Resistance to stripe rust in Triticum turgidum,T. tauschii and their synthetic hexaploids

Resistance to stripe rust (caused by Puccinia striiformis Westend.) of 34 Triticum turgidum L. var.durum, 278 T. tauschii, and 267 synthetic hexaploid wheats (T. turgidum x T. tauschii) was evaluated at the seedling stage in the greenhouse and at the adult-plant stage at two field locations. Mexican pathotype 14E14 was used in all studies. Seedling resistance, expressed as low infection type, was present in all three species. One hundred and twenty-eight (46%) accessions of T. tauschii, 8 (23%) of T. turgidum and 31 (12%) of synthetic hexaploid wheats were highly resistant as seedlings. In the field tests, resistance was evaluated by estimating area under disease progress curve (AUDPC). Synthetic hexaploid wheats showed a wide range of variability for disease responses in both greenhouse and field tests, indicating the presence of a number of genes for resistance. In general, genotypes with seedling resistance were also found to be resistant as adult plants. Genotypes, which were susceptible or intermediate as seedlings but resistant as adult plants, were present in both T. turgidum and the synthetic hexaploids. Resistances from either T. turgidum or T. tauschii or both were identified in the synthetic hexaploids in this study. These new sources of resistance could be incorporated into cultivated hexaploid wheats to increase the existing gene pool of resistance to stripe rust.     

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.     

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.     

Expression of morphological and flowering time variation through allopolyploidization: an empirical study with 27 wheat synthetics and their parental Aegilops tauschii accessions

It was recently shown that allopolyploidy brings novel epistatic interactions to genes belonging to different genomes. However, systematic studies of the phenotypic relationships between synthetic hexaploid wheats and their parental lines have not been conducted. In this study, 27 synthetic hexaploid wheats were produced by crossing the tetraploid wheat cultivar ‘Langdon’ with 27 accessions of Aegilops tauschii. Variations in 20 morphological and flowering traits were analysed in both the synthetic wheat lines and the parental Ae. tauschii accessions. The 20 traits exhibited large variations in the wheat lines. For many of the traits, the degree of variation in the parental accessions was reduced in the hexaploid derivatives. Principal component analysis of floret‐related traits divided the Ae. tauschii accessions into two subspecies, ssp. tauschii and ssp. strangulata, but this parental pattern of subspecific division was not detectable in the hexaploids. Our results suggest that the ‘Langdon’ genome may have an alleviating effect on the expression of D‐genome‐derived variations in derived synthetics.     

Molecular tagging of a stripe rust resistance gene in <Emphasis Type="Italic">Aegilops tauschii</Emphasis>

Stripe rust, caused by Puccinia striiformis f. sp. tritici (PST), is one of the most important diseases of common wheat (Triticum aestivum L.). China has the largest stripe rust epidemic areas in the world and yield losses can be large. Aegilops tauschii Coss, the D-genome progenitor of common wheat, includes two subspecies, tauschii and strangulata (Eig) Tzvel. The ssp. strangulata accession AS2388 is highly resistant to the prevailing physiological races of PST in China, and possesses a single dominant gene for stripe rust resistance. In order to tag this gene, AS2388 was crossed with the highly susceptible ssp. tauschii accession AS87. The parents, F₂ plants, and F_2:3 families were tested at adult plant stage in field trials with six currently prevailing races. Simple sequence repeat (SSR) primers were used to identify molecular markers linked to the resistance gene. SSR markers Xwmc285 and Xwmc617 were linked to the resistance gene on chromosome arm 4DS flanking it at 1.7 and 34.6 cM, respectively. Based on the chromosomal location, this gene temporarily designated as YrAS2388 is probably novel. The resistance in Ae. tauschii AS2388 was partially expressed in two newly developed synthetic hexaploid backgrounds.     

Global status of wheat leaf rust caused by <Emphasis Type="Italic">Puccinia triticina</Emphasis>

Leaf rust caused by Puccinia triticina is the most common and widely distributed of the three wheat rusts. Losses from leaf rust are usually less damaging than those from stem rust and stripe rust, but leaf rust causes greater annual losses due to its more frequent and widespread occurrence. Yield losses from leaf rust are mostly due to reductions in kernel weight. Many laboratories worldwide conduct leaf rust surveys and virulence analyses. Most currently important races (pathotypes) have either evolved through mutations in existing populations or migrated from other, often unknown, areas. Several leaf rust resistance genes are cataloged, and high levels of slow rusting adult plant resistance are available in high yielding CIMMYT wheats. This paper summarizes the importance of leaf rust in the main wheat production areas as reflected by yield losses, the complexity of virulence variation in pathogen populations, the role cultivars with race-specific resistance play in pathogen evolution, and the control measures currently practiced in various regions of the world.     

Transfer of rust resistance from Aegilops ovata into bread wheat (Triticum aestivum L.) and molecular characterisation of resistant derivatives

An interspecific cross was made to transfer leaf rust and stripe rust resistance from an accession of Aegilops ovata (UUMM) to susceptible Triticum aestivum (AABBDD) cv. WL711. The F₁was backcrossed to the recurrent wheat parent, and after two to three backcrosses and selfing, rust resistant progenies were selected. The C-banding study in a uniformly leaf rust and stripe rust resistant derivative showed a substitution of the 5M chromosome of Ae. ovata for 5D of wheat. Analysis of rust resistant derivatives with mapped wheat microsatellite makers confirmed the substitution of 5M for 5D. Some of these derivatives also possessed one or more of the three alien translocations involving 1BL, 2AL and 5BS wheat chromosomes which could not be detected through C-banding. A translocation involving 5DSof wheat and the substituted chromosome 5M of Ae. ovata was also observed in one of the derivatives. Susceptibility of this derivative to leaf rust showed that the leaf rust resistance gene(s) is/are located on short arm of 5M chromosome of Ae. ovata. Though the Ae. ovatasegment translocated to 1BL and 2AL did not seem to possess any rust resistance gene, the alien segment translocated to 5BS may also possess gene(s) for rust resistance. The study demonstrated the usefulness of microsatellite markers in characterisation of interspecific derivatives. This revised version was published online in August 2006 with corrections to the Cover Date.     

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.     

Development of diversity array technology (DArT) markers for assessment of population structure and diversity in Aegilops tauschii

Aegilops tauschii Coss. is the D-genome donor to hexaploid bread wheat (Triticum aestivum) and is the most promising wild species as a genetic resource for wheat breeding. To study the population structure and diversity of 81 Ae. tauschii accessions collected from various regions of its geographical distribution, the genomic representation of these lines were used to develop a diversity array technology (DArT) marker array. This Ae. tauschii array and a previously developed DArT wheat array were used to scan the genomes of the 81 accessions. Out of 7500 markers (5500 wheat and 2000 Ae. tauschii), 4449 were polymorphic (3776 wheat and 673 Ae. tauschii). Phylogenetic and population structure studies revealed that the accessions could be divided into three groups. The two Ae. tauschii subspecies could also be separately clustered, suggesting that the current taxonomy might be valid. DArT markers are effective to detect very small polymorphisms. The information obtained about Ae. tauschii in the current study could be useful for wheat breeding. In addition, the new DArT array from this Ae. tauschii population is expected to be an effective tool for hexaploid wheat studies.     

CIMMYT小麦材料的苗期和成株抗叶锈病鉴定

选择来自CIMMYT的103个小麦品种(系)及35个含有已知抗叶锈基因的对照品种,苗期接种17个中国小麦叶锈菌小种以鉴定这些品种中可能含有的抗叶锈病基因, 并于2008-2009和2009-2010连续2年对这些材料进行成株抗叶锈病鉴定。通过苗期鉴定结合系谱分析和分子检测,在46个品种中共鉴定出Lr26、Lr34、Lr42和Lr47等4个抗叶锈病基因,其中9个品种携带Lr26基因,28个品种含有成株抗叶锈病基因Lr34基因,Lr42可能存在于11个品种中,还有2个材料可能含有Lr47,其他57个品种(系)对供试的15个小种多数表现为高抗,没有鉴定出已知的抗叶锈病基因。通过两年的田间抗叶锈病鉴定共筛选出46个表现慢锈的品种。苗期和田间结果表明,CIMMYT材料中含有丰富的对我国叶锈菌小种有效的苗期和成株抗叶锈基因,这些材料均可应用于我国小麦的抗叶锈病育种。     

Applicability of Aegilops tauschii drought tolerance traits to breeding of hexaploid wheat

Few genes are available to develop drought-tolerant bread wheat (Triticum aestivum L.) cultivars. One way to enhance bread wheat’s genetic diversity would be to take advantage of the diversity of wild species by creating synthetic hexaploid wheat (SW) with the genomic constitution of bread wheat. In this study, we compared the expression of traits encoded at different ploidy levels and evaluated the applicability of Aegilops tauschii drought-related traits using 33 Ae. tauschii accessions along with their corresponding SW lines under well-watered and drought conditions. We found wide variation in Ae. tauschii, and even wider variation in the SW lines. Some SW lines were more drought-tolerant than the standard cultivar Cham 6. Aegilops tauschii from some regions gave better performing SW lines. The traits of Ae. tauschii were not significantly correlated with their corresponding SW lines, indicating that the traits expressed in wild diploid relatives of wheat may not predict the traits that will be expressed in SW lines derived from them. We suggest that, regardless of the adaptability and performance of the Ae. tauschii under drought, production of SW could probably result in genotypes with enhanced trait expression due to gene interactions, and that the traits of the synthetic should be evaluated in hexaploid level.     

Race non-specific resistance to rust diseases in CIMMYT spring wheats

Rust diseases continue to cause significant losses to wheat production worldwide. Although the life of effective race-specific resistance genes can be prolonged by using gene combinations, an alternative approach is to deploy varieties that posses adult plant resistance (APR) based on combinations of minor, slow rusting genes. When present alone, APR genes do not confer adequate resistance especially under high disease pressure; however, combinations of 4?C5 such genes usually result in ??near-immunity?? or a high level of resistance. Although high diversity for APR occurs for all three rusts in improved germplasm, relatively few genes are characterized in detail. Breeding for APR to leaf rust and stripe rust in CIMMYT spring wheats was initiated in the early 1970s by crossing slow rusting parents that lacked effective race-specific resistance genes to prevalent pathogen populations and selecting plants in segregating populations under high disease pressure in field nurseries. Consequently most of the wheat germplasm distributed worldwide now possesses near-immunity or adequate levels of resistance. Some semidwarf wheats such as Kingbird, Pavon 76, Kiritati and Parula show high levels of APR to stem rust race Ug99 and its derivatives based on the Sr2-complex, or a combination of Sr2 with other uncharacterized slow rusting genes. These parents are being utilized in our crossing program and a Mexico-Kenya shuttle breeding scheme is used for selecting resistance to Ug99. High frequencies of lines with near-immunity to moderate levels of resistance are now emerging from these activities. After further yield trials and quality assessments these lines will be distributed internationally through the CIMMYT nursery system.     

Wheat leaf rust resistance gene Lr59 derived from Aegilops peregrina

An Aegilops peregrina (Hackel in J. Fraser) Maire & Weiller accession that showed resistance to mixed leaf rust ( Puccinia triticina Eriks.) inoculum was crossed with, and backcrossed to, hexaploid wheat ( Triticum aestivum L.). During backcrossing a chromosome segment containing a leaf rust resistance gene (here designated Lr59 ) was spontaneously translocated to wheat chromosome 1A. Meiotic, monosomic and microsatellite analyses suggested that the translocated segment replaced most of, or the complete, 1AL arm, and probably resulted from centromeric breaks and fusion. The translocation, of which hexaploid wheat line 0306 is the appropriate source material, provided seedling leaf rust resistance against a wide range of South African and Canadian pathotypes.     

Identification of quantitative trait loci controlling grain size and shape in the D genome of synthetic hexaploid wheat lines

Synthetic hexaploid wheat is an effective genetic resource for transferring agronomically important genes from Aegilops tauschii to common wheat. Wide variation in grain size and shape, one of the main targets for wheat breeding, has been observed among Ae. tauschii accessions. To identify the quantitative trait loci (QTLs) responsible for grain size and shape variation in the wheat D genome under a hexaploid genetic background, six parameters related to grain size and shape were measured using SmartGrain digital image software and QTL analysis was conducted using four F₂ mapping populations of wheat synthetic hexaploids. In total, 18 QTLs for the six parameters were found on five of the seven D-genome chromosomes. The identified QTLs significantly contributed to the variation in grain size and shape among the synthetic wheat lines, implying that the D-genome QTLs might be at least partly functional in hexaploid wheat. Thus, synthetic wheat lines with diverse D genomes from Ae. tauschii are useful resources for the identification of agronomically important loci that function in hexaploid wheat.     

‘CPAN 1842’: a new source of adult plant leaf rust resistance in bread wheat

There is worldwide interest in adult plant resistance (APR) because of greater durability of APR to the cereal rusts. Peruvian bread wheat genotype ‘CPAN (Coordinated Project Accession Number) 1842’ (LM 50–53) has shown leaf rust resistance in disease screening nurseries since its introduction in 1977. However, it is susceptible at the seedling stage to several Puccinia triticina (Pt) pathotypes including the widely prevalent 77‐5 (121R63‐1) that infects bread wheat. Inheritance studies showed that CPAN 1842 carried a dominant gene for APR to pathotype 77‐5, which was different from Lr12, Lr13, Lr22a, Lr34, Lr35, Lr37, Lr46, Lr48, Lr49 and Lr68, based on the tests of allelism; and from Lr67, based on genotyping with the closely linked SSR marker cfd71. This gene should also be different from Lr22b as the latter is totally ineffective against pathotype 77‐5. CPAN 1842 therefore appears to be a new promising source of leaf rust resistance. Also having resistance to stem rust and stripe rust, this line can contribute to breeding for multiple rust resistances in wheat.     

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.     

Discovery, characterisation and mapping of wheat leaf rust resistance gene Lr71

A temporarily designated gene LrARK12c (identified from spelt wheat cv. Altgold Rotkorn) with an intermediate low infection type was found effective against prevalent Australian Puccinia triticina (Pt) pathotypes. The gene was mapped to chromosome 1B between markers Xgwm18 and Xbarc187, with linkage distances of 1.0 and 1.3 cM, respectively. While it was not possible to assign a definitive chromosomal arm location to LrARK12c, it maps close to the centromere based on physical mapping of SSR marker loci using deletion lines. Other genes conferring resistance to Pt in chromosome 1B include Lr33, Lr44 and Lr46. Genetic analysis showed that LrARK12c and Lr44 are genetically independent. Comparisons of markers linked to LrARK12c and Lr46 indicate that Lr46 should be well distal to the centromere. Lr33 is not effective in the seedling stage with Australian Pt pathotypes, therefore question of possible allelism of LrARK12c and Lr33 cannot be resolved using Australian Pt pathotypes. Genetic studies, chromosome mapping and allelism tests indicated that LrARK12c is a new and genetically independent leaf rust resistance locus, and hence it was designated Lr71 in accordance with the rules of wheat gene nomenclature.     

Detection of quantitative trait loci for leaf rust resistance in wheat––T. timopheevii/T. tauschii introgression lines

Advanced backcross QTL analysis was used to identify QTLs for seedling and adult plant resistance to leaf rust in introgression lines derived from a cross between the spring wheat cultivar ‘Saratovskaya 29’ and a synthetic allopolyploid wheat (T. timopheevii/T. tauschii). F₂ mapping populations involving two backcross selections (‘BC5’ and ‘BC9’ lines) were genotyped with microsatellite markers. Two significant QTL for adult plant resistance were identified in line ‘BC5’: one on chromosome 2B, but originating from chromosome 2G, explained 31% of the trait variance. The other, derived from T. tauschii and mapped to the short arm of chromosome 2D explained 19% of the trait variance. In the second line, one major seedling and adult plant resistance QTL was identified on chromosome 2B. Both QTL co-located to the same marker interval. Such introgression lines, resulting from the reconstruction of common wheat genome, are of interest both as initial material for breeding and improvement of current cultivars, and as a resource for the study of the interaction and transformation of genomes.     

Ph I-induced transfer of leaf and stripe rust-resistance genes from Aegilops triuncialis and Ae. geniculata to bread wheat

Leaf and stripe rusts are severe foliar diseases of bread wheat. Recently, chromosomes 5M^g from the related species Aegilops geniculata that confers resistance to both leaf and stripe rust and 5U^t from Ae. triuncialis conferring resistance to leaf rust have been transferred to bread wheat in the form of disomic DS5M^g(5D) and DS5U^t(5A) chromosome substitution lines. The objective of this study was to shorten the alien segments in these lines using Ph ^I-mediated, induced homoeologous recombination. Putativerecombinants were evaluated for their rust resistance, and by genomic in situ hybridization and microsatellite analyses. One agronomically useful wheat-Ae. geniculata recombinant resistant to leaf and stripe rust was identified that had only a small terminal segment of the 5M^gL arm transferred to the long arm of an unidentified wheat chromosome. This germplasm can be used directly in breeding programs. Only one leaf rust-resistant wheat-Ae. triuncialis recombinant, which consists of most of the complete 5U^t chromosome with a small terminal segment derived from 5AS, was identified. This germplasm will need further chromosome engineering before it can be used in wheat improvement. This revised version was published online in August 2006 with corrections to the Cover Date.     

Stripe rust resistance and genes in Chinese wheat cultivars and breeding lines

Stripe rust, caused by Puccinia striiformis f. sp. tritici (Pst), is one of the most important diseases on wheat in China. To assess resistance in wheat cultivars and breeding lines in China, 330 leading cultivars and 164 advanced breeding lines were evaluated with stripe rust. In the greenhouse tests, seedlings of the entries were inoculated separately with several Pst pathotypes. In the field tests, the entries were evaluated for stripe rust resistance in Yangling, Shaanxi Province artificially inoculated and in Tianshui, Gansu Province under natural infection of Pst. The oversummering/wintering and spring epidemic zones of resistance genes were postulated using molecular markers for Yr5, Yr9, Yr10, Yr15, Yr17, Yr18, and Yr26, in combination with resistance spectra. Out of the 494 wheat entries, 16 (3.24 %) entries had all-stage resistance (ASR) in all race tests, 99 (20.04 %) had adult-plant resistance (APR), 28 (5.67 %) were considered to have slow-rusting (SR), and 351 (71.05 %) were susceptible to one or more races in both seedling and adult-plant stages. Advanced breeding lines had a higher percentage (37.2 %) of resistant entries (The sum of ASR, APR and SR) than leading cultivars (24.85 %). Among the epidemic regions, southern Gansu had a higher percentage of resistant entries than any other regions. Based on stripe rust reactions and molecular markers, two cultivars were found to possibly have Yr5 while no entries have Yr10 or Yr15. Resistance genes Yr9, Yr17, Yr18, and Yr26 were found in 134 (29.4 %), 45 (9.1 %), 10 (2 %), and 15 (3 %) entries, respectively.