Suppression/expression of resistance to stripe rust in synthetic hexaploid wheat (Triticum turgidum×T. tauschii)

Seventy-four hexaploid wheats, synthesized by either crossing resistantTriticum turgidum L. var.durum with susceptible/intermediateT. tauschii or susceptible/intermediateT. turgidum with resistantT. tauschii, and their parents were evaluated as seedlings in the greenhouse and as adult-plants at two field locations in Mexico for resistance to pathotype 14E14 of stripe (or yellow) rust (caused byPuccinia striiformis Westend). The seedlings of different synthetic hexaploids showed high phenotypic diversity for resistance. However, the resistance level of only 15 of the 74 synthetic hexaploid wheats were similar to the low infection types of the respective donor parents. The remaining synthetic wheats displayed either intermediate or high infection types. A similar result was also obtained in field tests, where only 18 synthetic hexaploids were resistant as adult-plants. In general, genotypes with seedling resistance were also resistant as adult-plants. A few synthetic hexaploids, which displayed intermediate or susceptible infection types as seedlings were resistant as adult-plants, indicating that additional genes for adult-plant resistance were also present. The fact that resistance of some donor parents was not expressed, or only partially expressed, in a synthetic hexaploid background suggests the presence of suppressor genes in the both the A or B, and D genomes ofT. turgidum andT. tauschii, respectively. The resistance of a donor parent was expressed in a synthetic hexaploid only if the corresponding suppressor was absent in the second parent. Moreover, the suppressors appeared to be resistance gene specific.     

Assessment of genetic diversity in synthetic hexaploid wheats and their Triticum dicoccum and Aegilops tauschii parents using AFLPs and agronomic traits

Synthetic hexaploid wheats are of interest to wheat breeding programs, especially for introducing new genes that confer resistance to biotic and abiotic stresses. A group of 54 synthetic hexaploid wheats derived from crosses between emmer wheat(Triticum dicoccum, source of the A and B genomes) and goat grass (Aegilops tauschii, D genome donor) were investigated for genetic diversity. Using the AFLP technique, dendrograms revealed clear grouping according to geographical origin for the T. dicoccum parents but no clear groups for the Ae. tauschii parents. The geographical clustering of the T. dicoccum parents was also reflected in the dendrogram of their derived synthetic hexaploids. Diversity of the T. dicoccum parents and their derived synthetic hexaploids was further evaluated by measuring 18morphological and agronomic traits on the plants. Clustering based on morphological and agronomic data also reflected geographical origin. However, comparison of genetic distances obtained from AFLP and agronomic data showed no correlation between the two diversity measurements. Nevertheless, similarities among major clusters with the two systems could be identified. Based on percentage of polymorphic markers, the synthetic hexaploids had a considerably higher level of AFLP diversity (39%) than normally observed in cultivated hexaploid wheat (12–21%). This suggests that synthetic hexaploid wheats can be used to introduce new genetic diversity into the bread wheat gene pool. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of resistance to Tilletia indica (Mitra) in synthetic hexaploid wheat ×Triticum aestivum crosses

Inheritance of resistance to Karnal bunt was investigated in the crosses of four resistant synthetic hexaploid wheats (SH; Triticum turgidum×T. tauschii) and two susceptible T. aestivum cultivars. The resistance was dominant or partly dominant over susceptibility. The SH cultivars Chen/T. tauschii (205) and Chen/T. tauschii (224) have single dominant resistance genes which could be allelic to each other. ‘Altar 84’/T. tauschii (219) appeared to have two dominant genes for resistance. ‘Duergand’T. tauschii (214) possessed two complementary dominant genes for resistance. The work is being extended to involve diverse Karnal bunt-resistant SH and bread wheat cultivars.     

Suppression of stripe rust and leaf rust resistances in interspecific crosses of wheat

Stripe rust and leaf rust caused by Puccinia striiformis (Ps) Westend. and P. triticina (Pt) Eriks., respectively, are important foliar diseases of wheat worldwide. Breeding resistant wheat cultivars is the preferred strategy to control these diseases. Genes for resistance when introgressed from alien species or wheats of lower ploidy are frequently diluted effectiveness in the hexaploid wheat background or are completely suppressed. The objective of this study was to examine the expression of wheat stripe rust and leaf rust resistances derived from wild emmer wheat and Aegilops tauschii when combined in synthetic hexaploid lines. Eight amphidiploid wheat lines, synthesized by crossing five tetraploid wheats (AABB), viz. Triticum carthlicum var. darginicum, T. carthlicum var. fuligioscum, T. dicoccoides var. fuligioscum, T. durum with five lines of Ae. tauschii (DD), were evaluated in the seedling stage for resistance to five pathotypes of stripe rust caused by Ps and four pathotypes of leaf rust caused by Pt. Resistance in one or both parents was frequently suppressed in synthetic hexaploid lines, indicating the presence of suppressor genes in both Ae. tauschii and T. carthlicum var. darginicum. Specific suppression of resistance genes in the parental genotypes and to pathotypes of Ps and Pt were also observed. The presence and specificity of the suppressors for rust resistance obtained in this study provides useful knowledge for developing cultivars resistant to both rusts utilizing such genetic stocks in wheat breeding programs.     

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.     

Evaluation of seedling and adult plant resistance in European wheat cultivars to Australian isolates of <Emphasis Type="Italic">Puccinia striiformis</Emphasis> f. sp. <Emphasis Type="Italic">tritici</Emphasis>

A set of 105 European wheat cultivars was assessed for seedling resistance and adult plant resistance (APR) to stripe (yellow) rust in greenhouse and field tests with selected Australian isolates of Puccinia striiformis f. sp. tritici (Pst). Twelve cultivars were susceptible to all pathotypes, and among the remainder, 11 designated seedling genes (Yr1, Yr3, Yr4, Yr6, Yr7, Yr9, Yr17, Yr27, Yr32, YrHVII and YrSP) and a range of unidentified seedling resistances were detected either singly or in combination. The identity of seedling resistance in 43 cultivars could not be determined with the available Pst pathotypes, and it is considered possible that at least some of these may carry uncharacterised seedling resistance genes. The gene Yr9 occurred with the highest frequency, present in 19 cultivars (18%), followed by Yr17, present in 10 cultivars (10%). Twenty four cultivars lacked seedling resistance that was effective against the pathotype used in field nurseries, and all but two of these displayed very high levels of APR. While the genetic identity of this APR is currently unknown, it is potentially a very useful source of resistance to Pst. Genetic studies are now needed to characterise this resistance to expedite its use in efforts to breed for resistance to stripe rust. Colin R. Wellings seconded from NSW Department of Primary Industries.     

Starch characterisation and variability in GBSS loci of synthetic hexaploid wheats and their durum and <Emphasis Type="Italic">Aegilops tauschii</Emphasis> parents

Greater variability in starch properties is found in lower ploidy wheats than in commercial hexaploid wheats. This paper reports on the starch properties and variability in granule bound starch synthase (GBSS) loci of 17 diploid (Aegilops tauschii) and 12 tetraploid (durums) potential progenitors of wheat, compared with 29 synthetic hexaploid wheats produced from such progenitors. Starch properties examined were granule size distribution, swelling power, amylose content, gelatinisation and amylose-lipid dissociation properties. A PCR screening method was able to detect the presence or absence of each of the three GBSS genes. It also detected polymorphisms in eight diploids and nine hexaploids, all displaying the same 25 bases deletion in the D genome allele of GBSS. Two tetraploids and five hexaploids were null 4A for GBSS. There was little difference in the amylose contents and amylose-lipid dissociation peak temperatures of the synthetic hexaploids and the lower ploidy wheats. The synthetic hexaploids showed intermediate swelling power values with the durums giving the highest swelling powers. The durums also had higher B granule contents than the A. tauschii accessions, but not as high as the synthetics. However, the A. tauschii samples gave the highest gelatinisation peak temperatures. The presence of the null 4A mutation was positively correlated with swelling power, amylose content and DSC measurements. The new smaller D genome allele of GBSS was associated with slightly higher swelling power. These results confirm the value of wheat progenitor lines as sources of new starch properties for hexaploid wheat. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Responses of Israeli wild emmers to selected Australian pathotypes of Puccinia species

Summary Seedling responses to one Australian isolate of each of the stripe rust, stem rust and leaf rust pathogens were determined for 541 accessions of T. dicoccoides collected from 23 locations in Israel. Resistance to stripe rust was more frequent than resistance to stem rust. Stripe rust responses showed a wide range of variability indicative of a number of genes for resistance. Comparison of the present stem rust data and that reported for the same accessions tested in Israel indicated that different genes were operating in each country. Only moderately resistant responses to stem rust were obtained. This level of resistance is probably inadequate for transfer to commercial wheat cultivars. We found no potentially useful seedling resistance to leaf rust.     

Use of Triticum tauschii to improve yield of wheat in low-yielding environments

Triticum tauschii (Coss.) Schmal. is an ancestor of bread wheat (T. aestivum). This species has been widely used as a source ofsimply-inherited traits, but there are few reports of yield increases due tointrogression of genes from this species. Selections from F₂-derivedlines of backcross derivatives of synthetic hexaploid wheats (T.turgidum / T. tauschii) were evaluated for grain yield in diverseenvironments in southern Australia. Re-selections were made in theF₆ generation and evaluated for grain yield, yield componentsincluding grain weight, and grain growth characters in diverse environmentsin southern Australia and north-western Mexico. Re-selection was effectivein identifying lines which were higher yielding than the recurrent parent,except in full-irrigation environments. Grain yields of the selectedderivatives were highest relative to the recurrent parent in thelowest-yielding environments, which experienced terminal moisture deficitand heat stress during grain filling. The yield advantage of the derivativesin these environments was not due to a change in anthesis date orgrain-filling duration, but was manifest as increased rates of grain-filling andlarger grains, indicating that T. tauschii has outstanding potential forimproving wheat for low-yielding, drought-stressed environments.     

Stripe rust reactions of Australasian wheats

Summary Seedling and field reactions to four European races of stripe rust were determined for 254 wheat cultivars, mostly from Australia and New Zealand. Numerical analyses of the data employed a divisive classification procedure with termal reallocation to form 10 groups which were then further classified, ordinated and diagnosed by several procedures. The groups formed ranged from one comprising four wheats with low seedling reactions to all four rust strains and mean field rust of only 1.0%, to one comprising 105 wheats with high seedling reactions and mean field rust of 64.7%. Groups of cultivars with intermediate levels of rust resistance were classified both on relative level of overall rust and on differential reactions to rust strains. One group of 13 cultivars had moderately high seedling reactions but averaged only 1.5% rust in the field.Seedling response to gibberellic acid (GA) was also measured and diagnosed as an external attribute, along with grain type and region of origin. Principal co-ordinate analysis revealed that greater resistance to stripe rust was associated with the insensitivity to GA typical of semidwarf wheats and with an origin in northern Australia, where semidwarf wheats have been most used in the breeding program. In terms of numerical analysis of disease data in plant breeding programs, the study highlighted the advantages of using divisive classification with terminal reallocation of group members.     

Resistance to Karnal Bunt (Tilletia indica Mitra) in Synthetic Hexaploid Wheats Derived from Triticum turgidum×T. tauschii

Synthetic hexaploids (SH) developed at the International Maize and Wheat Improvement Center (CIMMYT), involving four Triticum turgidum and nine T. tauschii parents, were evaluated for resistance to Karnal bunt (KB) (Tilletia indica Mitra) during three crop seasons over three years at Ciudad Obregon, Sonora, Mexico. Ten tillers of each SH at boot stage, taken at random, were injected with a suspension of sporidia in water (10,000 spores/ml of water). At maturity the inoculated spikes were threshed individually and evaluated for the percentage KB-infected grains. Based on the mean KB score of each entry for three seasons, 49 % of the SH were immune (0 % infection) to KB. Highly resistant expressions characterized the SH which appeared to be influenced by the resistance of their T. turgidum and/or T. tauschii parents. The overall mean infection of the SH wheats was 0.24 % compared to 56.14 % in the susceptible bread wheat check cultivat ‘WL711’. Transfer of KB resistance genes from SH wheats into bread wheat is currently underway at CIMMYT.     

Application of Triticum monococcum for the improvement of triticale resistance to leaf rust (Puccinia triticina)

The aim of this work was to evaluate the leaf rust resistance introduced into introgressive triticale lines with Triticum monococcum genes, and to study the expression of these genes at the hexaploid level. The introgressive lines were developed by incorporating diploid wheat (T. monococcum s.s.) genes into hexaploid triticale LT 522/6 using the synthetic allotetraploid T. monococcum/Secale cereale (A^mA^mRR) as a bridging form. A group of 44 those lines, parental stocks and check cultivars were inoculated at the seedling stage (in a greenhouse) and at the adult‐plant stage (in the field) with four pathotypes of Puccinia triticina. At the seedling stage the assessment of infection type showed that four lines had resistance to all pathotypes as high as in the T. monococcum donor. Adult plant examinations showed some introgressive lines with complete resistance and also lines with partial resistance, expressed in area under the disease progress curve (AUDPC) calculations as slow rusting. Some lines comprise low AUDPC with complete resistance at seedling stage.     

Resistance to stripe rust in durum wheats, A-genome diploids, and their amphiploids

Stripe rust (caused by Puccinia striiformis Westend.) is a wheat disease of worldwide importance. Seedlings of 75 accessions of Triticum boeoticum, 12 of T. monococcum, 16 of T. urartu, 230 of durum wheat (T. turgidum L. var. durum), and 128 amphiploids (genome AAAABB) involving the crosses of the three diploid species (AA) with T. turgidum (AABB) were evaluated in the greenhouse for their reaction to P. striiformis race 14E14. Durum wheats and the amphiploids were also evaluated at two field locations in Mexico with the same race for their adult plant response. Resistant seedling reactions (infection types: 0-3 on a 0-9 scale) were seen for 10 (13%) accessions of T. boeticum, 19 (8%) accessions of T. turgidum and 32 (25%) amphiploids. The remaining accessions were either moderately resistant (ITs 4-6) or susceptible (ITs 7-9). The three amphiploids derived from the crosses of seedling resistant T. boeoticum and T. turgidum, were resistant as seedlings. Among the 51 amphiploids involving one resistant parent, 29 were resistant and the remaining 22 displayed intermediate to susceptible reactions. Suppressors for resistance were common in the A and AB genomes and suppression was resistance gene specific. Forty-five (20%) durums showed adequate field resistance (relative AUDPC <10% of the susceptible check ‘Morocco’). These included the 19 seedling resistant durums. Presence of genes involved in adult plant resistance was evident, because 26 of the remaining adult plant resistant durums had displayed intermediate-susceptible seedling reactions. Though the seedling reactions of the amphiploids varied from low to high, all involving the adult plant resistant durums possessed adequate field resistance. The resistant, newly produced, AAAABB amphiploids are useful genetic resources for stripe rust resistance which could be transferred to the cultivated T. turgidum. This revised version was published online in July 2006 with corrections to the Cover Date.     

Evaluation of seedling and adult plant resistance to leaf rust in European wheat cultivars

Summary A set of 105 European wheat cultivars, comprising 68 cultivars with known seedling resistance genes and 37 cultivars that had not been tested previously, was tested for resistance to selected Australian pathotypes of P. triticina in seedling greenhouse tests and adult plant field tests. Only 4% of the cultivars were susceptible at all growth stages. Twelve cultivars lacked detectable seedling resistance to leaf rust, and among the remaining cultivars, 10 designated genes were present either singly or in combination. Lr13 was the most frequently detected gene, present in 67 cultivars, followed by the rye-derived gene Lr26, present in 19 cultivars. Other genes present were Lr1, Lr3a, Lr3ka, Lr10, Lr14a, Lr17b, Lr20 and Lr37. There was evidence for unidentified seedling resistance in addition to known resistance genes in 11 cultivars. Field tests with known pathotypes of P. triticina demonstrated that 57% of the cultivars carried adult plant resistance (APR) to P. triticina. The genetic identity of the APR is largely unknown. Genetic studies on selected cultivars with unidentified seedling resistances as well as all of those identified to carry APR are required to determine the number and inheritance of the genes involved, to determine their relationships with previously designated rust resistance genes, and to assess their potential value in breeding for resistance to leaf rust.     

Enhanced leaf rust resistance in wheat conditioned by resistance gene pairs with Lr13

Summary Leaf rust resistance gene Lr13 is present in many North American hard red spring wheat cultivars that have shown durable resistance to leaf rust. Fifteen pair-wise combinations of Lr13 and seedling leaf rust resistance genes were developed by intercrossing near isogenic Thatcher lines. In both seedling and adult plant tests, homozygous paired combinations of specific resistance genes with Lr13 had enhanced resistance relative to either parent to rust isolates that had intermediate avirulent infection types to the additional genes. In field tests, homozygous lines were more resistant than either parent if the additional leaf rust gene conditioned an effective level of resistance when present singly.     

具特异性状人工合成小麦的条锈病抗性改良

为了解3份具特异优良性状但高感条锈病的人工合成小麦SHW-Z1、SHW-Z2和SHW-Z4感病性的遗传特点,进行更好的育种利用。用高抗条锈病的普通小麦材料5157与上述人工合成小麦分别进行正反杂交,对6个杂交组合的亲本、F¹世代的条锈病抗性与F²代的条锈病抗感分离情况进行了分析以探究其感病性的遗传特点,结果表明：（1）本研究的普通小麦和人工合成小麦杂交后代的条锈病抗性由多对基因控制,遗传上表现出加-显效应;(2)SHW-Z1条锈病的抗性改良效果优于SHW-Z2和SHW-Z4;（3）本研究材料的条锈病抗性基因可能还受到遗传背景的影响。本研究对这3份人工合成小麦的条锈病抗性改良和育种利用提供了理论依据,同时可为相关研究提供参考。     

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.     

Novel resistance to pre-harvest sprouting in Australian wheat from the wild relative Triticum tauschii

The diploid D-genome progenitor of hexaploid wheat, Triticum tauschii (Coss.) Schmahl., was screened to identify mechanisms for resistance to pre-harvest sprouting. A number of promising mechanisms were identified, and transferred to hexaploid wheat via wide-hybridisation. One identified mechanism, an inhibitory phenolic compound present in the bracts surrounding the grain, has been shown to function effectively in synthetic hexaploid wheats. A number of seed-borne dormancy mechanisms were also identified. Expression of embryo dormancy in synthetic hexaploid wheats was demonstrated when compared with non-dormant hexaploid wheat. Effects of the seed coat on dormancy were also studied, with the seed coat of synthetic hexaploids accelerating rather than inhibiting germination. Embryo dormancy was also demonstrated in two `direct-cross' hybrids. The results suggest that a combination of the described mechanisms may produce white wheats with resistance to pre-harvest sprouting adequate for most Australian climatic conditions. This revised version was published online in August 2006 with corrections to the Cover Date.     

Monosomic analyses of stripe rust and leaf rust resistance genes in winter wheat varieties Lüquyu and Yantar

Summary A set of 21 monosomics of Novosadska Rana-1 was used to locate the rust resistance genes of Lüqiyu, a stripe rust resistant line developed by BAU and Yantar, a leaf rust resistant wheat introduced from Bulgaria. The resistance of the former to p. striiformis race C25 was conditioned by a dominant gene located on chromosome 2B, whereas that of the latter to P. recondita race CL3 was controlled by two complementary dominant genes located on chromosomes 5A and 1D, respectively. The relationship of the stripe rust resistance gene in Lüqiyu to Yr5, Yr7 or Yr _Suwon' all located on chromosome 2B is unknown. The two complementary leaf rust resistance factors in Yantar appear to be new.     

Cytogenetical studies in wheat XIX. Location and linkage studies on gene Yr27 for resistance to stripe (yellow) rust

The stripe (yellow) rust resistance gene Yr27 was located in wheat (Triticum aestivum L.) chromosome 2B and shown to be closely linked to the leaf (brown) rust resistance genes Lr13 and Lr23 in the proximal region of the short arm. Gene Yr27 was genetically independent of Lr16, which is distally located in the same arm. While Yr27 was often difficult to score in segregating seedling populations, it is apparently quite effective in conferring resistance to avirulent cultures under field conditions. The occurrence of Yr27 in Mexican wheat germplasm and the current over-dependence on Yr27 for crop protection in Asia are discussed.