Leaf Rust and Stripe Rust Resistance Genes Derived from Aegilops Sharonensis

Summary Linked leaf and stripe rust resistance genes introgressed into hexaploid wheat from Aegilops sharonensis provided protection in the seedling stage to a wide range of pathotypes of the two diseases. Monosomic and telosomic analyses showed that the resistance genes occur on wheat chromosome 6A. This result could be confirmed making use of mapped chromosome 6A microsatellite markers. The introgressed chromatin appeared to involve the proximal part of 6AL and the complete 6AS arm and it was thus not possible to deduce the chromosome arm harbouring the resistance genes. The resistance showed non-Mendelian transmission. The genetic background of a heterozygote interacted with the introgressed region to result in either preferential or impaired female transmission. Male transmission appeared to be affected in a different way from female transmission and was exclusive in the genetic background studied. Symbols Lr56 and Yr38 are proposed to designate the respective genes of which line 0352-4 is the appropriate source material.     

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.     

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.     

Mapping of the leaf rust resistance gene <Emphasis Type="Italic">Lr38</Emphasis> on wheat chromosome arm 6DL using SSR markers

Leaf rust caused by the fungus Puccinia triticina is one of the most important diseases of wheat (Triticum aestivum) worldwide. The use of resistant wheat cultivars is considered the most economical and environment-friendly approach in controlling the disease. The Lr38 gene, introgressed from Agropyron intermedium, confers a stable seedling and adult plant resistance against multiple isolates tested in Europe. In the present study, 94 F₂ plants resulting from a cross made between the resistant Thatcher-derived near-isogenic line (NIL) RL6097, and the susceptible Ethiopian wheat cultivar Kubsa were used to map the Thatcher Lr38 locus in wheat using simple sequence repeat (SSR) markers. Out of 54 markers tested, 15 SSRs were polymorphic between the two parents and subsequently genotyped in the population. The P. triticina isolate DZ7-24 (race FGJTJ), discriminating Lr38 resistant and susceptible plants, was used to inoculate seedlings of the two parents and the segregating population. The SSR markers Xwmc773 and Xbarc273 flanked the Lr38 locus at a distance of 6.1 and 7.9 cM, respectively, to the proximal end of wheat chromosome arm 6DL. The SSR markers Xcfd5 and Xcfd60 both flanked the locus at a distance of 22.1 cM to the distal end of 6DL. In future, these SSR markers can be used by wheat breeders and pathologists for marker assisted selection (MAS) of Lr38-mediated leaf rust resistance in wheat.     

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.     

Associations and genetics of three components of slow rusting in leaf rust of wheat

Summary Forty F₆ lines, the two parental lines, and a susceptible check cultivar of wheat (Triticum aestivum L.) were inoculated in the young flag leaf stage with leaf rust (Puccinia recondita f.sp. tritici) and evaluated for latent period, receptivity, and uredinium size in a greenhouse experiment. Genotypic (r_g) and phenotypic (r_p) correlations between latent period and uredinium size were –0.81 and –0.62, respectively. A negative correlation (r_g=–0.50, r_p=–0.41) was found between latent period and receptivity and a positive correlation (r_g=0.28, r_p=0.26) between uredinium size and receptivity was found. Area under the disease progress curve (AUDPC) and final rust severity (FRS) obtained from a subsequent field study with common entries were negatively correlated with latent period and positively correlated with uredinium size. Correlations of receptivity with both AUDPC and FRS were not significant. The distributions of F₆ family mean uredinia size and latent period were continuous between slow rusting and fast rusting parents: however, the distribution for receptivity was discrete. Narrow-sense heritability estimates were 63%, 57%, and 47% for uredinium size, latent period, and receptivity, respectively. Estimates of the minimum number of effective factors were three for latent period and three or four for the uredinium size and receptivity. The components are controlled by closely linked genes or due to pleotropic effects of the same gene.Abbeviations AUDPC - Area under the disease progress curve - FRS - Final rust severity     

Analysis of durable resistance to stem rust in barley

Summary Since the mid-1940's, barley cultivars grown in the northern Great Plains of the USA and Canada have been resistant to stem rust caused byPuccinia graminis f. sp.tritici. This durable resistance is largely conferred by a single gene,Rpg1, derived from a single plant selection of the cultivar Wisconsin 37 and an unimproved Swiss cultivar. At the seedling stage, barley genotypes withRpg1 generally exhibit low mesothetic reactions at 16–20° C and slightly higher mesothetic reactions at 24–28° C to many stem rust pathotypes. This resistance is manifested by a low level of rust infection and mostly incompatible type uredia on adult plants.Rpg1 reacts in a pathotype-specific manner since some genotypes ofP. g. f. sp.tritici are virulent on cultivars carrying this gene in the field. Several factors may have contributed to the longevity of stem rust resistance in barley, a) since barley is planted early and matures early, it can sometimes escape damage from stem rust inoculum carried from the south; b) one or more minor genes may augment the level of resistance already provided byRpg1; c) the cultivation of resistant wheat cultivars and eradication of barberry have reduced the effective population size and number of potential new pathotypes ofP. g. f. sp.tritici, respectively; and d) virulent pathotypes ofP. g. f. sp.tritici andP. g. f. sp.secalis have not become established. This situation changed in 1989 when a virulent pathotype (Pgt-QCC) ofP. g. f. sp.tritici became widely distributed over the Great Plains. However,Rpg1 may still confer some degree of resistance to pathotype QCC because stem rust severities have been low to moderate and yield losses light on barley cultivars carrying the gene during the last four seasons (1989–1992). Several sources of incomplete resistance to pathotype QCC have been identified in barley. To facilitate the transfer of resistance genes from these sources into advanced breeding lines, molecular marker assisted selection is being employed.     

The inheritance of stem rust and leaf rust resistance in some Ethiopian wheat collections

Summary Common and durum wheat populations obtained from Sweden and originally collected in Ethiopia were screened for resistance to steum rust and leaf rust. Resistant selections of common wheat were crossed and backcrossed with either stem rust susceptible RL6071, or leaf rust susceptible Thatcher. Genetic studies, based largely on tests of backcross F₂ families, showed that four of the selections had in common a recessive gene SrA. Plants with this gene were resistant (1⁺ infection type) to all stem rust races tested. This gene was neither Sr26 nor Sr29. The resistance of other selections, based on tests with an array of rust isolates, was due to various combinations of Sr6, 8a, 9a, 9d, 9c, 11, 13, 30, and 36. One of the selections had linked genes, Lr19/Sr25. Another selection had a dominant gene for resistance (;1 infection type) to all the races of leaf rust. With the possible exception of this gene for leaf rust resistance and SrA, no obviously new resistance was found.     

DNA fingerprint of F1 interspecific hybrids from the Triticeae tribe using ISSRs

Inter-simple-sequence repeat (ISSR) were used to fingerprint three kinds of F1 interspecific hybrids: the Portuguese durum wheat cultivar ‘Candial’ × tritordeum, the Portuguese bread wheat cultivar ‘Barbela’ × tritordeum and the Portuguese triticale cultivar ‘Douro’ × tritordeum. Among the 30 primers tested, the number of discriminative primers varies from 13 to 20 in each hybrid combination, the total number of ISSR loci analysed ranged from 101 to 183, and the number of discriminative ISSR loci varied from 50 to 71. ISSR-PCR technique revealed to be a valuable fingerprint system for all F1 hybrid combinations and respective parents analysed here. The complementary use of molecular cytogenetic techniques and molecular markers could allow a more accurately evaluation and characterisation of different plant species.     

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.     

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.     

Characterisation and origin of rust and powdery mildew resistance genes in VPM1 wheat

Summary The expression of rust resistances conferred by closely linked genes derived from VPM1 varied with environmental conditions and with genetic backgrounds. Under low light and low temperature conditions seedlings carrying Yr17 showed susceptible responses. Stem rust and leaf rust resistance genes Sr38 and Lr37 tended to confer more resistance at 17±2° C than at normal temperatures above > 20° C. These studies supported the hypothesis that Yr17, Lr37 and Sr38 were derived from Aegilops ventricosa, whereas Pm4b was probably derived from T. persicum. Studies on certain addition lines and parental stocks indicated that wheat cytoplasm may enhance the expression of Sr38.     

More precise map position and origin of a durable non-specific adult plant disease resistance against stripe rust (Puccinia striiformis) in wheat

Recently a major gene determining non-specific adult plant disease resistance against stripe rust (Puccinia striiformis) designated Yrns-B1 was mapped in wheat Triticum aestivum L. by using a cross between ‘Lgst. 79-74’ (resistant) and ‘Winzi’ (susceptible). Linkage to five Gatersleben wheat microsatellite (GWM) markers was discovered, previously mapped on chromosome arm 3BS. In the present study this map was improved by the incorporation of four additional GWM markers. QTL-analysis revealed high LOD values for the resistance at all nine loci, whereas the largest LOD (20.76) was found for the newly mapped marker Xgwm1329. Microsatellite analysis and resistance tests of a collection of old German/UK wheat varieties, including probable ancestors of ‘Lgst.79-74’ were carried out. A high coincidence of non-specific adult plant disease resistance against stripe rust and the presence of ‘Lgst. 79-74’ allele (117 bp) of the marker Xgwm533 was observed among the varieties tested. Linkage during the inheritance of both the resistance and the 117 bp allele of Xgwm533 was demonstrated. The probable origin of Yrns-B1 is discussed. Carriers of this resistance gene were grown on large areas since more than 100 years. To estimate the capability of Xgwm533 as a diagnostic marker for non-specific adult plant disease resistance against stripe rust, microsatellite analysis and resistance tests of a collection of Russian spring wheat varieties were performed. The 117 bp allele of Xgwm533 was found in about 35% of the Russian cultivars analysed, however, none of them possessed the expected disease resistance. Thus, the utilisation of Xgwm533 as diagnostic marker seems to be restricted to certain genepools.     

Molecular characterization of a novel powdery mildew resistance gene Pm30 in wheat originating from wild emmer

Powdery mildew caused by Erysiphe graminis f. sp. tritici is one of the most important wheat diseases in many regions of theworld. A powdery mildew resistance gene, originating from wild emmerwheat (Triticum dicoccoides) accession `C20', from Rosh Pinna, Israel,was successfully transferred to hexaploid wheat through crossing andbackcrossing. Genetic analysis indicated that a single dominant genecontrols the powdery mildew resistance at the seedling stage. SegregatingBC₁F₂ progenies of the cross 87-1/C20//2*8866 wereused for bulked segregant analysis (BSA). The PCR approach was used togenerate polymorphic DNA fragments between the resistant and susceptibleDNA pools by use of 10-mer random primers, STS primers, and wheatmicrosatellite primers. Three markers, Xgwm159/430,Xgwm159/460, and Xgwm159/500, were found to be linked tothe resistance gene. After evaluating the polymorphic markers in twosegregating populations, the distance between the markers and the mildewresistance gene was estimated to be 5–6 cM. By means of ChineseSpring nullisomic-tetrasomics and ditelosomics, the polymorphic markersand the resistance gene were assigned to chromosome arm 5BS and werephysically mapped on the gene rich regions of fragment length (FL) 0.41–0.43 by Chinese Spring deletion lines. As no powdery mildew resistancegene has been reported on chromosome arm 5BS, the mildew resistancegene originating from C20 should be a new gene and is designated Pm30.     

The reaction of x Tritordeum and its Triticum spp. and Hordeum chilense parents to rust diseases

Summary Hexaploid and octoploid tritordeums and their parents Hordeum chilense and Triticum spp. were screened for resistance to isolates of wheat and barley yellow and brown rusts. All H. chilense lines were highly resistant to both wheat and barley brown rust, few lines were susceptible to wheat yellow rust while susceptibility to barley yellow rust was common. In general the resistance of tritordeum is predominantly contributed by the wheat parent and apparently the genes for resistance in H. chilense are inhibited in their expression by the presence of the wheat genome.Abbreviations WYR wheat yellow rust - WBR wheat brown rust - BYR barley yellow rust - BBR barley brown rust     

Alien introgression of spring habit dominant genes into bread wheat

Summary Alien dominant genes of spring habit were introgressed into bread wheat. The introgression was undertaken by simple crossing of winter bread wheat to related spring species or genera, followed by backcrossing to winter bread wheat, and did not involve the use of the ph mutants or embryo culture. The introgressed genes were located mostly on chromosomes of homoeologous group 5, and were allelic to the known Vrn genes in bread wheat. Nevertheless three groups of lines were discovered with the genes possibly located on other chromosomes. These genes were non-allelic to each other and to known Vrn genes and were designated Vrn6 ^Sc , Vrn7 ^Sc (introgressed from Secale cereale) and Vrn8 ^Ts (from Triticum sphaerococcum).     

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.     

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.     

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.     

Cytological and microsatellite mapping of the genes determining liguleless phenotype in durum wheat

Liguleless phenotypes of wheat lack ligule and auricle structures on all leaves of the plant. Two recessive genes principally control the liguleless character in tetraploid wheat. The F₂ progenies of k17769 (liguleless mutant)/Triticum dicoccoides and k17769/T. dicoccum segregated in a 15:1 ratio, whereas the F₂ progenies of k17769/T. durum and k17769/T. turgidum segregated in a 3:1 ratio. A new gene, lg₃, was found on chromosome 2A. Segregation of F₂ progenies between k17769 and chromosome substitution lines for homoeologous group 2 chromosomes suggested that the liguleless genotype had occurred by mutation at the lg₃ locus on chromosome 2A, and then by mutation at the lg₁ locus on chromosome 2B, in the process of domestication of tetraploid wheat. The gene (lg₁) was linked to Tc₂ (11.9 cM), which determines phenol colour reaction of kernels, on the long arm of chromosome 2B. The distance of lg₁ to the centromere was found to be 60.4 cM, and microsatellite mapping established the gene order, centromere – Xgwm382 – Xgwm619 – Tc₂ – lg₁ on the long arm of chromosome 2B.