Quantitative-genetic analysis of leaf-rust resistance in seedling and adult-plant stages of inbred lines and their testcrosses in winter rye

Leaf rust is the most frequent leaf disease of winter rye in Germany. All widely grown population and hybrid varieties are susceptible. This study was undertaken to estimate quantitative‐genetic parameters of leaf‐rust resistance in self‐fertile breeding materials with introgressed foreign leaf‐rust resistances and to analyze the relative importance of seedling and adult‐plant resistance. Forty‐four inbred lines and their corresponding testcrosses with a highly susceptible tester line were grown in a field in four different environments (location‐year combinations) with artificial inoculation. Plots were separated by a nonhost to promote autoinfections and minimize interplot interference. Leaf‐rust severity was rated on three leaf insertions at three dates. The testcrosses showed a considerably higher disease severity than the lines. High correlations (r ? 0.9, P = 0.01) existed among the leaf insertions and the rating dates. Large genotypic variation for resistance was found in both the inbred and testcross populations. Genotype‐environment interaction and error variances were of minor importance, thus high entry‐mean heritabilities were achieved. A tight correlation between the inbreds and their corresponding testcrosses was found (r = 0.88, P = 0.01). Heterosis for resistance was significant (P = 0.05), but not very important. In a seedling test with 20—30 single‐pustule isolates, 34 out of 44 inbreds reacted race‐specifically. From the remaining inbred lines, three were medium and seven highly susceptible. In a further greenhouse test with 16 inbreds, seven were susceptible and five were resistant in both seedling and adult‐plant stages. The remaining four lines had adult‐plant resistance. In conclusion, race‐specific leaf‐rust resistance can be selected among inbred lines per se. Lines should also be tested in the adult‐plant stage.     

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.     

The role of Lr34 in imparting durable resistance to wheat leaf rust through gene interaction

Summary The leaf rust responses of wheat lines carrying the complementary genes Lr27 and Lr31 and the same genes in a Chinese Spring background which contains Lr34, indicate that Lr34 interacts with the complementary genes to give enhanced levels of field resistance to leaf rust. Lr34, particularly in combination with other genes, is considered to be an important gene for imparting a high degree of durable resistance to leaf rust. Its similarity to Sr2, an adult plant gene for resistance to stem rust and its association with adult plant resistances to stem and stripe rusts are discussed.     

Monosomic analyses of stripe rust and leaf rust resistance genes in winter wheat varieties Lüquyu and Yantar

Summary A set of 21 monosomics of Novosadska Rana-1 was used to locate the rust resistance genes of Lüqiyu, a stripe rust resistant line developed by BAU and Yantar, a leaf rust resistant wheat introduced from Bulgaria. The resistance of the former to p. striiformis race C25 was conditioned by a dominant gene located on chromosome 2B, whereas that of the latter to P. recondita race CL3 was controlled by two complementary dominant genes located on chromosomes 5A and 1D, respectively. The relationship of the stripe rust resistance gene in Lüqiyu to Yr5, Yr7 or Yr _Suwon' all located on chromosome 2B is unknown. The two complementary leaf rust resistance factors in Yantar appear to be new.     

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.     

Genes Lr48 and Lr49 for hypersensitive adult plant leaf rust resistance in wheat (Triticum aestivum L.)

The genetic bases of leaf rust resistance in wheat (Triticum aestivum L.) line CSP44, selected from the Australian cultivar Condor, and Indian cultivar VL404, were studied. The reaction patterns of CSP44 and VL404 against Indian races 12, 77, 77-1, 77-2, 77-3, 77-4, 77-5 and 108 were different from reaction patterns shown by near-isogenic lines with known adult plant resistance (APR) genes, viz. Lr12, Lr13, Lr22b and Lr34. Although the reaction patterns of CSP44 and VL404 were similar to the near-isogenic line Tc+Lr22a, tests of allelism indicated absence of Lr22a in both CSP44 and VL404. On the basis of genetic studies, their resistances in field tests against race 77-5, the most virulent race from the Indian sub-continent, were each ascribed to two genes. One of the two genes in each wheat was identified to be the non-hypersensitive APR gene Lr34. The second APR genes in CSP44 and VL404 gave hypersensitive reaction types and were recessive and dominant, respectively. The gene in CSP44 was designated Lr48and the gene in VL404, Lr49. This revised version was published online in August 2006 with corrections to the Cover Date.     

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。     

Quantitative trait loci for temperature-sensitive resistance to <Emphasis Type="Italic">Puccinia striiformis</Emphasis> f. sp. <Emphasis Type="Italic">tritici</Emphasis> in wheat cultivar Flinor

Stripe (yellow) rust, caused by Puccinia striiformis Westend. f. sp. tritici Eriks. (Pst), is an important disease of wheat (Triticum aestivum L.) globally. Use of host resistance is an important strategy to manage the disease. The cultivar Flinor has temperature-sensitive resistance to stripe rust. To map quantitative trait loci (QTLs) for these temperature-sensitive resistances, Flinor was crossed with susceptible cultivar Ming Xian 169. The seedlings of the parents, and F₁, F₃ progeny were screened against Chinese yellow rust race CYR32 in controlled-temperature growth chambers under different temperature regimes. Genetic analysis confirmed two genes for temperature-sensitive stripe rust resistance. A linkage map of SSR markers was constructed using 130 F₃ families derived from the cross. Two temperature-sensitive resistance QTLs were detected on chromosome 5B, designated QYr-tem-5B.1 and QYr-tem-5B.2, respectively, and are separated by a genetic distance of over 50 cM. The loci contributed 33.12 and 37.33% of the total phenotypic variation for infection type, respectively, and up to 70.45% collectively. Favorable alleles of these two QTLs came from Flinor. These two QTLs are temperature-sensitive resistance loci and different from previously reported QTLs for resistance to stripe rust.     

Alleles on the two dwarfing loci on 4B and 4D are main drivers of FHB‐related traits in the Canadian winter wheat population “Vienna” × “25R47”

Fusarium head blight (FHB), leaf rust and stem rust are among the most destructive wheat diseases. High‐yielding, native disease resistance sources are available in North America. The objective of this study was to map loci associated with FHB traits, leaf rust, stem rust and plant height in a “Vienna”/”25R47” population. DArT markers were used to generate a genetic map, and quantitative trait loci (QTL) analysis was performed by evaluating 113 doubled haploid lines across three environments in Ontario, Canada. FHB resistance QTL were identified on chromosomes 4D, 4B, 2D and 7A, while a QTL for leaf and stem rust resistance was identified on chromosome 1B. The dwarfing alleles of both Rht‐B1 and Rht‐D1 were associated with increased FHB index and DON content.     

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.     

Detection and inheritance of leaf rust resistance in common wheat lines Agra Local and IWP94

Genetic studies were undertaken to determine the number and identities of leaf rust resistance genes in common wheat lines Agra Local and IWP94. The infection type arrays of the two lines with eight pathotypes (pt.) of P. triticina were different from those of lines possessing known leaf rust resistance (Lr) genes. Agra Local possessed two recessive resistance genes, one conditioning resistance to pathotype 4R9-7, and the other, a temperature-sensitive factor, gave resistance to pt. 121R127 at high temperature (27°C). IWP94 was previously demonstrated to carry Lr23. From the present study IWP94 was determined to have at least four leaf rust resistance genes. The first of these was the same recessive gene conferring resistance to pathotype 4R9-7 which was found in Agra Local. A second partially dominant gene conferred resistance to pathotype 121R127 at high temperature and two additional recessive genes governed resistance to pathotype 93R15. When present together, these two recessive genes complemented each other and provided resistance to pathotype 69R13 as well. One of the two recessive genes conferring resistance to pathotypes 93R15 and 69R13 was Lr23.     

Sources of resistance to Indian pathotypes of Puccinia graminis tritici and P. triticina in durum wheat

Stem rust and leaf rust are important diseases affecting durum wheat production in India. Inheritance and extent of diversity in resistance were studied in five durum genotypes, viz. ?B 662', ?ED 2398‐A', ?HG 110', ?IWP 5019' and ?Line 1172? using Pgt pathotypes 40A and 117‐6, and Pt pathotypes 12‐2 and 104‐2. F₂ and F₃ analyses showed that resistance was conferred by one or two genes in each line. In all, four genes for resistance to 40A, and eight each for pathotypes 117‐6 and 12‐2 were identified among the five genotypes, and three for resistance to 104‐2 among B 662, ED 2398‐A and IWP 5019 were indicated by tests of allelism. Although the gene identities are not known, at least some of them should be different from Sr2, Sr7b, Sr8a, Sr8b, Sr9e, Sr9g, Sr11, Sr12, Sr13, Sr14, Sr17, Sr23 and Sr28; and from Lr3, Lr14a, Lr23 and Lr27 + Lr31. These genotypes enrich the diversity of resistance to stem rust and leaf rust for durum wheat improvement.     

Association of leaf rust and powdery mildew resistance in a recombinant derived from a Hordeum vulgare × Hordeum bulbosum hybrid

While studying powdery mildew resistance in a recombinant line (code 81882) derived from a Hordeum vulgare (cv. ‘Vada’) ×Hordeum bulbosum hybrid, a low infection type of resistance to leaf rust was observed. To determine the mode of inheritance of the leaf rust resistance and whether there was linkage between the two resistances, F₂ and F₃ progenies from crosses between 81882 and ‘Vada’ were inoculated with the leaf rust and powdery mildew pathogens. Southern blots were prepared using restricted DNA extracted from leaves of 82 F₂ plants and four chromosome 2HS sequences were hybridized with the blots to define the length of the introgression. The leaf rust resistance appears to be inherited as a single dominant gene on chromosome 2HS, which co-segregates with the powdery mildew resistance. There was an almost complete association between the resistances and the respective molecular markers, but it is likely that the strong linkage results from the frequent inheritance of the introgressed H. bulbosum DNA as an intact segment of chromatin with only low levels of recombination within the segment.     

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.     

A test of supposed mutants for stem rust resistance in wheat

Summary Eight stem rust (Puccinia graminis tritici Eriks. and Henn.) resistant lines (designated TICENA lines) that had been selected by Veiga et al. (1981) following gamma radiation of BH-1146 wheat (Triticum aestivum L.) were studied. Six of the lines were resistant to race 15B-1 of stem rust and susceptible to race 56, and proved to carry the gene Sr7a. TICENA 4 carries two unidentified genes, each giving resistance to one of the two races. TICENA 10 carries Sr6, Sr7a and an unidentified gene giving resistance to race 56 but not 15B-1. The results raise doubts about the supposed origin of the lines as mutants.     

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.     

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.     

Identification and mapping of adult‐plant stripe rust resistance in soft red winter wheat cultivar ‘USG 3555’

Little is known about the extent or diversity of resistance in soft red winter wheat (Triticum aestivum L.) to stripe rust, caused by the fungal pathogen Puccinia striiformis f.sp. tritici. The soft red winter (SRW) wheat cultivar ‘USG 3555’ has effective adult‐plant resistance to stripe rust, which was characterized in a population derived from ‘USG 3555’/‘Neuse’. The mapping population consisted of 99 recombinant inbred lines, which were evaluated for stripe rust infection type (IT) and severity to race PST‐100 in field trials in North Carolina in 2010 and 2011. Genome‐wide molecular‐marker screenings with 119 simple sequence repeats and 560 Diversity Arrays Technology (DArT) markers were employed to identify quantitative trait loci (QTL) for stripe rust resistance. QTL on chromosomes 1AS, 4BL and 7D of ‘USG 3555’ explained 12.8, 73.0 and 13.6% of the variation in stripe rust IT, and 13.5, 72.3 and 10.5% of the variation in stripe rust severity, respectively. Use of these and additional diagnostic markers for these QTL will facilitate the introgression of this source of stripe rust resistance into SRW wheat lines via marker‐assisted selection.     

Possible New Genes for Resistance to Powdery Mildew, Septoria Glume Blotch and Leaf Rust of Wheat

An Ethiopian wheat collection consisting of 293 tetraploid and hexaploid entries was investigated for resistance to powdery mildew, Septoria glume blotch, and leaf rust with the aim of finding probable new genes for resistance to these diseases. Seedlings were screened with isolates of these diseases in the greenhouse or growth chamber. The material was also scored for field resistance to powdery mildew after the fifth leaf stale. The diversity of the reaction types to powdery mildew and Septoria glume blotch was estimated by the Shannon-Weaver diversity index. Thirty-nine entries (13%) of the collection were resistant to moderately resistant co the mildew isolates, 14S-77 and 46—77, that had: a combined virulence spectrum effective against nine identified genes for resistance to powdery mildew. One hundred and et till TV-tour entries (63 %) of the collection showed field resistance to mildew. One hundred and eighty-one entries (62 %) of the collection were at least moderately resistant in an aggressive isolate of Sartorial nodorum. Resistance to a race of leaf rust was detected in one hundred and sixty-eight entries or 58.% of the collection. Generally, resistance to these diseases is concentrated in Central and Southern Ethiopia. The different reaction types of the resistant entries to these diseases and the high estimates of diversity for reaction types indicated the presence of many different probable new genes and genetic backgrounds for resistance to these diseases.     

Variation of characters in near-isogenic lines of wheat with added genes for leaf rust resistance

Summary Using the cultivar Arina as the recurrent parent, six backcrosses were made with two donor lines carrying the leaf rust resistance genes Lr1 and Lr9, respectively. Selection for leaf rust resistance occurred at the seedling stage in the greenhouse; the first plants transferred to the field were BC₆F₄s. Frequency distribution of the 332 Lr1/7 × Arina and the 335 Lr9/7 × Arina lines showed continuous variation for yellow rust resistance and heading date in these leaf rust near-isogenic lines (NILs). Similar results were also obtained for plant height, for resistance to powdery mildew and glume blotch, as well as for baking quality characters in another set of more advanced NILs. The available information on the behaviour of one of the parents of cultivar Arina led to the conclusion that the expressed yellow rust resistance is quantitative and might possibly be durable.