Effects of inoculum density and temperature on three components of leaf rust resistance controlled by Lr34 in wheat

Summary Different inoculum densities had negligible effects on latent period, uredinium density and uredinium size measured on flag leaves of adult RL6058 (Thatcher^*6/PI58548[Lr34]) plants kept at low (17.5°C) post-infection temperatures in a glasshouse. In a qualitative assessment of rust severity at higher (24.6°C) temperatures, all three components of resistance indicated a susceptible flag leaf response on RL6058. In the latter environment, precise estimations of receptivity to different inoculum densities showed that adult RL6058 plants supported significantly less pustules than the leaf rust-susceptible cultivar Thatcher. In tillering plants, statistically equal numbers of uredinia developed on RL6058 and Thatcher in all paired temperature-inoculum density combinations. Growth stage-related susceptibility, and higher temperatures conducive to a shorter latent period and larger uredinia, could result in high terminal severities of Puccinia recondita f. sp. tritici on wheat genotypes containing Lr34. The reduction in receptivity associated with this gene may contribute, however, to delayed disease increase on cultivars or lines with monogenic Lr34 resistance.     

Genetics of stem rust resistance in the spring wheat cultivar Thatcher and the enhancement of stem rust resistance by <Emphasis Type="Italic">Lr34</Emphasis>

Three recombinant inbred line populations from the crosses RL6071/Thatcher, RL6071/RL6058 (Thatcher Lr34), and Thatcher/RL6058, were used to study the genetics of stem rust resistance in Thatcher and TcLr34. Segregation of stem rust response in each population was used to determine the number of genes conferring resistance, as well as the effect of the leaf rust resistance gene Lr34 on stem rust resistance. The relationship between resistance in seedling and adult plants was also examined, and an attempt was made to identify microsatellite markers linked to genes that were effective in adult plants. In field plot tests at least three additive resistance genes segregated in the RL6071/RL6058 population, whereas two resistance genes segregated in the RL6071/Thatcher population. The presence of the gene Lr34 permitted the expression of additional stem rust resistance in Thatcher-derived lines both at the seedling and adult plant stages. Seedling resistance to races TPMK and RKQQ was significantly associated with resistance in adult plants, whereas seedling resistance to races QCCD and QCCB may have made a minor contribution. The seedling resistance genes Sr16 and Sr12 may have contributed to resistance in adult plants. A molecular marker linked to resistance in adult plants was identified on chromosome 2BL.     

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.     

The influence of leaf rust resistance genes Lr29, Lr34, Lr35 and Lr37 on breadmaking quality in wheat

Leaf rust, caused by Puccinia triticina, is considered one of the most important diseases of wheat. In South Africa the genes Lr29, Lr34, Lr35 and Lr37 confer effective resistance to leaf rust, qualifying them for use in cultivar improvement. To study possible secondary effects of these genes, a collection of BC6 lines containing each of the genes singly, was evaluated for breadmaking quality. The recurrent parent Karee, and Thatcher NILs used as resistance donors in the primary crosses, as well as Thatcher, were included as checks. The presence of Lr29, Lr34, Lr35 and Lr37 caused a significant increase in flour protein and water absorption. For most of the other characteristics the NILs performed statistically similar to the recurrent parent. Some sib lines performed significantly better than others, emphasising the value of selecting for improved quality among backcross lines. This revised version was published online in August 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.     

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.     

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.     

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.     

Genetic Diversity for Adult Plant Resistance to Leaf Rust (Puccinia recondita) in Near-Isogenic Lines and in Indian Wheats

Variation for adult plant resistance in near-isogenic wheat lines carrying Lrl4b, Lrl4ab and Lr30 in a ‘Thatcher’ background indicated the possible presence of novel adult plant resistance genes effective against the Indian leaf rust population. Sixty-one wheats released for cultivation in India were grown in isolated nurseries. Each nursery was separately inoculated with one of four leaf rust pathotypes which had been selected to aid identification of resistance effective only in the adult plant stage. Seven distinct response groups were recognised and a minimum of six sources of adult plant resistance were postulated. In a group of 14 wheats, resistance was explained on the basis of the seedling response genes that were identified. Similar results for two years with pathotype 77-1 gave support to the reliability of field tests. Adult plant resistance (APR) sources were either race-specific or effective against all pathotypes used. Seedlings of cultivars with APR showed susceptible reactions. The possible presence of Lr34 in Indian wheats and its role in durable leaf rust resistance are discussed.     

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.     

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.     

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.     

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.     

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.     

Development and Validation of SCAR Markers Co-Segregating with an Agropyron Elongatum Derived Leaf Rust Resistance Gene Lr24 in Wheat

Summary An Agropyron elongatum-derived leaf rust resistance gene Lr24 located on chromosome 3DL of wheat was tagged with six random amplified polymorphic DNA (RAPD) markers which co-segregated with the gene. The markers were identified in homozygous resistant F₂ plants taken from a population segregating for leaf rust resistance generated from a cross between two near-isogenic lines (NILs) differing only for Lr24. Phenotyping was done by inoculating the plants with pathotype 77-5 of Puccinia triticina. To enable gene-specific selection, three RAPD markers (S1302₆₀₉, S1326₆₁₅ and OPAB-1₃₈₈) were successfully converted to polymorphic sequence characterized amplified region (SCAR) markers, amplifying only the critical DNA fragments co-segregating with Lr24. The SCAR markers were validated for specificity to the gene Lr24 in wheat NILs possessing Lr24 in 10 additional genetic backgrounds including the Thatcher NIL, but not to 43 Thatcher NILs possessing designated leaf rust resistance genes other than Lr24. This indicated the potential usefulness of these SCAR markers in marker assisted selection (MAS) and for pyramiding leaf rust resistance genes in wheat.     

56个小麦品种（系）的苗期和成株抗叶锈鉴定

为了研究中国小麦品种中所携带的抗叶锈基因,对56个小麦品种（系）进行苗期接种推导其中所含有的抗叶锈基因,同时连续2年对供试材料进行田间成株抗叶锈鉴定。通过苗期基因推导结合分子标记辅助检测,结果表明,在36个小麦品种中共鉴定出Lr26、Lr34、Lr1、Lr2a、Lr11、Lr20、Lr30、Lr33和Lr44等9个抗叶锈基因,其中28个品种含有Lr26,Lr1和Lr20分别存在于6个品种中,4个品种含有Lr30,Lr11和Lr44各存在于2个品种中,Lr2a、Lr33和Lr34各自在1个品种中出现。经过2年的田间抗叶锈鉴定共筛选出46个慢锈品种。筛选到的这些苗期和成株抗病品种均可用于小麦持久抗叶锈品种的培育。     

‘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.     

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.     

Leaf rust resistance in selected late maturity, common wheat cultivars from Uruguay

Leaf rust (caused by Puccinia triticina) is one of the most important diseases of wheat in Uruguay, and breeding for resistance to this disease is a priority for the INIA wheat program. Knowledge of the effective resistance genes present in the germplasm is relevant when selecting for effective and more durable resistance. The leaf rust resistance present in six adapted wheat cultivars that are parents of many advanced lines was studied. Races of P. triticina with different virulence combinations were used to determine which seedling resistance genes might be present in the six cultivars and/or derived lines. Genetic analysis of seedling and adult plant resistance (APR) was conducted on BC₁F₂ and F₃ generations from crosses of four cultivars with the susceptible cultivar Thatcher. The presence of APR genes Lr13 and Lr34 was confirmed with crosses of the four cultivars and Thatcher lines with these genes. A genetic marker associated with Lr34 was used to postulate the presence of this gene in all cultivars. The cultivars and resistance genes postulated to be present were: Estanzuela Calandria Lr3bg, Lr16 and Lr24; Estanzuela Federal Lr10; Estanzuela Halcón Lr10, Lr14a, and Lr16; INIA Tijereta and INIA Garza Lr16, Lr24 and Lr34; and INIA Torcaza Lr10 and Lr24. Only Lr16 and Lr34 remain effective to the predominant pathotypes. Additional ineffective seedling resistance that could not be identified was present in E. Federal, I. Tijereta and I. Torcaza. Unknown APR gene(s) could be present in E. Calandria and E. Federal.     

28个小麦微核心种质抗叶锈性分析

选取在成株期表现高、中、低抗叶锈的28个小麦微核心种质,利用39个以Thatcher为背景的近等基因系(或单基因系)作为已知基因的鉴别寄主,接种8个小麦叶锈菌致病型进行苗期抗叶锈基因推导,结合成株期抗病鉴定,初步明确了这些品种(系)的抗性和可能携带的抗病基因。利用19个与Lr基因紧密连锁或共分离的分子标记,对28个微核心种质进行抗叶锈病基因的进一步鉴定,推测新克旱9号可能含有Lr17、Lr2b、Lr14a和Lr33;兴义4号可能含有Lr26、Lr36和Lr37;紫皮可能含有Lr2b和Lr34;大白皮含有Lr1;毕红穗含有Lr1、Lr10和Lr34;中优9507含有Lr10;小白麦、红粒当年老、老麦、蝉不吱、苏麦3号和车锏子含有Lr1和Lr34;红花早可能含有Lr1、Lr34、Lr14a和Lr2b;江西早、泡子麦、三月黄、有芒扫谷旦、阜阳红、成都光头和酱麦可能含有Lr34;敦化春麦和甘肃96可能含有Lr28;欧柔可能含有Lr34、Lr16、Lr11、Lr3bg和Lr33;此外,新克旱9号、兴义4号、红花早、红粒当年老、欧柔、有芒扫谷旦、成都光头、甘肃96、小红皮、定兴寨、中优9507和红冬麦中可能含有未知抗病基因;在这28份种质中,不含Lr9、Lr19、Lr20、Lr21、Lr24、Lr29、Lr35、Lr38和Lr47基因。研究结果表明,测试的微核心种质中含有比较丰富的抗叶锈病基因,可为育种提供丰富的抗源。