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.     

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.     

Genes for Resistance to Yellow Rust in Seedlings of Wheat Cultivars from Pakistan Tested with British Isolates of Puccinia striiformis

Seedlings of 26 wheat caltivars from Pakistan were tested with 18 British races of P. striiformis. It is postulated that the race-specific genes Yr6, Yr7, Yr9 and perhaps Yr2 were present among the cultivars, and that there was other resistance not controlled by these genes.     

Sources of seedling and adult plant resistance to Puccinia striiformis f.sp. tritici in European wheats

One-hundred-and-forty-one wheat cultivars were tested at the seedling stage using up to 16 yellow rust isolates of diverse origin. Sixty-five resistance spectra were observed, including 20 spectra defined by differential cultivars with specific genes for resistance to Puccinia striiformis f.sp. tritici . Yr1 , Yr2 , Yr3 , Yr4 , Yr6 , Yr9 , Yr15 , Yr17 , Yr25 , Yr32 , and several additional sources of resistance were recognized. The resistance spectra were often conferred by Yr -genes and resistance factors with an unresolved genetic basis. All cultivars carried resistance and 27 had resistance for which no fully compatible isolate was detected. Yr15 was detected in four cultivars, a resistance from wild Emmer not previously reported in commercial wheat. There was no indication of Yr5 , Yr7 , Yr8 , Yr10 and Yr24 in any cultivar. Most cultivars were also investigated in field nurseries using up to nine isolates of Danish origin. Fifty-six cultivars displayed high or very high levels of resistance to any isolate in the field, and 18 of these showed full compatibility at the seedling stage to at least one isolate, i.e. revealing components of adult plant resistance.     

Double dose efficiency of the yellow rust resistance gene Yr17 in bread wheat lines

Yellow rust, caused by Puccinia striiformis f. sp. tritici, is one of the most severe wheat disease worldwide. Crop losses have ranged from 10% to 70% and up to 100% in extreme conditions. Eighty-two resistance genes, designated Yr, have been identified. Among them, Yr17 derived from Aegilops ventricosa and located on chromosome 2A has been widely used in wheat breeding. However, it had been overcome already. Through recombination of the Ae. ventricosa Yr17-carrying 6N^v chromosome with 2D of wheat, we introduced Yr17 onto chromosome 2D. Then, lines carrying Yr17 on both 2A and 2D were generated. Seedlings of the latter, as well as those carrying a single dose of Yr17 either on 2A or on 2D, were inoculated with virulent or avirulent strains on wheat seedlings. The different genotypes were fully susceptible for the two pathotypes that are virulent on Yr17. In the case of avirulent pathotypes, the Yr17 double dose lines were fully resistant, while those with the Yr17 gene only on either 2A or 2D had intermediate resistance reactions towards one or the other or both pathotypes.     

SSR and STS markers for wheat stripe rust resistance gene <Emphasis Type="Italic">Yr26</Emphasis>

Stripe (yellow) rust, caused by Puccinia striiformis f. sp. tritici (Pst), is one of the most devastating wheat diseases worldwide. Triticum aestivum-Haynaldia villosa 6VS/6AL translocation lines carrying the Yr26 gene on chromosome 1B, are resistant to most races of Pst used in virulence tests. In order to better utilize Yr26 for wheat improvement, we attempted to screen SSR and EST-based STS markers closely linked with Yr26. A total of 500 F₂ plants and the F_2:3 progenies derived from a cross between 92R137 and susceptible cultivar Yangmai 5 were inoculated with race CYR32. The analysis confirmed that stripe rust resistance was controlled by a single dominant gene, Yr26. Among 35 pairs of genomic SSR markers and 81 pairs of STS markers derived from EST sequences located on chromosome 1B, Yr26 was flanked by 5 SSR and 7 STS markers. The markers were mapped in deletion bins using CS aneuploid and deletion lines. The closest flanking marker loci, Xwe173 and Xbarc181, mapped in 1BL and the genetic distances from Yr26 were 1.4 cM and 6.7 cM, respectively. Some of these markers were previously reported on 1BS. Eight common wheat cultivars and lines developed from the T. aestivum-H. villosa 6VS/6AL translocation lines by different research groups were tested for presence of the markers. Five lines with Yr26 carried the flanking markers whereas three lines without Yr26 did not. The results indicated that the flanking markers should be useful in marker-assisted selection for incorporating Yr26 into wheat cultivars.     

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.     

Resistance in spelt wheat to yellow rust

Summary Seven spelt wheat accessions of different origin were hybridized with the susceptible bread wheat cultivar Taichung 29 in order to study the genetics of their resistance to yellow rust (Puccinia striiformis Westend. f. sp. tritici). One Iranian and five European accessions were found to carry Yr5 of Triticum aestivum ssp. spelta var. album, whereas a factor for resistance in the Iranian accession 415 was confirmed to be genetically distinct from Yr5. The alleles for resistance in each of the accessions studied showed a monogenic dominant mode of inheritance. Twenty-eight spelt wheat accessions, including those studied for their resistance to yellow rust, were subjected to polyacrylamide-gel-electrophoresis to study variation for gliadin storage protein patterns. Thirteen distinct patterns were revealed, implying the presence of duplicates within the studied spelt wheat collection.     

Molecular mapping and detection of the yellow rust resistance gene Yr26 in wheat transferred from Triticum turgidum L. using microsatellite markers

Yellow rust (stripe rust), caused by Puccinia striiformis Westend f. sp. tritici, is one of the most devastating diseases of wheat throughout the world. Wheat-Haynaldia villosa 6AL.6VS translocation lines R43, R55, R64 and R77, derived from the cross of three species, carry resistance to both yellow rust and powdery mildew. An F₂ population was established by crossing R55 with the susceptible cultivar Yumai 18. The yellow rust resistance in R55 was controlled by a single dominant gene, which segregated independently of the powdery mildew resistance gene Pm21 located in the chromosome 6VS segment, indicating that the yellow rust resistance gene and Pm21 are unlikely to be carried by the same alien segment. This yellow rust resistance gene was considered to beYr26, originally thought to be also located in chromosome arm 6VS. Bulked Segregation Analysis and microsatellite primer screens of the population F₂ of Yumai 18 × R55 identified three chromosome 1B microsatellite locus markers, Xgwm11, Xgwm18 and Xgwm413, closely linked to Yr26. Yr26 was placed 1.9 cM distal of Xgwm11/Xgwml8, which in turn were 3.2 cM from Xgwm413. The respective LOD values were 21 and 36.5. Therefore, Yr26 was located in the short arm of chromosome 1B. The origin and distribution of Yr26 was investigated by pedigree, inheritance of resistance and molecular marker analysis. The results indicated that Yr26 came from Triticum turgidum L. Three other 6AL.6VS translocation lines, R43, R64 and R77, also carried Yr26. These PCR-based microsatellite markers were shown to be very effective for the detection of the Yr26 gene in segregating populations and therefore can be applied in wheat breeding. This revised version was published online in July 2006 with corrections to the Cover Date.     

Leaf rust and stripe rust resistance genes Lr54 and Yr37 transferred to wheat from Aegilops kotschyi

The tendency of unpaired meiotic chromosomes to undergo centric misdivision was exploited to translocate leaf rust and stripe rust resistance genes from an Aegilops kotschyi addition chromosome to a group 2 chromosome of wheat. Monosomic and telosomic analyses showed that the translocation occurred to wheat chromosome arm 2DL. The introgressed region did not pair with the corresponding wheat 2DL telosome during meiosis suggesting that a whole arm may have been transferred. Female transmission of the resistance was about 55% whereas male transmission was strongly preferential (96%). The symbols Lr54 and Yr37 are proposed to designate the new resistance genes.     

Microsatellite marker for yellow rust resistance gene Yr5 in wheat introgressed from spelt wheat

Yellow rust of wheat caused by Puccinia striiformis f sp. tritici has been periodically epidemic and severely damaged wheat production in China and throughout the world. Breeding for resistant cultivars has been proved to be an effective way to resolve the problem. A yellow rust resistance gene, Yr5, derived from Triticum spelta shows immunity or high resistance to the most popular isolates Tiaozhong 30 and 31 in China. Establishment of DNA markers for the Yr5 gene will facilitate marker‐assisted selection and gene pyramiding in the breeding programme. Since the Yr5 gene was cytologically located on the long arm of chromosome 2B, By33, the donor of Yr5, was crossed and backcrossed with the susceptible line 441, and BC₃F₂ and BC₃F₃ segregating populations were screened for polymorphism by using 11 microsatellite primers mapped on chromosome 2B. A marker, Xgwm501‐195 bp/160 bp, was found to be linked to Yr5, with a genetic distance of 10.5‐13.3 cM.     

Stripe rust resistance gene Yr18 and its suppressor gene in Chinese wheat landraces

The slow‐rusting and mildewing gene Yr18/Lr34/Pm38/Sr57 confers partial, durable resistance to multiple fungal pathogens and has its origins in China. A number of diagnostic markers were developed for this gene based on the gene sequence, but these markers do not always predict the presence of the resistant phenotype as some wheat varieties with the gene are susceptible to stripe rust in China. We hypothesized that these varieties have a suppressor of Yr18. This study was undertaken to determine the presence of Yr18, the suppressor and/or another resistance gene in 144 Chinese wheat landraces using molecular markers and stripe rust field data. Forty‐three landraces were predicted to have Yr18 based on the presence of the markers, but had final disease severities higher than 70%, indicating that this gene may be under the influence of a suppressor. Four of these landraces, ‘Sichuanyonggang 2’, ‘Baikemai’, ‘Youmai’ and ‘Zhangsihuang’, were chosen for genetic studies. Crosses were made between the lines and ‘Avocet S’, with further crosses of Sichuanyonggang 2 ×  ‘Huixianhong’ and Sichuanyonggang 2 ×  ‘Chinese Spring’. The F₁ plants of Sichuanyonggang 2/Chinese Spring was susceptible indicating the presence of a dominant suppressor gene. The results of genetic analyses of F_2:3 and BC₁F₂ families derived from these crosses indicated the presence of Yr18, a Yr18 suppressor and another additive resistance gene. The Yr18 region in Sichuanyonggang 2 was sequenced to ensure that it contained the functional allele. This is the first report of a suppressor of Yr18/Lr34/Pm38/Sr57 gene with respect to stripe rust response.     

Molecular genetics of race non-specific rust resistance in wheat

Over 150 resistance genes that confer resistance to either leaf rust, stripe rust or stem rust have been catalogued in wheat or introgressed into wheat from related species. A few of these genes from the ‘slow-rusting’ adult plant resistance (APR) class confer partial resistance in a race non-specific manner to one or multiple rust diseases. The recent cloning of two of these genes, Lr34/Yr18, a dual APR for leaf rust and stripe rust, and Yr36, a stripe rust APR gene, showed that they differ from other classes of plant resistance genes. Currently, seven Lr34/Yr18 haplotypes have been identified from sequencing the encoding ATP Binding Cassette transporter gene from diverse wheat germplasm of which one haplotype is commonly associated with the resistance phenotype. The paucity of well characterised APR genes, particularly for stem rust, calls for a focused effort in developing critical genetic stocks to delineate quantitative trait loci, construct specific BAC libraries for targeted APR genes to facilitate robust marker development for breeding applications, and the eventual cloning of the encoding genes.     

Development and characterization of a new 1BL.1RS translocation line with resistance to stripe rust and powdery mildew of wheat

The 1BL.1RS wheat-rye translocation from Petkus rye has contributed substantially to the world wheat production. However, following the breakdown of disease resistance genes in 1RS, its importance for wheat improvement decreased. We have developed a new 1BL.1RS line, R14, by means of crossing rye inbred line L155, selected from Petkus rye to several wheat cultivars. One new gene each, for stripe rust and powdery mildew resistance, located on 1RS of the line R14, are tentatively named YrCn17 and PmCn17. YrCn17 and PmCn17 confer resistance to Puccinia striiformis f. sp. tritici pathotypes that are virulent on Yr9, and Blumeria graminis f. sp. tritici pathotypes virulent on Pm8. These two new resistances, YrCn17 and PmCn17, are now available for wheat improvement programs. The present study indicates that rye cultivars may carry yet untapped variations as potential sources of resistance.     

Evolution of virulence patterns in yellow rust races and its implications for Ereeding for resistance in wheat in Kenya

Summary Virulence patterns of yellow rust isolates collected in Kenya between 1986–1989 were compared with earlier results. The number of virulence factors per race and the range in virulence factors both increased considerably. Before 1976 races carried on average 4.5 to 5.0 virulence factors, whereas the races after 1986 had a mean of 6.5 virulence factors. The range in the number of virulence factors increased from some seven to eight in the first period to 12 in the second out of the 17 evaluated. In the period 1986–1989 another three virulence factors (2, 9 and A) were assessed. All three occurred at a high frequency.Virulence neutralizing the resistance genes Yr2, Yr2+, Yr6, Yr6+, Yr7, Yr7+, Yr8, Yr9, Yr9+ and those in the cultivars Anza (A), Strubes Dickkopf (SD) and Suwon92/Omar (SU) occurred at a high frequency, while virulence for Yr3V, Yr4+, Yr5, CV and SP (resistance in Carstens V and Spaldings Prolific resp.) were not found. The remaining three virulence factors for Yr1, 10 and 3N were rare.In the past ten years the resistance of most released cultivars became ineffective in less than six years. They were shown to carry race-specific major resistance genes such as Yr7+, Yr9+, SD and A. However, in the field, the resistance of the cultivars was not completely neutralized. A residual resistance, ranging from moderate to fairly high, was observed in all cultivars in which the major gene resistances were neutralized by corresponding virulence genes.Other wheat cultivars such as Africa Mayo, Kenya Kudu, Enkoy, Kenya Leopard, Bounty, Frontatch, Bonny and Kenya Plume appeared to keep their resistance over a condiserable period of time. They are considered to be durably resistant to the Kenyan yellow rust populations. This form of resistance, together with the residual resistance, can be recommended for use in breeding programmes.     

Reduction of <Emphasis Type="Italic">Aegilops sharonensis</Emphasis> chromatin associated with resistance genes <Emphasis Type="Italic">Lr56</Emphasis> and <Emphasis Type="Italic">Yr38</Emphasis> in wheat

The Lr56/Yr38 translocation consists primarily of alien-derived chromatin with only the 6AL telomeric region being of wheat origin. To improve its utility in wheat breeding, an attempt was made to exchange excess Ae. sharonensis chromatin for wheat chromatin through homoeologous crossover in the absence of Ph1. Translocation heterozygotes that lacked Ph1 were test-crossed with Chinese Spring nullisomic 6A tetrasomic 6B and nullisomic 6A-tetrasomic 6D plants and the resistant (hemizygous 6A) progeny were analyzed with four microsatellite markers. Genetic mapping suggested general homoeology between wheat chromosome 6A and the translocation chromosomes, and showed that Lr56 was located near the long arm telomere. Thirty of the 53 recombinants had breakpoints between Lr56 and the most distal marker Xgwm427. These were characterized with additional markers. The data suggested that recombinants #39, 157 and 175 were wheat chromosomes 6A with small intercalary inserts of foreign chromatin containing Lr56 and Yr38, located distally on the long arms. These three recombinants are being incorporated into adapted germplasm. Attempts to identify the single shortest translocation and to develop appropriate markers are being continued.     

Cytogenetic studies in wheat XVII. Monosomic analysis and linkage relationships of gene Yr15 for resistance to stripe rust

Summary The highly effective stripe rust resistance gene, Yr15, derived from Triticum dicoccoides, was located in chromosome 1BS. Yr15 showed linkage of 0.30 (34 cM) with Yr10 and 0.07 with the centromere. Yr15 was preferentially transmitted relative to its alternate allele.     

Combination of resistance tests and molecular tests to postulate the yellow rust resistance gene Yr17 in bread wheat lines

Yellow rust caused by Puccinia striiformis is a wheat disease of worldwide importance. The Yr17 resistance gene introgressed from Aegilops ventricosa was effective, in France, against all yellow rust isolates until 1998. The SC‐Y15 marker is one of three molecular markers closely linked to Yr17. In this paper, results obtained are compared with the molecular marker SC‐Y15 and with resistance tests performed at the seedling and adult plant stages on 31 lines from five populations derived from recurrent selection programmes. The resistance tests showed that Yr17 controlled the resistance in seven lines, but that others had additional resistance at the adult stage (18 lines). The molecular test corresponded well with the resistance test in most lines (98% of 156 plants tested), including individual plants that were resistant or susceptible in heterogeneous lines. It also indicated the presence of Yr17 in lines in which it could not be identified by the resistance test because of the presence of other genes. Three of the 156 plants tested appeared to have the gene Yr17 according to the resistance tests, but lacked the molecular marker. These could have resulted from breakage of the linkage, the number being consistent with the estimate of linkage already published. This indicated the need for a resistance test, at least in later stages of breeding programmes, if it is considered essential to have the Yr17 gene present. The use of the selected lines in breeding programmes is also discussed.     

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.     

Evaluating the contribution of Yr genes to stripe rust resistance breeding through marker-assisted detection in wheat

Numerous stripe rust resistance genes have been identified from wheat, and new virulent races of Puccinia striiformis f. sp. tritici have also emerged in recent years. Deployment of diverse combinations of resistance genes is an efficient way to combat virulent evolution of strip rust pathogen. In this study, publically available molecular markers were used to identify the distribution of 36 Yr genes in 672 wheat accessions. The effectiveness of Yr genes individually and in combinations was also evaluated in field conditions. The result showed effective resistance of some recently applied genes, such as Yr15 and Yr65. It also showed the lost efficacy of some once widely used genes, such as Yr9 and Yr10. Moreover, significant additive effects were observed in some gene combinations, such as Yr9 + Yr18 and Yr30 + Yr46. Proper deploying of Yr genes and utilizing the positive interactions will be helpful for durable resistance breeding in wheat.