Genetic basis of seedling-resistance to leaf rust in bread wheat 'Thatcher'

The bread wheat cultivar ‘Thatcher’ is documented to carry the gene Lr22b for adult‐plant resistance to leaf rust. Seedling‐resistance to leaf rust caused by Puccinia triticina in the bread wheat cultivar ‘Thatcher’, the background parent of the near‐isogenic lines for leaf rust resistance genes in wheat, is rare and no published information could be found on its genetic basis. The F₂ and F₃ analysis of the cross ‘Agra Local’ (susceptible) × ‘Thatcher’ showed that an apparently incompletely dominant gene conditioned seedling‐resistance in ‘Thatcher’ to the three ‘Thatcher’‐avirulent Indian leaf rust pathotypes – 0R8, 0R8‐1 and 0R9. Test of allelism revealed that this gene (temporarily designated LrKr1) was derived from ‘Kanred’, one of the parents of ‘Thatcher’. Absence of any susceptible F₂ segregants in a ‘Thatcher’ × ‘Marquis’ cross confirmed that an additional gene (temporarily designated LrMq1) derived from ‘Marquis’, another parent of ‘Thatcher’, was effective against pathotype 0R9 alone. These two genes as well as a second gene in ‘Kanred’ (temporarily designated LrKr2), which was effective against all the three pathotypes, but has not been inherited by ‘Thatcher’, seem to be novel, undocumented leaf rust resistance genes.     

Genetics of leaf rust resistance in three Americano landrace-derived wheat cultivars from Uruguay

A genetic analysis of the landrace‐derived wheat accessions Americano 25e, Americano 26n, and Americano 44d, from Uruguay was conducted to identify the leaf rust resistance genes present in these early wheat cultivars. The three cultivars were crossed with the leaf rust susceptible cultivar ‘Thatcher’ and approximately 80 backcross (BC1) F₂ families were derived for each cross. The BC1F₂ families and selected BC1F₄ lines were tested for seedling and adult plant leaf rust resistance with selected isolates of leaf rust, Puccinia triticina. The segregation and infection type data indicated that Americano 25e had seedling resistance genes Lr3, Lr16, an additional unidentified seedling gene, and one adult plant resistance gene that was neither Lr12 nor Lr13, and did not phenotypically resemble Lr34. Americano 26n was postulated to have genes Lr11, Lr12, Lr13, and Lr14a. Americano 44d appeared to have two possibly unique adult plant leaf rust resistance genes.     

Resistance and tolerance to leaf rust in Ethiopian tetraploid wheat landraces

Infection type to leaf rust (Puccinia recondita f. sp. tritici) in Ethiopian tetraploid wheat (Triticum turgidum L.) landraces is almost invariably susceptible and disease severity is usually high. However, such wheats produce a stable yield and seeds are less shrivelled, relative to disease severity. Using 10‘pure-line’genotypes, the effect of the disease on kernel weight and grain yield was studied to detect whether some form of incomplete resistance or tolerance was involved. Genotypes exhibited significant differences for mean disease severity. Percentage losses varied from 3.9 to 14.4% for kernel weight and from 2.8 to 31.3% for grain yield. Regression analysis revealed a significant linear relationship (r = 0.66, P < 0.05) between mean percentage loss and mean disease severity for kernel weight but not for grain yield. Only one genotype sustained significant losses of both kernel weight and grain yield, and was therefore classified as susceptible. Genotypic differences for both traits and the negative correlation (r = -0.87, P < 0.01) between them complicated interpretation of the data. Generally, however, the results indicated that incomplete resistance, possibly partial resistance, is present when one considers kernel weight. For grain yield, tolerance was also implicated since the variation could not be explained by differences in disease severity, and percentage loss did not appear to be a function of disease severity.     

Inheritance of seedling and adult plant resistance to leaf rust of selected Australian spring and English winter wheat varieties

Genetic studies were conducted to gain an understanding of the inheritance of adult plant resistance (APR) to leaf rust in six common wheat varieties. The Australian varieties ‘Cranbrook’ and ‘Harrier’ each carry two genes for APR to leaf rust. These genes are genetically independent of the seedling resistance genes Lr23 and Lrl7b, carried by the respective varieties. Adult plant resistance in ‘Suneca’ was conferred by at least two genes, in addition to the seedling genes Lr1 and Lrli. It is likely that the APRs in ‘Cranbrook’, ‘Harrier’ and ‘Suneca’ are conferred by uncharacterized gene(s). Tests of allelism confirmed that seedling resistances in the varieties ‘Avocet R’, ‘Hereward’, ‘Moulin’ and ‘Pastiche’ are conferred by Lrli. Adult plant resistance in the variety ‘Hereward’ was inherited monogenically, whereas varieties ‘Moulin’ and ‘Pastiche’ each carried two dominant genes. On the basis of rust specificity and pedigree analysis, it would seem likely that the APR genes in ‘Hereward’, ‘Moulin’ and ‘Pastiche’ are also currently uncharacterized.     

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.     

A leaf rust resistance gene, different from Lr34, associated with leaf tip necrosis in wheat

The gene Lr34 has contributed to durable resistance to leaf rust caused by Puccinia triticina in wheat worldwide. The closely associated leaf tip necrosis is generally used as the gene's marker. Lr34 has been postulated in many Indian bread wheat cultivars including ‘C 306’, based on the associated leaf tip necrosis and a few other field and glasshouse observations. The present study showed monogenic control of adult‐plant resistance in ‘C 306’ to leaf rust pathotype 77‐5 (121R63‐1). The F₂ segregation in the crosses between ‘C 306’ and the two known carriers of Lr34, ‘Line 897’ and ‘Jupateco 73’‘R’ fitted a digenic ratio. The F₃ families derived from the susceptible F₂ segregants were true breeding for susceptibility, proving the absence of Lr34 in ‘C 306’. The cross between ‘Line 897’ and ‘Jupateco 73’‘R’ did not segregate for susceptibility. Resistance in the cross ‘Agra Local’ (susceptible) × ‘C 306’ was associated with leaf tip necrosis, showing that the leaf rust resistance gene in ‘C 306’ was associated with leaf tip necrosis, but was different from Lr34. This gene is being temporarily designated as Lr‘C 306’. Hence, leaf tip necrosis cannot be considered as an exclusive marker for selecting Lr34 in wheat improvement.     

Production of haploids in bread wheat, durum wheat and hexaploid triticale crossed with pearl millet

Pearl millet is an efficient alternative to maize as a pollen source for haploid production in bread wheat. To compare haploid production frequencies in other Triticeae species, the crossabilities of two genotypes each of bread wheat, durum wheat and hexaploid triticale with four pearl millet genotypes and a maize control were examined. Embryos were obtained from crosses of all three species with both pearl millet and maize. However, significant differences in crossability were found among the three species (10.5–79.8% seed development and 1.4–15.8% embryo formation), as well as among genotypes of durum wheat (7.2–23.7% and 2.1–6.4%) and hexaploid triticale (0.3–20.6% and 0.1–2.7%). Crossability of bread wheat with pearl millet was relatively high. Haploid plants were regenerated from crosses of all three species with pearl millet. As in the case of maize crosses, low crossabilities of durum wheat and hexaploid triticale with pearl millet can be attributed to the absence of D-genome chromosomes.     

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.     

Components of quantitative resistance to yellow rust in ten spring bread wheat cultivars and their relations with field assessments

Latency period, infection frequency, lesion length, lesion growth, disease severity and percentage of infected leaf parts were assessed on 10-day-old seedling leaves and flag leaves of ten bread wheat (Triticum aestivum L.) cultivars after inoculation with urediospores of Puccinia striiformis Westend. f. sp. tritici. For all components significant genotypic differences were detected. Components of resistance tended to be associated. A long latency period was associated with a low infection frequency, small lesions, a low disease severity and a low percentage of infected leaf parts. The latency period, measured as time period until first pustule appearance (LP1), was highly correlated with the latency period measured as time period until 50% of the pustules appeared (LP50). Assessment of latency period of large numbers of cultivars could therefore be reliably done by measuring LP1 which is less time consuming than measuring LP50. Latency period, infection frequency and disease severity were highly correlated with disease development data from field experiments. These results suggest that selection in the greenhouse for one of these components should result in cultivars with high levels of quantitative resistance. Disease severity after uniform inoculation in the greenhouse can be used for monocyclic evaluations because it is the easiest to assess. This revised version was published online in July 2006 with corrections to the Cover Date.     

A microsatellite marker linked to leaf rust resistance transferred from Aegilops triuncialis into hexaploid wheat

Aegilops triuncialis (UUCC) is an excellent source of resistance to various wheat diseases, including leaf rust. Leaf rust‐resistant derivatives from a cross of a highly susceptible Triticum aestivum cv.‘WL711’ as the recurrent parent and Ae. triuncialis Ace.3549 as the donor and with and without a pair of acrocentric chromosomes were used for molecular tagging. The use of a set of sequence tagged microsatellite (STMS) markers already mapped to different wheat chromosomes unequivocally indicated that STMS marker gwm368 of chromosome 4BS was tightly linked to the Ae. triuncialis leaf rust resistance gene transferred to wheat. The presence of the Ae. Triuncialis‐specific STMS gwm368 homoeoallele along with the non‐polymorphic 4BS allele in the rust‐resistant derivatives with and without the acrocentric chromosome indicates that the resistance has been transferred from the acrocentric chromosome to either the A or the D genome of wheat. This alien leaf rust resistance gene has been temporarily named as LrTr.     

Chromosomal locations in common wheat of three new leaf rust resistance genes from Triticum monococcum

Monosomic analysis was conducted to determine chromosomal locations of three new leaf rust resistance genes recently transferred to common wheat (Triticum aestivum) from T. monococcum. The resistance gene in wheat germplasm line KS92WGRC23 was transferred from T. monococcum ssp. monococcum. The resistance genes found in KS93U3 and KS96WGRC34 were transferred from T. monococcum ssp. aegilopoides. Allelism tests showed that the three resistance genes were unlinked. The three lines were crossed with each of the seven A-genome Wichita monosomic lines. The leaf rust resistance genes in KS92WGRC23, KS93U3, and KS96WGRC34 were located on chromosomes 6A, 1A, and 5A, respectively, by monosomic analysis. These results demonstrate that the three new genes derived from T. monococcum are each different. They also differ from previously reported Lr genes. This information on chromosome location and the development of mapping populations will facilitate molecular tagging of the new genes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Partial resistance to wheat leaf rust in 18 spring wheat cultivars

Summary Eighteen spring wheat cultivars were tested in microfields and race nurseries for their partial resistance PR to wheat leaf rust under low and high disease pressure respectively. Large differences existed between the 18 cultivars, Skalavatis 56 being the most susceptible and Ponta Grossa 1 being the most resistant cultivar. Of the three epidemic parameters, disease severity (DS) at the time that the susceptible check was severely diseased and area under the transformed disease severity curve (AUTC) and the logistic growth rate (r), AUTC and DS were highly correlated. Both seemed to be reliable estimators of PR but DS should be preferred for economical reasons. The logistic growth rate seemed to be unsuitable as an estimator of partial resistance.High and low disease pressure gave similar cultivar ranking. PR can be screened and selected equally well in race nurseries with low space, low time and low cost input as in microfields with high space, time and cost input.Cultivar differences in development rate had a large impact on the cultivar differences for amount of disease and can therefore greatly bias the estimation of cultivar resistance. The resistance of early cultivars tended to be underestimated whereas the resistance of late cultivars tended to be overestimated. The effect of differences in developmental rate was most pronounced in the flag leaf. It is advisable to avoid the assessment of disease levels on the flag leaf only and to incorporate in the tests several susceptible and resistant checks that cover the range of development rates in the material to be selected, because otherwise selection for resistance will tend to select also for lateness.Regression of the epidemiological parameters on three components of partial resistance revealed that latency period (LP) is an important factor in determining the resistance observed in the field explaining on average 67% of the observed variation. Adding infection frequency (IF) and urediosorus size (US) to the linear model increased the proportion of the observed variation in the field explained by the components to 80%. This result supports the idea that the components of PR inherit independently, at least, in part.     

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.     

First report of leaf rust pathotypes virulent to highly effective Lr-genes transferred from Agropyron species to bread wheat

The gene pool of effective sources of leaf rust resistance used in the breeding of wheat (Triticum aestivum L.) includes several species of the genus Agropyron. The genes deriving therefrom (Lr 19, 19d, 29, Agⁱ1, Agⁱ2, 38) are highly effective to pathotypes of Puccinia recondita Rob. ex Desm. In the Saratov and Orenbhurg districts of Russia, however, pathotypes virulent to these genes have been discovered. These pathotypes are virulent to Saratov-bred cultivars carrying Lr 19, to ‘Indis’ (Lr 19d) and RL 6097 (Lr 38). The distribution of virulence on the ‘Thatcher’ near-isogenic lines with different Lr genes shows that most of the Lr genes tested are susceptible to these new pathotypes of P. recondita, but the Lr genes Lr 9, 23, 24, 26 were exceptions. The inoculation of Mexican bread wheat cultivars, which carry widespread Lr gene combinations, by these pathotypes disclosed different infection types. Out of 10 Lr-gene combinations, four were highly effective; namely the combinations Lr 13 + 26, Lr 26 +?, Lr 23+26 and Lr 23+26+34.     

The control of adult-plant resistance to yellow rust by the translocated chromosome 5BS-7BS of bread wheat

The reciprocal translocation 5BL-7BL and 5BS-7BS was widespread in West European wheats 30 years ago, and is probably present in many of their descendants today. In varieties with a history of durable adult-plant resistance to yellow rust and carrying this translocation, removal of the 5BS-7BS chromosome gave adult plants which were much more susceptible. It was suggested that this chromosome might therefore carry the gene(s) responsible for a major part of their resistance and possibly their durability. To test this, a series of lines was developed in which 5BS-7BS chromosomes from both resistant and susceptible varieties were substituted into a number of the durably resistant varieties. In every case, the substituted 5BS-7BS chromosome, irrespective of origin, was found to produce the resistant phenotype, indicating that background chromosomes were responsible for the differences between the varieties. The resistance and durability of the resistant varieties cannot therefore be due solely to the translocated chromosome. In similar experiments, the 5BS and 7BS arms from varieties not carrying the translocation were substituted into a variety carrying the translocation. In each instance, the lines with the substituted arms were much more susceptible than their recipient, confirming the major effect of the 5BS-7BS chromosome on resistance. The complete correlation between the translocation and resistance and between increased susceptibility and its absence suggests that the gene(s) for adult-plant resistance, located on the 5BS-7BS chromosome, may be closely linked to the break point. Alternatively, it may be a consequence of the close relatedness of some of the varieties. The presence of this gene(s) might be a factor explaining the prevalence of this translocation in some West European wheats.     

Cytogenetical studies in wheat. XVIII. Gene Yr24 for resistance to stripe rust

A new gene, Yr24, for resistance to stripe rust was transferred from a durum accession to common wheat via an amphiploid (synthetic wheat) with Aegilops tauschii. Yr24 was located in chromosome 1B by monosomic analysis. Its genetic linkage of 4 cM with Yr15 indicated its localization to the short arm.     

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.     

Novel complementary genes for adult plant leaf rust resistance in a wheat stock carrying the 1BL-1RS translocation

The wheat-rye translocation (IBL-IRS) that carries the tightly linked genes Lr26/Sr31/Yr9, has been widely exploited in the development of wheat cultivars worldwide. This resistance, however, has become ineffective owing to the evolution of new pathotypes of Puccinia recondita that neutralize the resistance of Lr26. Inheritance studies on ‘Federation4′/‘Kavkaz’ revealed complementary genes derived separately from ‘Federation’ and ‘Kavkaz’ for adult plant resistance. This previously undescribed source of resistance appears to be widely effective and could therefore be used to broaden the genetic base for resistance in India. Its effectiveness in other geographical areas is unknown.     

Development of SCAR markers for identification of stem rust resistance gene Sr31 in the homozygous or heterozygous condition in bread wheat

The stem rust resistance gene Sr31, transferred from rye (Secale cereale) into wheat (Triticum aestivum L.) imparts resistance to all the virulent pathotypes of stem rust (Puccinia graminis f. sp. tritici) found in India. Wheat genotypes including carriers and non‐carriers of the Sr31 gene were analysed using arbitrary primed polymerase chain reaction (AP‐PCR). AP‐PCR markers viz. SS30.2_580(H) associated with the Sr31 gene and SS26.1₁₁₀₀ associated with the allele for susceptibility were identified. Linkage between the markers and phenotypes was confirmed by analysing an F₂ population obtained from a cross between a resistant and a susceptible genotype. The markers were tightly linked to the respective alleles. Both the AP‐PCR markers were converted into sequence characterized amplified region (SCAR) markers, viz. SCSS30.2₅₇₆ and SCSS26.1₁₁₀₀ respectively. The markers were validated in two more segregating populations and 49 wheat genotypes. Using both markers it was possible to distinguish the homozygous from the heterozygous carriers of the Sr31 gene in the F₂ generation. The markers developed in this study can be used for pyramiding of the Sr31 gene with other rust resistance genes and in marker‐assisted selection.