Cytogenetic studies in wheat XIX. Chromosome location and linkage studies of a gene for leaf rust resistance in the Australian cultivar 'Harrier'

Monosomic analysis indicated that a seedling leaf rust resistance gene present in the Australian wheat cultivar ‘Harrier’(tentatively designated LrH) is located on chromosome 2A. LrH segregated independently of the stripe rust resistance gene Yr1 located in the long arm of that chromosome, but failed to recombine with Lr17 located in the short arm. LrH was therefore designated Lr17b and the allele formerly known as Lr17 was redesignated as Lr17a. The genes Lr17b and Lr37 showed close repulsion linkage. Tests of allelism indicated that Lr1 7b is also present in the English wheats ‘Dwarf A’(‘Hobbit Sib’), ‘Maris Fundin’ and ‘Norman’. Virulence for Lr17b occurs in Australia, and pathogenicity studies have also demonstrated virulence in many western European isolates of the leaf rust pathogen. Despite this, it is possible that the gene may be of value in some regions if used in combination with other 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.     

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.     

Use of a synthetic hexaploid Triticum miguschovae for transfer of leaf rust resistance to common wheat

Summary Triticum miguschovae, a genome addition synthetic, was used as a source for transfer of leaf rust (Puccinia recondita tritici) resistance to common wheat. This synthetic, developed from two wild species Triticum militinae and Aegilops squarrosa, proves a valuable donor of the genes for leaf rust resistance. Leaf rust resistance was transferred from T. miguschovae by both dominant and recessive genes. Stable lines phenotypically similar to their recurrent parents Kavkaz and Bezostaya 1 but differing from them in a high level of leaf rust resistance were obtained. The genes for resistance in 3 selected lines differed from each other and from the known effective genes Lr9, Lr19, and Lr24. The resistance of one of them (line 1229) is controlled by two complementary interacting genes located on chromosome 7B and 1D was revealed by monosomic analysis.     

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.     

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.     

Intergeneric Transfer (Rye to Wheat) of a Gene(s) for Russian Wheat Aphid Resistance

An octoploid triticale was derived from the F, of a Russian wheat aphid-resistant rye, ‘Turkey 77’, and ‘Chinese Spring’ wheat. The alloploid was crossed to common wheat, and to ‘Imperial’ rye/‘Chinese Spring’ disomic addition lines. F₂, progeny from these crosses were tested for Russian wheat aphid resistance and C-banded. A resistance gene(s) was found to be associated with chromosome arm IRS of the ‘Turkey 77’ rye genome. A monotelosomic IRS (‘Turkey 77’) addition plant was then crossed with the wheat cultivar ‘Gamtoos’, which has the 1BL.1RS ‘Veery’ translocation. Unlike the IRS segment in ‘Gamtoos’, the ‘Turkey 77’-derived 1 RS telosome did not express the rust resistance genes Sr31 and Ar26, which could then be used as markers. From the F, a monotelosomic 1 RS addition plant that was also heterozygous for the 1BL. 1 RS translocation was selected and testerossed with an aphid-susceptible common wheat, ‘Inia 66’ Meiotic pairing between the rye arms resulted in the recovery of five euploid Russian-wheat-aphid-resistant plants. One recombinant also retained Sr31 and Lr26 and was selfed to produce translocation homozygotes.     

The Aegilops ventricosa segment on chromosome 2AS of the wheat cultivar 'VPM1' carries the cereal cyst nematode resistance gene Cre5

Previous studies showed that the intermediate level of resistance in bread wheat line ‘VPM1’ to pathotype Ha12 of the cereal cyst nematode could be conferred by an Aegilops ventricosa‐derived gene, CreX, in chromosome arm 2AS, which also carries the rust resistance genes Yrl7, Lr37 and Sr38. Near isogenic lines (NILs) differing for the presence and absence of the Ae. ventricosa‐derived linked genes Yrl7/Lr37/Sr38 were tested with cereal cyst nematode. Lines carrying Yr17 produced significantly fewer nematode cysts than the controls. An infested soil experiment produced better differentiation among resistant and susceptible genotypes. Susceptibility of ‘Trident’ indicated that linkage between CreX and Yr17 is incomplete. Microsatellite markers did not differentiate between ‘Trident’ and CreX‐carrying genotypes. However, Xgwm636 (104) was associated with the presence of Yr17 in all six genetic backgrounds. Since none of the reported cereal cyst nematode resistance genes is located in chromosome 2AS, CreX was designated as Cre5.     

Molecular markers for the identification of resistance genes and marker-assisted selection in breeding wheat for leaf rust resistance

The resistance genes Lr9, Lr24, Lr25, Lr29, Lr35 and Lr37, which were not previously utilised in Hungary, have been incorporated into four Martonvásár winter wheat cultivars using marker-assisted selection with PCR-based markers. In the course of a backcross programme, the genes were transferred into Martonvásár wheat varieties and various BC generations were produced. Work aimed at pyramiding resistance genes is currently underway in Martonvásár, and plants containing the gene combinations Lr9 + Lr24, Lr9 + Lr25 and Lr9 + Lr29 are now available. From the BC₂F₄ generation of the ‘Mv Emma’*3/’R.L.6010’ combination (‘R.L.6010’ is the donor of the Lr9 gene) 287 lines were tested for leaf rust resistance in an artificially inoculated nursery. A co-dominant primer combination was designed to identify both resistant and susceptible offsprings. The results of resistance tests and molecular marker detection agreed in most cases. Designated leaf rust resistance genes were identified with molecular markers in wheat varieties and breeding lines. The Lr26 and Lr34 resistance genes occur frequently in the Martonvásár gene pool, and the presence of the Lr37 gene has also been detected in a number of Hungarian genotypes.     

‘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.     

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.     

Identification of a molecular marker linked to an Agropyron elongatum-derived gene Lr19 for leaf rust resistance in wheat

The leaf rust resistance gene Lr19, transferred from Agropyron elongatum into wheat (Triticum aestivum L.) imparts resistance to all pathotypes of leaf rust (Puccinia recondita f.sp. tritici) in South‐east Asia. A segregating F₂ population from a cross between the leaf rust resistant parent ‘HW 2046’ carrying Lr19 and a susceptible parent ‘Agra Local’ was screened in the phytotron against a virulent pathotype 77‐5 of leaf rust with the objective of identifying the molecular markers linked to Lr19. The gene was first tagged with a randomly amplified polymorphic DNA (RAPD) marker S73₇₂₈. The RAPD marker linked to the gene Lr19 which mapped at 6.4 ± 0.035 cM distance, was converted to a sequence characterized amplified region (SCAR) marker. The SCAR marker (SCS73₇₁₉) was specific to Lr19 and was not amplified in the near‐isogenic lines (NILs) carrying other equally effective alien genes Lr9, Lr28 and Lr32 enabling breeders to pyramid Lr19 with these genes.     

Quantitative trait loci mapping of adult‐plant resistance to leaf rust in a Fundulea 900 × ‘Thatcher’ wheat cross

Wheat leaf rust (LR), caused by the obligate biotrophic fungus Puccinia triticina (Pt), is a destructive foliar disease of common wheat (Triticum aestivum L.) worldwide. The most effective, economic means to control the disease is resistant cultivars. The Romanian wheat line Fundulea 900 showed high resistance to LR in the field. To identify the basis of resistance to LR in Fundulea 900, a population of 188 F_2:3 lines from the cross Fundulea 900/‘Thatcher’ was phenotyped for LR severity during the 2010–2011, 2011–2012 and 2012–2013 cropping seasons in the field at Baoding, Hebei Province. Bulked segregant analysis and simple sequence repeat markers were used to identify the quantitative trait loci (QTLs) for LR adult‐plant resistance in the population. Three QTLs were detected and designated as QLr.hebau‐1BL, QLr.hebau‐2DS and QLr.hebau‐7DS. Based on the chromosome positions and molecular marker tests, QLr.hebau‐1BL is Lr46, and QLr.hebau‐7DS is Lr34. QLr.hebau‐2DS was derived from ‘Thatcher’ and was close to Lr22. This result suggests that Lr22b may confer residual resistance on field nurseries when challenged with isolates virulent on Lr22b, or another gene linked to Lr22b confers this resistance from ‘Thatcher’. This study confirms the value of Lr34 and Lr46 in breeding for LR resistance in China; the contribution of the QTL to chromosome 2D needs further validation.     

Adult plant leaf rust resistance from 111 wheat (Triticum aestivum L.) cultivars

Adult plant resistance against Indian leaf rust race 77 and five of its highly virulent variants have been identified from 111 bread wheat cultivars originating from 12 countries. The adult plant resistance of only 16 of these cultivars is due to hypersensitive seedling or adult plant resistance genes. All others expressed nonhypersensitive type of resistance characteristic of the genes Lr34 and Lr46.Forty five of the 111 cultivars showed tip necrosis on flag leaves, a trait linked to the gene Lr34. Therefore, the nonhypersensilive type of resistance of these 45 cultivars is attributed to Lr34. The nonhypersensitive resistance of the remaining cultivars is likely to be due to the gene(s) different than Lr34. The reaction pattern of these 111 cultivars to six races suggests the presence of at least six to seven new hypersensitive adult plant resistance genes and at least three new hypersensitive seedling resistance genes. The known genes Lr10, Lr23 and Lr26 were detected frequently but these genes did not contribute towards the adult plant resistance of any of the 111 cultivars. Based on the presence of new genes for hypersensitive and nonhypersensitive type of resistance, the 111 cultivars have been classified into 31 diverse resistance groups. This revised version was published online in July 2006 with corrections to the Cover Date.     

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

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

Specific resistance to leaf rust expressed at the seedling stage in cultivars grown in France from 1983 to 2007

Host resistance is the most economical means to reduce yield losses caused by wheat leaf rust. Knowledge of the effective specific resistance genes is a prerequisite for analysis of the non-specific components of resistance, assumed to be more durable than specific resistance. Lr genes were inferred from seedling response phenotype of 275 wheat cultivars and 21 standard isolates of Puccinia triticina. Enough cultivars were selected for analysis so that findings would account for at least two-thirds of the French agricultural land dedicated to wheat from 1983 to 2007. In this paper, genes Lr13, Lr37, Lr10, Lr14a, Lr3, Lr26, Lr1, Lr24, Lr20 are postulated, alone or in combinations, in, respectively, 67%, 45%, 34%, 20%, 8%, 7%, 6%, 1%, and 1% of the cultivars. Forty five phenotypic arrays were found, the most frequent being (Lr10, Lr13, Lr37) and (Lr13) in 45 and 37 cultivars, respectively. Over the period, the combinations became increasingly complex. Isolates with virulence corresponding to most of the Lr gene combinations were identified in the pathogen population, except for combinations involving Lr24 and some unidentified genes. These findings will help breeders and extension service staff (Arvalis) in diversifying sources of resistance to wheat leaf rust. This information is also crucial for research programs aiming, on the one hand, to identify sources of quantitative resistance, and, on the other hand, to quantify selection pressure exerted on pathogen populations.     

Postulation of leaf (brown) rust resistance genes in 70 wheat cultivars grown in the United Kingdom

Multi-pathotype tests on 70 U.K. wheat cultivars permitted postulation of eight known seedling genes for resistance to Puccinia recondita f. sp.tritici either singly or in combinations. The most commonly detected gene was Lr13 (present in approximately 57% of cultivars), followed by Lr26 (22%), Lr37 (20%), Lr10 (17%), Lr17b (LrH) (10%), Lr1 (7%), Lr3a (6%) and Lr20(4%). This information permitted assessments of adult plant resistance (APR) in some cultivars, in field nurseries inoculated with pathotypes of P. recondita f. sp. tritici of known pathogenicities for characterized seedling resistance genes. APR was identified in eleven cultivars, including Avalon and Maris Ranger, which lacked detectable seedling resistance genes. The results provided a better understanding of specific resistances in the cultivars tested than was available from previous reports. This revised version was published online in July 2006 with corrections to the Cover Date.     

SCAR marker tagged to the alien leaf rust resistance gene Lr19 uniquely marking the Agropyron elongatum-derived gene Lr24 in wheat: a revision

A sequence characterized amplified region (SCAR) marker tagged to an Agropyron elongatum‐derived leaf rust resistance (Lr) gene Lr19 was validated on 18 known alien Lr gener in near‐isogenic lines (NILs) in the variety ‘Thatcher’, along with three wheat cultivers carrying Lr24 and two carrying Lr19. The marker was expressed only in the Lr24 lines confirming that the marker tagged the geneLr24. The monomorphic expression of the SCAR marker in 10NIL pairs for Lr19 and Lr24 revealed that each NIL pair possessed the same gene, Lr24. The donor parents used in the NIL pairs for Lr19 (‘Sunstar*6/C80‐1′) and Lr24 (‘TR380‐14*7/3Ag#14′) amplified the same fragment. Nonsegregation for leaf rust in the F₂ population of the cross between the above donor parents confirmed the presence of the same gene in the two parents. Apparently, a genuine parent stock of ‘Sunstar*6/C80‐1’ was not involved in the development of the NIL pairs for Lr19 due to an improper maintence bredding protocol either at source or destination which went undetected in the absence of signs of virulence for either gene in the region.     

Regional phenotypic diversity of Puccinia triticina and wheat host resistance in western Europe, 1995

Pathogenicity data from surveys of Puccinia triticina (formerly P. recondita f. sp. tritici) conducted in western Europe in 1995 were analysed to compare the structure of regional populations of the pathogen. Many of the populations differed in phenotypic diversity and pathotypic composition, even though they occurred within a single epidemiological unit, suggesting that local factors may influence the establishment and propagation of individual pathotypes in the regional populations. Neighbouring regions were more similar than distant regions, and all regions shared at least one pathotype, except populations in northern Italy and Scotland. A high degree of similarity was found between populations in northern France and Great Britain, providing strong evidence of free movement of inoculum between these regions. Resistance genes were postulated for a selection of 91 wheat cultivars, representing those most commonly grown in western Europe in 1995. Thirteen cultivars lacked detectable seedling resistance genes and the remaining 78 possessed from one to three resistance genes; those detected were Lr1, Lr3a, Lr10, Lr13, Lr14a, Lr17b, Lr20, Lr26 and Lr37. The most commonly detected resistance gene was Lr13, which was present singly or in combination with other resistance genes in 48 cultivars (53%). The gene Lr14a was detected in 18 cultivars, Lr26 was present in 16 cultivars. The role of host selection in the composition of the regional populations of P. triticina in western Europe in 1995 was difficult to assess on the basis of the results obtained, since virulence data were not available for Lr13 and Lr14a. This revised version was published online in August 2006 with corrections to the Cover Date.     

Resistance to leaf rust in cultivars of bread wheat and durum wheat grown in Spain

A set of bread wheat and durum wheat cultivars adapted to Spanish conditions was tested for resistance against leaf rust caused by different pathotypes of Puccinia triticina in field trials and in growth chamber studies. Lower levels of resistance were found in durum wheat than in bread wheat. The most frequent Lr genes found in bread wheat were Lr1, Lr10, Lr13, Lr20, Lr26 and Lr28. In durum wheat, additional resistance genes that differed from the known Lr genes were identified. The level of partial resistance to leaf rust was in general low, although significant levels were identified in some bread wheat and durum wheat cultivars.