Genes Governing Resistance to Puccinia hordei in Thirteen Spring Barley Accessions

ABSTRACT Leaf rust, caused by Puccinia hordei, is an important disease of barley in many parts of the world. In the eastern United States, this disease was effectively controlled for over 20 years through the deployment of cultivars carrying the resistance gene Rph7. Isolates of P. hordei with virulence for Rph7 appeared in this region in the early 1990s rendering barley cultivars with this gene vulnerable to leaf rust infection. From a preliminary evaluation test, 13 accessions from diverse geographic locations possessed resistance to P. hordei isolate VA90-34, which has virulence for genes Rph1, 2, 4, 6, 7, 8, and 11. Each of these 13 accessions was crossed with susceptible cvs. Moore or Larker to characterize gene number and gene action for resistance to P. hordei. Additionally, the 13 accessions were intercrossed and crossed to host differential lines possessing genes Rph3, Rph5, and Rph9 to determine allelic relationships of resistance genes. Seedlings of F(1), F(2), and BC(1)F(1) populations were evaluated in the greenhouse for their reaction to P. hordei isolate VA90-34. Leaf rust resistance in six of the accessions including Collo sib, CR270.3.2, Deir Alla 105, Giza 119, Gloria, and Lenka is governed by a single dominant gene located at or near the Rph3 locus. All accessions for which the gene Rph3 was postulated to govern leaf rust resistance, except for Deir Alla 105, likely possess an allele different than Rph3.c found in Estate based on the differential reaction to isolates of P. hordei. The resistance gene in Grit and Donan is located at or near the Rph9 locus. Alleles at both the Rph3 and Rph9 loci confer resistance in Femina and Dorina. In addition to Rph3, Caroline and CR366.13.2 likely possess a second unknown recessive gene for leaf rust resistance. Resistance in Carre 180 is governed by a recessive gene that is different from all other genes considered in this study. Identification of both known and unique genes conferring leaf rust resistance in the barley germplasm included in this study provides breeding programs with the knowledge and opportunity to assess currently used sources of leaf rust resistance and to incorporate new sources of resistance into their programs.     

Rph20: adult plant resistance gene to barley leaf rust can be detected at early growth stages

Rph20 is the only designated gene conferring adult plant resistance to leaf rust in barley, and to date, it has not displayed race specificity. It is not known at what particular growth stage the onset of Rph20 resistance occurs, or if the expression of the gene could be observed in controlled greenhouse assays. The present studies demonstrated that Rph20 can be identified reliably under greenhouse conditions in 5-week-old seedlings under normal temperature conditions (22?±?2 °C), although the resistance becomes effective at 4 weeks in certain genetic backgrounds. The resistance conferred by Rph20 was strongest and best expressed at cooler temperatures (18?±?2 °C). The effectiveness of Rph20 at cooler temperatures may have important implications in the epidemiology of barley leaf rust under field conditions, by preventing the carryover of inoculum through winter and early spring, leading to a delayed onset of rusting and subsequently lower, if any, yield losses. The detection of Rph20 in early growth stages could help simplifying the selection of the gene in breeding programs and in monitoring Puccinia hordei populations for pathogenicity.     

Molecular mapping of the leaf rust resistance gene rph6 in barley and its linkage relationships with rph5 and rph7

ABSTRACT The barley cv. Bolivia carries two leaf rust (Puccinia hordei) resistance genes, Rph2 and Rph6, and is the only known source of the latter gene. A resistant line (Bolivia-Rph6) carrying Rph6 only was obtained in the F(4) generation of a cross between cv. Bolivia and the susceptible cv. Bowman via progeny testing with differential isolates of the leaf rust pathogen. Genetic analyses and bulk segregant analysis using amplified fragment length polymorphism (AFLP) and restriction fragment length polymorphism (RFLP) markers localized Rph6 on the short arm of barley chromosome 3H at a distance of 4.4 centimorgans (cM) distal from RFLP marker MWG2021 and 1.2 cM proximal from RFLP marker BCD907. The allelic relationship of Rph6 to other leaf rust resistance genes mapping to this region of chromosome 3H (namely Rph5 and Rph7) were tested using crosses among cvs. Magnif 102 (carrying Rph5), Bolivia-Rph6 (Rph6), and Cebada Capa (Rph7). Segregation analyses indicated that Rph6 is allelic to Rph5 and closely linked to Rph7. The data generated from this study will facilitate breeding for leaf rust resistance via marker-assisted selection and provide a starting point for positional gene cloning.     

Susceptibility of German spring barley cultivars to loose smut populations from different European origins

Forty-two registered spring barley cultivars from the German official list were tested under natural infection conditions for susceptibility to loose smut (Ustilago tritici f.sp. hordei) during two test cycles at two locations. Only cv. Steffi was found to be resistant to the local loose smut population. Cultivar Sigrid showed lowest susceptibility because of flowering inside the leaf sheath. Less than 1% infection at all sites showed up in cvs Auriga, Jacinta and Hendrix. Twenty-one cultivars had an infection rate of less than 2%. Cultivar Danuta displayed the highest susceptibility with an average of 12.6%. Another 23 spring barley accessions with expected loose smut resistance were inoculated artificially with loose smut populations obtained from 11 locations in Germany and neighbouring countries. Only Jet with the resistance Un3/6, CDC Freedom with Un8, CIho9973 with quantitative resistance, as well as Lino and GangTuoQuingKeHao1 remained disease-free. In addition to these, another eight accessions in this test group are recommended to become part of a differential tester set to distinguish origins of loose smut. Statistical analysis showed that for scoring of cultivars more importance has to be given to the number of locations for infestation than to the number of test locations to determine the degree of attack. In view of the existing inspection limits for production of certified seed in European countries, the currently registered German barley cultivars put organic seed producers and breeders at high risk in respect to loose smut infection, if the number of generations for multiplication under organic farming increases.     

Appresoorium formation of Puccinia hordei on partially resistant barley and two non-host species

One of the components of partial resistance of barley to leaf rust,Puccinia hordei, is a reduced infectibility. It was investigated whether this low infectibility may rest on a hampered appressorium formation of the leaf rust fungus. The appressorium formation on the primary leaves of 11 barley genotypes with an intermediate-to-low infectibility was compared with that on the highly infectible L94. The number of stomata per cm² leaf area occupied by appressoria ofP. hordei was determined per genotype by means of fluorescence microscopy. No cosistent differences could be detected, indicating that the mechanisms causing a low infectibility of partially resistant barley seedlings act at a phase later than the formation of the appressoria. On the non-host wheat not fewer appressoria were formed than on L94, but no appressoria were found on a lettuce genotype. The latter probably lacks the stimuli that enable the fungus to find stomata.Samenvatting Eén van de componenten van partiële resistentie van gerst tegen dwergroest,Puccinia hordei, is een verminderde infectiedichtheid. Het mechanisme, dat hieraan ten grondslag ligt, is onbekend. Een experiment werd uitgevoerd om na te gaan of bij partieel resistente rassen een verminderde appressoriumvorming optreedt. Na inoculatie in een inoculatietoren en een zorgvuldig uitgevoerde incubatie werd het aantal huidmondjes per cm² bladoppervlak bepaald dat bezet was door appressoria vanP. hordei. De elf weinig vatbare gerstlijnen uit deze studie bleken niet reproduceerbaar te verschillen van de zeer vatbare gerstlijn L94 in de mate van appressoriumbezetting. Dit wijst erop dat infectiedichtheidsverschillen t.g.v. partiële resistentie veroorzaakt worden door mechanismen die werken na de appressoriumvorming. In een tweede experiment werd aangetoond dat zelfs op de niet-waardsoort tarwe, waaropP. hordei geen symptomen veroorzaakt, niet minder appressoria worden gevormd dan op L94. Op een sla-genotype trad echter geen appressoriumvorming op. Op deze laatste niet-waardsoort ontbreken waarschijnlijk de stimuli die de schimmel in staat stellen huidmondjes te vinden.     

Barley lesion mimics,supersusceptible or highly resistant to leaf rust and net blotch

Lesion mimic mutants of plants have the feature of spontaneously displaying necrotic spots or bands on their leaves. Lesion mimics have often displayed enhanced resistance to biotrophic pathogens whilst showing increased susceptibility to necrotrophs. This paper identifies three novel, non‐allelic mutants of barley (Hordeum vulgare), which spontaneously form necrotic leaf lesions: Necrotic leaf spot 9.3091 (nec9.3091), Mottled leaf 8.1661 (mtt8.1661) and Mottled leaf 9.2721 (mtt9.2721). The Necrotic leaf spot 8.3550 mutant (nec8.3550), formerly known as bst1, was included in the study because it is a lesion mimic mutant belonging to the same original pool. The reactions of the mutants to the biotroph Puccinia hordei and the necrotroph Pyrenophora teres f. sp. teres were investigated. Mutants nec8.3550 and mtt8.1661 were more resistant than the parental Bowman near‐isogenic line with the Rph3.c gene (Bowman Rph3.c, NGB 22452) to leaf rust, caused by P. hordei. Mutants nec8.3550, mtt8.1661 and mtt9.2721 were more susceptible than Bowman Rph3.c to net blotch, caused by P. teres f. sp. teres. Autofluorescence was detected in leaf tissues of all mutants. Based on the high expression of the PR1 and Hv‐HIR genes, combined with the low susceptibility to P. hordei, nec8.3550 appears to have entered a state of systemic acquired resistance, which is quite distinct from the resistance expressed in mtt8.1661. The latter mutant has low or no expression of PR1 and Hv‐HIR genes, yet it is highly resistant to rust. It is also extremely susceptible to net blotch. These mutants can serve as genetic sources of novel disease resistance for barley improvement.     

Resistance in Australian barley (Hordeum vulgare) germplasm to the exotic pathogen Puccinia striiformis f. sp. hordei,causal agent of stripe rust

Eighty‐eight Australian and 10 international barley cultivars were assessed for resistance to the barley stripe (yellow) rust pathogen, Puccinia striiformis f. sp. hordei (Psh). All cultivars were tested for seedling resistance to two UK‐derived isolates of Psh (11.01 and 83.39) that were shown to differ in virulence based on responses on 16 differential barley genotypes. The 98 barley cultivars differed substantially in stripe rust response; 45% were susceptible to Psh 11.01, 53% to Psh 83.39 and 44% to both isolates. The observed diverse infection types (ITs) suggest the presence of both known and uncharacterized resistance. However, further multipathotype tests are required for accurate gene postulation. The Yerong × Franklin (Y×F) doubled haploid (DH) population was phenotypically assessed as seedlings using both Psh isolates. Yerong and Franklin were immune and highly resistant, respectively, to both isolates used in this study. Marker‐trait and QTL mapping identified a major effect on the long arm of chromosome 7H contributed by Franklin in response to all isolates. The resistance of Yerong was mapped to 113·96 and 169·38 cM on chromosome 5HL in response to Psh 11.01 and 83.39, respectively. The Psh resistance sources identified in this study can be used for further genetic analysis and introgression for varietal improvement.     

Virulence spectrum of Puccinia hordei in barley production systems in Ethiopia

Barley leaf rust caused by Puccinia hordei is an important disease of barley in Ethiopia. In the 2003 and 2004 cropping seasons, surveys of P. hordei were conducted on fields in the main rainy, residual moisture and short rainy season-barley production systems. A total of 381 isolates were analysed on 12 barley differential hosts carrying different Rph resistance genes ( Rph1 – Rph12 ). Based on infection phenotypes on leaf rust ( Rph ) resistance genes, seven pathotypes were identified, namely ETPh7611, ETPh7631, ETPh6611, ETPh7651, ETPh7671, ETPh7653 and ETPh7633, with frequencies of 63·0, 21·5, 6·8, 2·9, 2·6, 2·1 and 1·2%, respectively. ETPh7611 and ETPh7631 were the most common pathotypes found in all the surveyed areas of the three production systems. ETPh7653 was found in the small rainy season production system only. ETPh7671, ETPh7653 and ETPh7633 were the most virulent, but the least frequent, pathotypes. All isolates were virulent on resistance genes Rph1 , Rph4 , Rph8 , Rph9 , Rph11 and Rph12 . Virulence against Rph3 and Rph7 was absent. Genes Rph2 , Rph6+2 , Rph5 and Rph10 were effective against 96·3, 88·9, 65 and 2·4% of the rust isolates tested, respectively.     

Prehaustorial Resistance against Alfalfa Rust (Uromyces striatus) in Medicago truncatula

The legume species Medicago truncatula is gaining interests as a plant for structural and functional genomics that can be used to identify agronomically important genes in crop legumes. Resistance to the alfalfa rust (Uromyces striatus) was studied in a germplasm collection of M. truncatula. Accessions varied in resistance, as expressed by disease severity, but none showed macroscopically visible necrosis. Histological investigations, in selected lines covering the whole range of resistance reactions, revealed little difference in spore germination and none in orientation of germtubes on the leaf surface. However, appressorium formation on the stoma was significantly reduced in some accessions. Differences in resistance among accessions were more evident once the stoma were penetrated by the infection structures. Resistance was mainly due to a restriction of haustorium formation with varying levels of early abortion of the colonies, a reduction in the number of haustoria per colony, and hampered colony growth. In addition, necrosis of the host cells associated with infection hyphae was detectable in some accessions from the beginning of colony development. This information will be useful for eventual mapping and cloning analyses of resistance genes in M. truncatula that will in turn be useful for understanding other legume/rust interactions.     

A Greenhouse Test for Screening Sugar Beet (Beta Vulgaris) for Resistance to Rhizoctonia Solani

Rhizoctonia solani Kühn is a serious plant pathogenic fungus, causing various types of damage to sugar beet (Beta vulgaris L.). In Europe, the disease is spreading and becoming a threat for the growing of this crop. Plant resistance seems to be the most practical and economical way to control the disease. Experiments were carried out to optimise a greenhouse procedure to screen plants of sugar beet for resistance to R. solani. In the first experiment, two susceptible accessions were evaluated for root and leaf symptoms, after being grown in seven different soil mixtures and inoculated with R. solani. The fungus infected all plants. It was concluded that leaf symptoms were not reliable for the rating of disease severity. Statistically significant differences between the soil mixtures were observed, and there were no significant differences between the two accessions. The two soil mixtures, showing the most severe disease symptoms, were selected for a second experiment, including both resistant and susceptible accessions. As in the first experiment, root symptoms were recorded using a 1–7 scale, and a significant expression of resistance was observed. The average severity of the disease in the greenhouse experiment generally was comparable with the infection in field experiments, and the ranking of the accessions was the same in the two types of experiments. It was concluded that evaluation procedures in the greenhouse could be used as a rapid assay to screen sugar beet plants for resistance to R. solani.     

Partial stem and leaf resistance against the fungal pathogen <Emphasis Type="Italic">Botrytis cinerea</Emphasis> in wild relatives of tomato

Tomato (Solanum lycopersicum) is one of many greenhouse crops that can be infected by the necrotrophic ascomycete Botrytis cinerea. Commercial cultivation of tomato is hampered by the lack of resistance. Quantitative resistance has been reported in wild tomato relatives, mostly based on leaf assays. We aimed to identify wild tomato relatives with resistance to B. cinerea based on quantitative assays both on leaves and stem segments, monitoring infection frequency and disease expansion rate as parameters. A quantitative tomato stem segment assay was developed. This stem assay and a previously described leaf assay were used to screen a collection of 22 Solanum accessions. Significant differences in disease parameters were observed among accessions. Resistance to B. cinerea was observed in a number of wild Solanum species, including accessions of S. chilense, S. habrochaites and S. neorickii, both in the leaf assay and the stem segment assay. A number of resistant and susceptible accessions were evaluated as adult plants under greenhouse conditions. The data obtained in greenhouse assays confirmed the leaf and stem disease data. The expression of several defence-related genes was studied in a subset of accessions. There was no apparent correlation between the expression levels of the genes tested and the quantitative resistance level to B. cinerea. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Mechanism and molecular markers associated with rust resistance in a chickpea interspecific cross (Cicer arietinum × Cicer reticulatum)

A gene that controls resistance to chickpea rust (Uromyces ciceris-arietini) has been identified in a recombinant inbred line (RIL) population derived from an interspecific cross between Cicer arietinum (ILC72) × Cicer reticulatum (Cr5-10), susceptible and resistant to rust, respectively. Both parental lines and all RILs displayed a compatible interaction but differed in the level of infection measured as Disease Severity (DS) and Area Under the Disease Progress Curve (AUDPC). Histological studies of the seedlings of resistant parental Cr5-10 line revealed a reduction in spore germination, appressorium formation, number of haustoria per colony and colony size, with little host cell necrosis, fitting the definition of partial resistance. A Quantitative Trait Locus (QTL) explaining 31% of the total phenotypic variation for DS in seedlings and 81% of the AUDPC in adult plants in the field was located on linkage group 7 of the chickpea genetic map. The AUDPC displayed a bimodal distribution with high frequency of susceptible lines and both the AUDPC and markers showed the same distorted segregation. Consequently, it was hypothesised that a single dominant gene (proposed as Uca1/uca1) controlled resistance to rust in adult plants. This allowed us to locate the gene on the genetic linkage map. Two Sequence Tagged Microsatellite Sites (STMS) markers, TA18 and TA180 (3.9 cM apart) were identified that flank the resistance gene. These findings could be the starting point for a Marker-Assisted Selection (MAS) programme for rust resistance in chickpea.     

Differential Responses to Pea Bacterial Blight in Stems, Leaves and Pods Under Glasshouse and Field Conditions

Resistance to pea bacterial blight (Pseudomonas syringae pv. pisi) in different plant parts was assessed in 19 Pisum sativum cultivars and landraces, carrying race-specific resistance genes (R-genes) and two Pisum abyssinicum accessions carrying race-nonspecific resistance. Stems, leaves and pods were inoculated with seven races of P. s. pv. pisi under glasshouse conditions. For both race-specific and nonspecific resistance, a resistant response in the stem was not always associated with resistance in leaf and pod. Race-specific genes conferred stem resistance consistently, however, there was variability in the responses of leaves and pods which depended on the matching R-gene and A-gene (avirulence gene in the pathogen) combination. R2 generally conferred resistance in all plant parts. R3 or R4 singly did not confer complete resistance in leaf and pod, however, R3 in combination with R2 or R4 enhanced leaf and pod resistance. Race-nonspecific resistance conferred stem resistance to all races, leaf and pod resistance to races 2, 5 and 7 and variable reactions in leaves and pods to races 1, 3, 4 and 6.Disease expression was also studied in the field under autumn/winter conditions. P. sativum cultivar, Kelvedon Wonder (with no R genes), and two P. abyssinicum accessions, were inoculated with the most frequent races in Europe under field conditions (2, 4 and 6). Kelvedon Wonder was very susceptible to all three races, whereas P. abyssinicum was much less affected. The combination of disease resistance with frost tolerance in P. abyssinicum enabled plants to survive through the winter. A breeding strategy combining race-nonspecific resistance derived from P. abyssinicum with race-specific R-genes should provide durable resistance under severe disease pressure.     

Rhamnus lycioides in Tunisia is a new aecial host of oat crown rust

Puccinia coronata was not previously described on Rhamnus spp. in Tunisia. Three sites in the northwest of Tunisia, where Rhamnus is reported to be abundant, were surveyed for the presence of pycnia and aecia of oat crown rust caused by Puccinia coronata f. sp. avenae. Two Rhamnus species (R. lycioides and R. alaternus) were encountered in the sites. Pycnia with viable pycniospores and aecia with viable aeciospores were found on R. lycioides. However, no characteristic structures of crown rust were found on R. alaternus. Aeciospores collected from leaves of R. lycioides were used to inoculate oat plants usually susceptible to oat crown rust. Typical uredinia containing oat crown rust urediniospores appeared on the leaves of these plants. Moreover, the sixteen Pc-gene differential oat lines, used by oat researchers to study the virulence pattern in oat crown rust populations, were artificially inoculated with aeciospores from R. lycioides. These inoculated lines showed resistance/susceptibility similar to the registered resistance level of these lines to crown rust under field conditions in Tunisia. These results indicate that R. lycioides, a common and endemic part of the vegetation in the northwest of Tunisia, is a new aecial host of oat crown rust. The aeciospores produced on this forest plant could constitute the source of the virulence diversity already detected via the Pc-gene line trials.     

Genetics of host-pathogen interactions in the <Emphasis Type="Italic">Pyrenophora teres</Emphasis> f. <Emphasis Type="Italic">teres</Emphasis> (net form) – barley (<Emphasis Type="Italic">Hordeum vulgare)</Emphasis> pathosystem

The genetics of host-pathogen interactions in the Hordeum vulgare – P. teres f. teres pathosystem was studied in twelve resistant barley accessions, i.e. CI 9825, CI 9819, Diamond, CI 4922, CI 5401, Harbin, c-8755, c-21849, c-8721 c-23874, c-19979, c-15811. F₂ analyses of crosses with susceptible genotypes employing various isolates (from Europe, USA, Canada, and Australia) revealed that resistance is mostly isolate-specific and controlled by one or two genes. Segregation in ascospore progeny from two crosses between isolates of different origin revealed that avirulence in P. teres is also determined by one or two genes. An epistatic effect of suppressor genes on avirulence genes is proposed for the genetics of virulence to Diamond, Harbin, CI 5401 and c-8721 in the fungal crosses D (181-6 × A80) and F (H-22 × 92-178/9). Segregation in F₂ of crosses of three new sources of resistance (c-23874, c-19979, c-15811) to the susceptible cv. Pirkka was studied in laboratory and greenhouse tests by using seven P. teres isolates, i.e. 181-6, d8-3, d8-4, d9-1, d9-4, F4 and F74. In addition, virulence to these barley accessions of ascospore progeny from crosses of the same isolates was studied. Based on these studies it was concluded that depending on the isolate used, resistance of c-23874 is determined at least by two genes and in c-19979 and c-15811 by three genes. The results of this parallel analyses of genetics of resistance and genetics of virulence allows the postulation of a gene–for–gene interaction in the P. teres – H. vulgare pathosystem.     

Antioxidant,ethylene and membrane leakage responses to powdery mildew infection of near-isogenic barley lines with various types of resistance

Leaves of powdery mildew-susceptible barley (Hordeum vulgare cv. Ingrid) and related near-isogenic lines bearing various resistance genes (Mla12, Mlg or mlo5) were inoculated with Blumeria graminis f. sp. hordei race A6. Fungal attack induced several-fold increases in ethylene emission and electrolyte leakage in leaves of susceptible Ingrid beginning 3 days after inoculation. Activities of peroxidase, superoxide dismutase, glutathione S-transferase, ascorbate peroxidase and glutathione reductase enzymes were induced markedly in susceptible leaves 5–7 days after inoculation. Similar, but less pronounced pathogen-induced changes were detected in inoculated leaves of Mla-type resistant plants that show hypersensitive cell death upon inoculation, and, to an even lesser extent, in the Mlg and mlo lines, where no visible symptoms accompanied the incompatible interaction. Glutathione content increased only in susceptible barley 7 days after inoculation. Catalase activity, total ascorbate content and redox state were not influenced by inoculation in any of the genotypes. The activity of dehydroascorbate reductase was significantly reduced 3–5 days after inoculation in the susceptible parental plants and after 5 days in Mla and Mlg lines, while it was stable in the mlo barley. Slightly elevated levels of H₂O₂ were observed in the inoculated resistant plants. In contrast, H₂O₂ content decreased in the susceptible line 7 days after pathogen attack. These data indicate that high levels of antioxidants are involved in the compatible interaction of susceptible barley and powdery mildew by protecting the pathogen from oxidative damage.     

Relative tolerances of wild and cultivated barley to infection by Blumeria graminis f.sp. hordei (Syn. Erysiphe graminis f.sp. hordei). II—the effects of infection on photosynthesis and respiration

The relative levels of tolerance of two wild barley lines (Hordeum spontaneum), B19909 and I-17-40, and one cultivated barley (Hordeum vulgare), cv. Prisma, to Blumeria graminis f.sp. hordei were determined by comparing the effects of different levels of infection on the photosynthesis and respiration rates of the third leaf. Infection caused the early onset of senescence in all three lines, and in particular in cv. Prisma, and was accompanied by decreases in gross and net photosynthesis rates, increases in respiration rates, and loss of chlorophyll. The onset of senescence occurred at approximately the same time in infected leaves of the two wild lines, but once triggered, photosynthesis rates and chlorophyll levels declined more rapidly in I-17-40 than in B19909. A burst of respiratory activity accompanied the onset of senescence, and this was greatest in cv. Prisma. Conidial production was higher in B19909, indicating a higher level of tolerance in this line.     

Chromosomal location and genetic relationship of leaf rust resistance genes rph9 and rph12 in barley

ABSTRACT Barley lines Hor 2596 and Triumph are the sources of leaf rust resistance genes Rph9 and Rph12, respectively. An allelism test was performed with F(2) progeny of the cross Triumph/Hor 2596 inoculated with Puccinia hordei. No recombinants were found in a population of 3,858 progeny, indicating Rph9 and Rph12 are alleles. Molecular and morphological markers were used to identify the chromosomal location of these genes in the crosses Bowman/Hor 2596 and Triumph/I91-533-va. A linkage was detected between Rph9 and the flanking sequence-tagged site (STS) markers ABC155 and ABG3 on chromosome 7(5H) at a distance of 20.6 and 20.1 centimorgans (cM), respectively, and to the microsatellite marker dehydrin-9 (HVDHN9) at a distance of 10.2 cM in the Bowman/ Hor 2596 cross. Analysis of isozymes in bulks of the same population showed that Rph9 may be closely linked to the Est9 locus on chromosome 7(5H). The Rph12 locus was linked to the morphological trait locus va (controlling variegated leaf color) on chromosome 7(5H) at a distance of 22.6 cM in the Triumph/I91-533-va cross. Rph12 also was linked with STS marker ABC155 (24.4 cM) and RAPD marker OPA19 (1.5) (17.8 cM). These data indicate that Hor 2596 and Triumph carry a leaf rust resistance gene at the same locus on the long arm of chromosome 7(5H) of barley.     

四川58个小麦品种苗期抗条锈基因推导及成株期抗性表现

为了解四川小麦品种所含抗条锈基因及其在田间的抗性表现,将29个毒性谱各异的小麦条锈菌鉴别菌株于苗期接种35个已知抗条锈基因载体品系和58份四川小麦品种（系）,通过抗病谱对比分析和各品种（系）系谱分析,推导了四川小麦品种（系）的抗条锈基因型。在田间,选用中国小麦条锈菌优势小种CY32、CY33、Su-4和对Yr10、Yr24和Yr26致病的贵农22致病类型田间突变株进行混合接种,在乳熟期对各品种（系）的抗条锈性进行了鉴定。结果显示,Yr2、Yr5、Yr7、Yr9、Yr10、Yr24、Yr27、YrSpp、YrAlba等9个基因以单基因或基因组合的形式存在于44个四川小麦品种（系）中。有6个品种（系）含有Yr9基因,10个品种（系）含有Yr10基因,11个品种（系）含有Yr24基因,其中1个同时含有Yr10和Yr24基因。23个品种（系）含有未知基因及其组合。共有19个小麦品种（系）成株期对包含上述混合菌种在内的田间流行菌系表现抗病,其中5个品种抗性受成株抗性基因控制。