume color was a good predictor of gluten strength in the F₃ generation. Selection for glume color appears to be an effective breeding strategy for improving gluten strength in those environments where glume color differences are easily detected.     

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.     

A comparison of marker-assisted and phenotypic selection for high grain protein content in spring wheat

Wheat grain protein content (GPC) is a primary end-use quality determinant for hard spring wheat (Triticum aestivum L.), and marker-assisted selection (MAS) could help plant breeders to develop high GPC cultivars. Two experiments were conducted using two populations developed by crossing low GPC cultivars (Ember) and (McVey) with (Glupro), which contains a high GPC QTL from Triticum dicoccoides (DIC). In one experiment, MAS and phenotypic selection (PS) were employed to select high GPC genotypes, and the selected genotypes were grown in six North Dakota (ND), USA environments. In a second experiment, molecular markers were used to select BC₂F₂ plants from each marker class for the DIC allele from each population. These plants were twice self-pollinated to produce BC₂F₄ plants, which were grown in single ND and Minnesota (MN) environments. Mean GPC was highest among lines using PS at two environments and not significantly different between MAS and PS in the other four environments. Lines presumably homozygous for DIC alleles had significantly higher GPC than their respective low GPC parents. The phenotypic GPC variation explained by the markers (r ²) was 30% at the ND and 15% at the MN environment. The use of PS was as effective as MAS in selecting for high GPC genotypes and more effective in some environments. This likely can be attributed to PS enabling selection for both the major QTL and other genes contributing to GPC. The use of molecular markers might be more advantageous for transferring the high GPC DIC QTL in a backcrossing program during parent development.     

Transfer of high kernel weight and high protein from wild tetraploid wheat (Triticum turgidum dicoccoides) to bread wheat (T. aestivum) using homologous and homoeologous recombination

Summary Wheat pentaploids were produced by hybridizing a high kernel weight (1000 grain wt=56 g), high protein (25.4%) line of wild tetraploid wheat (Triticum turgidum dicoccoides) as male parent, with the three hexaploids (T. aestivum) — normal Chinese Spring and its two homoeologous pairing mutants, ph _1b and ph ₂. The pentaploids were crossed as female parents to the two commercial hexaploid cultivars Warigal and Barkaee and 42-chromosome stable plants selected from the F₁ of the pentaploid x hexaploid crosses.Mean protein content of certain F₃ lines from all six pentaploid x hexaploid crosses was significantly higher than Chinese Spring and the respective commercial hexaploid parent (p<0.005) indicating high protein had been transferred from the tetraploid to the hexaploid level.Kernel weight amongst certain F₃ lines of the three pentaploids x Barkaee was significantly (p<0.0005) higher than either Chinese Spring or Barkaee, indicating the transfer also of high kernel weight from the tetraploid to the hexaploid level. However kernel weight was not significantly increased over Warigal in any F₃ lines of its crosses with the three pentaploids.High levels of homoeologous chromosome pairing in the ph-mutant pentaploids, plus evidence for significant modification of the composition of high-molecular weight (HMW) glutenin subunits of grain protein in certain F₃ derivatives of the ph-mutant pentaploid x hexaploid, crosses indicates that the ph-mutant-derived lines may possess novel (intergenome) genetic recombination, at least for high protein, and possibly kernel weight.     

Intra- and inter-population variations in grain protein percentage in wild tetraploid wheat,Triticum turgidum var. dicoccoides

Summary Grain protein percentage (GPP) was studied in 910 accessions of the wild tetraploid wheat, Triticum turgidum var. dicoccoides, collected from 22 populations representing different ecogeographical conditions in Israel. High values of GPP were found, ranging from 19.7% to 28.0% for population means, and from 14.1% to 35.1% for single accessions. Marginal populations had usually lower GPP and smaller intra-population variation than central ones. Repeated sampling of some central populations for four consecutive years revealed relatively large intra-population fluctuations. A high and significant genetic component of variation was found within and between populations by a nested analysis of variance in two nurseries. However, the regression coefficients of parents vs. offsprings were relatively low, indicating a smaller genetic component of variation which may be accounted for by a significant genotype × environment interaction. No correlation was found between GPP and ecological factors, except for soil type: accessions growing on terra-rossa had higher GPP than those growing on basaltic soil. Accessions with black glumes, or with glabrous auricles, or with large grains exhibited high GPP values. A strategy for collecting accessions with high GPP is presented, and the potential use of high GPP genotypes in breeding programs is discussed.     

Association of RFLP markers with trait loci affecting clubroot resistance and morphological characters in Brassica oleracea L.

Summary Resistance to Plasmodiophora brassicae Wor. race 7, the causal agent of the disease clubroot, was examined in an F₂ population of a cross between a clubroot resistant broccoli (Brassica oleracea var. italica) and a susceptible cauliflower (B. oleracea var. botrytis). A genetic linkage map was constructed in the same population based on the segregation of 58 dispersed restriction fragment length polymorphism (RFLP) markers. Associations between the inheritance of RFLP marker genotypes and segregation for disease resistance, morphological and maturity characteristics were examined. For each triat examined, several chromosomal regions marked by RFLP probes appeared to contain trait loci, suggesting that each trait was under polygenic control. RFLP marker linkage to a major factor imparting dominance for clubroot resistance from the broccoli parent was observed in this population. Additionally, RFLP marker linkage to an independently segregating factor contributing clubroot resistance from the cauliflower parent was observed, indicating that it should be possible to use RFLP markers to facilitate selection of transgressive segregants having the combined resistance from both parental sources. In some instances, RFLP markers from the same or closely linked chromosomal regions were associated with both clubroot resistance and morphological traits. Analysis of RFLP marker genotypes at linked loci should facilitate the selection of desired disease resistant morphotypes.     

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.     

Additive gene action for resistance to Puccinia striiformis F.SP. Tritici in Triticum dicoccoides

Summary Seven single-plant selections of wild emmer, with temperature-sensitive minor-effect genes for stripe rust resistance, were intercrossed in eight combinations. The resulting progenies were studied for a possible additive gene action.The transgressive segregation towards resistance in F₂ observed in all the combinations indicates that additive gene action for resistance indeed occurs in wild emmer. The common occurrence of this phenomenon in random combinations suggests further that several minor-effect genes are involved.Following selection of the most resistant plants in F₂, a marked shift towards resistance was noted in F₃, which demonstrates a positive response to selection. In some instances, additive resistance selected for (in F₂) at the high temperature-profile was expressed (in F₃) also at the low temperature-profile. This kind of resistance, when utilized in breeding programmes, promises therefore to be effective over a range of temperatures.     

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 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.     

Predicting transgressive segregants in early generation using single seed descent method-derived <Emphasis Type="Italic">micro-macrosperma</Emphasis> genepool of lentil (<Emphasis Type="Italic">Lens culinaris</Emphasis> Medikus)

In a self-fertilised crop like lentil, the identification of transgressive segregants for economically important trait such as seed yield is an important aspect of any practical breeding programme. The prediction of expected transgressive segregants in F₁ generation obtained as a ratio of additive genic effect [d] and additive variance (D) i.e. [d]/√D was studied in 28 crosses of lentil generated in a diallel fashion involving four parents each of macrosperma (exotic) and microsperma (Indian) types, respectively, resulting in three hybridization groups. The seed material advanced to F₂, F₃ and F₄ generations through single seed descent method was evaluated to determine the observed transgressive segregants for seed yield/plant. The observed frequency of crosses showing more than 20% transgressive segregants in F₂ to F₄ generations were exhibited in 9(32%) crosses, of which 7(77%) crosses were of macrosperma × microsperma type. Genotypes Precoz and HPL-5 of the exotic group (macrosperma) produced maximum number of transgressive segregants with the genotypes L-259, L-4145 and PL-406 of the Indian origin (microsperma). Goodness of fit (non-significant χ² value) in F₂ generation was observed for 19(68%) crosses of the total genepool, out of which 9(56%) crosses each in F₃ and F₄ generation belonged to the macrosperma × microsperma group, depicting it as the gene pool of paramount importance to obtain maximum transgressive segregants, therefore establishing the efficacy of the method used.     

Inheritance of Cold Hardiness in Triticum aestivum × Synthetic Hexaploid Wheat Crosses

Synthetic hexaploid wheat, produced by combining tetraploid wheat (AB genome) with Triticum tauschii (D genome), was crossed to modern hexaploid wheat (Triticum aestivum ABD genome) in an attempt to introduce new cold hardiness genes into the common hexaploid wheat gene pool. The cold hardiness levels of F) hybrids ranged from similar to parental means to equal to the hardy parent, indicating that cold hardiness was controlled by both additive and dominant genes. As expected when dominant gene action is involved, differences between F₂ and parental means were smaller than comparable differences in the F., Frequency distributions of F₂—derived F₃ lines also suggested that dominant genes were involved in the control of cold hardiness in some crosses. Heritability estimates for cold hardiness ranged from 63 to 70 % indicating that selection for cold hardiness should be effective in populations arising from crosses between common and synthetic hexaploid wheat. However, high selection pressure on the progeny of crosses that included the most hardy T. aestivum, T. durum, and T. tauschii accessions as parents did not identify transgressive segregates for improved cold hardiness. These observations indicate that the close wheat relatives, sharing common genomes with T. aestivum, are not promising sources of new genes to increase the maximum cold hardiness potential of common hexaploid wheat.     

Increased grain protein content and its association with agronomic and end-use quality in two hard red spring wheat populations derived from Triticum turgidum L. var. dicoccoides

Suitability of wheat (Triticum aestivum L.) for many food products depends on its unique protein. Elevated grain protein content (GPC) and its quality influences the bread making properties of wheat. The objective of this study was to examine the association of elevated GPC with agronomic and end-use quality in two hard red spring wheat recombinant inbred (RI) populations derived from wild emmer (Triticum turgidum L. var. dicoccoides). The two hard red spring populations (ND683/Bergen and Glupro/Bergen) were developed using a single-seed-descent method. ND683 and ‘Glupro’ are high in GPC (180 g kg^-1), presumably due to the introgression of gene(s) from Triticum turgidum L. var. dicoccoides and ‘Bergen’ is low in GPC (145 g kg^-1). From each of the two populations 12 high- and 12 low-GPC RI lines (F_5:7) were selected for replicated testing at two North Dakota (ND) locations in 1995. In both populations, the high-GPC lines had significantly (p < 0.05) higher values compared to the low-GPC lines for mean GPC and water absorption. Mean grain yield of the high-GPC lines was not significantly different from the low-GPC lines in both populations. In the ND683/Bergen population, the high-GPC lines had significantly (p < 0.05) higher values than the low-GPC lines for mean plant height, days to heading, and flour extraction. GPC was significantly (p < 0.05)and negatively associated with test weight and also significantly (p < 0.01) and positively associated with water absorption in the Glupro/Bergen population. In these populations, results suggested that it may be possible to select lines that combine higher GPC and acceptable yield level, but later in maturity and taller in plant height. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

Chromosome locations of leaf rust resistance genes in selected tetraploid wheats through substitution lines

Langdon durum D-genome disomic substitution lines were used to study the chromosome locations of adult-plant leaf rust resistance genes identified from tetraploid wheat accessions. The accessions are 104 (Triticum turgidum subsp. dicoccum var. arras) and 127 (T. turgidum subsp. durum var. aestivum). The complete sets of the substitution lines were crossed as female parents with the accessions and F₁ double monosomic individuals selected at metaphase I. Segregating F₂ individuals were inoculated during the flag leaf stage with pathotype UVPrt2 of Puccinia triticina. The substitution analysis involving accession 104 showed that the gene for leaf rust resistance is located on chromosome 6B. The analysis with accession 127 indicated that chromosome 4A carries a gene for leaf rust resistance. The two novel genes are temporarily designated as Lrac104 and Lrac127, respectively from accessions 104 and 127.     

Wheat pre-breeding using wild progenitors

To facilitate the use of wheat wild relatives in conventional breedingprograms, a wheat pre-breeding activity started at ICARDA in 1994/1995season. Preliminary results of gene introgression from wild diploidprogenitors, Triticum urartu, T. baeoticum, Aegilops speltoides andAe. tauschii and tetraploid T. dicoccoides are described. Crosseswith wild diploid Triticum spp. yielded high variation in plant andspike morphology. Synthetic hexaploids were produced from crosses of alocal durum wheat landrace `Haurani' with two Ae. tauschiiaccessions. Both Ae. tauschii accessions carry hybrid necrosis allelesthat gave necrotic plant phenotypes in crosses with some bread wheats.Backcross progenies with agronomical desirable traits, i.e. high spikeproductivity, short plant stature, earliness, drought tolerance and highproductive tillering, were identified in crosses of durum wheat with wild Triticum spp. and in a cross of one of the hexaploid synthetics with alocally adapted bread wheat cv. `Cham 6'. Resistance to yellow rust wasfound in durum wheat crosses with the three wild Triticum spp. andAe. speltoides and leaf rust resistance was identified in crosses withT. baeoticum and Ae. speltoides. The results show that wheatimmediate progenitors may be a valuable and readily accessible source ofnew genetic diversity for wheat improvement.     

Use of the gene pools of Triticum turgidum ssp. dicoccoides and Aegilops squarrosa for the breeding of common wheat (T. aestivum), through chromosome-doubled hybrids

Summary Triticum turgidum ssp. dicoccoides (wild emmer wheat, AABB, 2n=28) and Aegilops squarrosa (goat grass, DD, 2n=14) comprise a rich reservoir of valuable genetic material, which could be useful for the breeding of common wheat (T. aestivum, AABBDD, 2n=42). Many accessions of both wild species, most of them selected for resistance to stripe rust, were used to make amphiploids. Two strategies were applied: (1) the production of autopolyploid cytotypes of the wild species, followed by hybridisation, and (2) the production of allotriploid interspecific hybrids, followed by doubling of the number of chromosomes. The first route was unsuccessful because of failure of the crosses between the autopolyploid cytotypes, possibly due to incongruity between the two species and to reduced fertility in the autopolyploid cytotypes. The second route yielded the desired synthetic hexaploids. However, the rate of success of the crosses was low and there were great differences between years, and within years between crosses. Embryo rescue was applied to obtain the primary hybrids (2n=21), which were highly sterile and had on average 0.3 bivalents and 20.4 univalents per pollen mother cell. Various abnormalities were recorded. Doubling of the number of chromosomes sometimes occurred spontaneously or was brought about by colchicine treatment. The large scale of the interspecific hybridisation programme ensured that one-third of the female and one-sixth of the male accessions were represented in the synthetic hexaploids.