Path coefficient analysis of the effects of stripe rust and cultivar mixtures on yield and yield components of winter wheat

Four club wheat cultivars and three two-component cultivar mixtures, planted at five frequencies, were grown in three environments in both the presence and absence of stripe rust. The effect of stripe rust on wheat yield was through the yield components, with weight of individual seed being the component most affected by rust. In some cases, yield component compensation was indicated by the presence of negative correlations among the yield components. Path analysis of the yield components revealed that components with the highest correlations to yield also had the largest direct effects on yield. Of the yield components, number of heads per unit area exerted the largest direct influence on yield. The direct effects of number of seeds per head and weight of individual seed were similar, although number of seeds per head was more important in the absence of rust than in its presence. The pure stands and mixtures differed considerably with respect to correlation coefficients, but were very similar for direct effects of yield components on yield. Most of these discrepancies were due to opposing indirect effects, which were not evident from correlation coefficients alone.Paper No. 10,788 of the Journal Series of the Oregon Agricultural Experiment Station     

Mixing ability analysis of wheat cultivar mixtures under diseased and nondiseased conditions

Summary Mixing ability analyses, adapted from combining ability analyses used in plant breeding, were performed on yield and stripe rust (Puccinia striiformis) severity data for two-way mixtures among either four or five club wheat (Triticum aesitivum) cultivars grown in five environments. Initially, two statistics were calculated for each trait: general mixing ability (GMA), the average performance of a cultivar over all of the mixtures, and specific mixing ability (SMA), the deviation of a mixture from the estimated performance of the pair based on its average performance in mixtures. General mixing ability was further divided into two components: genotype performing ability (GPA), the innate ability of a cultivar to yield and resist disease in pure stand, and true general mixing ability (TGMA), the average ability of a cultivar to influence yield and disease when mixed with other cultivars. Significant mean squares for genotypes, GMA, SMA, and TGMA were found for all of the traits in most environments. Examination of TGMA and SMA revealed cultivars and cultivar combinations that were statistically better mixers than the others. Some of the significant effects were probably due to the use of cultivars that differed in height and stripe rust resistance, but for other combinations there was no apparent explanation for enhanced mixing ability.Paper No. 9132 of the Oregon Agricultural Experiment Station. Supported in part by USDA Grants 88-34106-3631 and 88-37151-3662     

Genetics of adult plant stripe rust resistance in CSP44, a selection from Australian wheat

Wheat line CSP44, a selection from an Australian bread wheat cultivar Condor, has shown resistance to stripe rust in India since the last twenty years. Seedlings and adult plants of CSP44 showed susceptible infection types against stripe rust race 46S119 but displayed average terminal disease severity of 2.67 on adult plants against this race as compared to 70.33 of susceptible Indian cultivar, WL711. This suggests the presence of nonhypersensitive adult plant stripe rust resistance in the line CSP44. The evaluation of F₁, F₂ and F₃ generations and F₆ SSD families from the cross of CSP44 with susceptible wheat cultivar WL711 for stripe rust severity indicated that the resistance in CSP44 is based on two genes showing additive effect. One of these two genes isYr18 and the second gene is not yet described.     

Resistance of wild wheat to stripe rust: Predictive method by ecology and allozyme genotypes

From 114 accessions of wild emmer wheat from 11 sites in Israel, known for their allozymic variation (Nevo & al. 1982), individual genotypes were tested for resistance to one isolate of stripe rust both in the seedling stage in a growth chamber and in the adult plant stage in the field. The results indicate that resistance to stripe rust in seedlings and adults are significantly correlated (r_s = 0.40, p < 0.001). Genetic polymorphisms of resistance to stripe rust vary geographically and are predictable by climatic, as well as allozymic markers. Three variable combinations of rainfall, evaporation, and temperature explain significantly 0.40–0.53 of the spatial variance in disease resistance to stripe rust, suggesting the operation of natural selection. Several allozyme genotypes are significantly associated with disease resistance. We conclude that natural populations of wild emmer wheat in Israel contain large amounts of disease resistance genes. These populations could be effectively screened and then utilized by the phytopathologist for identifying resistant genotypes and producing new resistant cultivars.Patterns of Resistance of Wild Wheat to Pathogens in Israel II.     

Stripe Rust Partial Resistance Increases Spring Bread Wheat Yield in South‐eastern Anatolia,Turkey

Stripe rust caused by Puccinia striiformis f.sp. tritici is the most serious disease of wheat globally including south‐eastern Anatolia of Turkey, where wheat originated. In this study, 12 spring wheat genotypes were artificially inoculated and preserved in two locations, Diyarbak?r and Ad?yaman, during the 2011–2012 season to investigate loss in yield and yield components. Genotypes were evaluated at the adult plant stage using two partial resistance parameters: final disease severity and area under the disease progress curve (AUDPC). AUDPC ranged from 14.8 to 860 in Diyarbak?r, and 74 to 760 in Ad?yaman. Yield loss ranged from 0.6 to 68.5% in Diyarbak?r and 9.8 to 56.8% in Ad?yaman. Genotypes G1, G5, G7 and G8 were found to lose less yield, while higher yield loss was observed in G3, G4 (Nurkent), G5 and G9 (Karacada?‐98). The highest loss in thousand kernel weight was observed in a susceptible cultivar Karacada?‐98 in Diyarbak?r followed by 43.4 and 24.4% in Ad?yaman. Test weight loss reached 8.89% in Diyarbak?r and 20.8% in Ad?yaman. Yield loss and AUDPC had a positive significant relationship. Based on the values of AUDPC, final disease severity and yield loss, three major clusters were formed for 12 wheat genotypes. Partially resistant genotypes were found to lose less grain yield and seemed to be stronger against severe stripe rust pressure.     

Competition in mixtures of susceptible and resistant genotypes of Chondrilla juncea differentially infected with rust

Plants of two genotypes of Chondrilla juncea (skeleton weed), one susceptible to and the other resistant to one isolate of the rust Puccinia chondrillina, were grown as pure and mixed populations both in the presence and absence of rust. Weights of individual plants were obtained at two harvests, one when the plants were rosettes and the other when flowering had begun. Distributions of plant weights of each genotype became progressively more positively skewed with time, with rust infection of plants of the susceptible genotype and with increasing competition between plants of both genotypes. The results show that genetic differences may be an important factor in determining which individuals become dominant or are suppressed in competing mixtures, and that differential disease pressure may alter dominance of individuals in plant populations.     

Impact of density and disease on frequency-dependent selection and genetic polymorphism: experiments with stripe rust and wheat

Frequency-dependent disease impacts may contribute to the maintenance of genetic diversity and sexual reproduction in plant populations. In earlier work with experimental wheat (Triticum aestivum) populations at a single density, we found that stripe rust (caused by Puccinia striiformis) created frequency-dependent selection on its host but competitive interactions between host genotypes reduced the potential for disease to maintain genetic polymorphisms in this highly self-pollinated species; the weaker competitor actually exhibited positive disease-mediated frequency-dependent selection. Based on these results we predicted that at low density, where the overall level of competition is lower, disease would have a stronger impact relative to competition and thus be more likely to maintain genetic polymorphisms; at low densities the greatest effect of disease for negative frequency-dependent selection should be seen in the weak competitor. Here we report on results with wheat stripe rust in which we altered both the frequency and density of host genotypes in factorial combinations of two-way mixtures where each host genotype was attacked by its own specialized race of rust. Within each density disease levels increased with genotype frequencies, creating frequency-dependent disease attack at all densities. Similarly, disease created negative frequency-dependent selection on its host at all densities, as a genotype’s fitness was often greater at low than high frequency when disease was present. Disease levels increased with plant density in 1997 but decreased in 1998. While increasing plant density reduced absolute fitness, presumably as a result of increased competition, a genetic polymorphism was not more likely to be maintained at low than high density as we had predicted. Within each density, the impact of disease was insufficient to reverse the slope of the relationship between absolute fitness and planted frequency from positive to negative for the less competitive host genotype, thus preventing the maintenance of a genetic polymorphism.     

Effects of stripe rust on the evolution of genetically diverse wheat populations

Summary Eighteen populations, composed of four wheat (Triticum aestivum) varieties that were originally mixed together at equal frequencies, were grown for one-to-three generations at two locations. In addition, pure stands of the four varieties were grown in each year. Populations were either exposed to two stripe rust (Puccinia striiformis) races, protected from stripe rust, or exposed to alternating years of diseased and disease-free conditions. Regression of the logit of a variety's frequency versus generation number was used to calculate the relative fitness of each variety in each population. These analyses suggest that the relative fitnesses of the wheat varieties were affected by disease and geographic location and were constant over time. However, frequency-changes of varieties in the mixtures were negatively correlated with their planting frequencies (0.0001 < P < 0.085 in 14 out of 16 cases), suggesting that fitnesses were frequency-dependent in both the presence and absence of disease. We hypothesize that failure to detect frequency-dependence of fitness in the logit analyses was due to a limited number of generations and a limited range of initial variety frequencies. This is supported by data from longer-term studies in the literature that provide evidence for frequency-dependence of fitness in plant mixtures. Analyses of currently available field data suggest that stable equilibria may be a more likely outcome for mixtures of varieties that are more closely related and/or more uniformly adapted to the environment in which they are grown.Paper No. 9820 of the journal series of the Oregon Agricultural Experiment Station.     

Control of Mn status and infection rate by genotype of both host and pathogen in the wheat take-all interaction

Take-all is a world-wide root-rotting disease of cereals. The causal organism of take-all of wheat is the soil-borne fungus Gaeumannomyces graminis var tritici (Ggt). No resistance to take-all, worthy of inclusion in a plant breeding programme, has been discovered in wheat but the severity of take-all is increased in host plants whose tissues are deficient for manganese (Mn). Take-all of wheat will be decreased by all techniques which lift Mn concentrations in shoots and roots of Mn-deficient hosts to adequate levels. Wheat seedlings were grown in a Mn-deficient calcareous sand in small pots and inoculated with four field isolates of Ggt. Infection by three virulent isolates was increased under conditions which were Mn deficient for the wheat host but infection by a weakly virulent isolate, already low, was further decreased. Only the three virulent isolates caused visible oxidation of Mn in vitro. The sensitivity of Ggt isolates to manganous ions in vitro did not explain the extent of infection they caused on wheat hosts. In a similar experiment four Australian wheat genotypes were grown in the same Mn-deficient calcareous sand and inoculated with one virulent isolate of Ggt. Two genotypes were inefficient at taking up manganese and were very susceptible to take-all, one was very efficient at taking up manganese and was resistant to take-all, and the fourth genotype was intermediate for both characters. All genotypes were equally resistant under Mn-adequate conditions.     

Comparing single- vs. mixed-genotype infections of Mycosphaerella graminicola on wheat: effects on pathogen virulence and host tolerance

Mixed-genotype infections (infections of a host by more than one pathogen genotype) are common in plant-pathogen systems. However their impact on the course of the infection and especially on pathogen virulence and host response to infection is poorly understood. We investigated the effects of mixed-genotype infections on several parameters: host resistance and tolerance, as well as pathogen aggressiveness and virulence. For these purposes, we inoculated three wheat lines with three Mycosphaerella graminicola genotypes, alone or in mixtures, in a greenhouse experiment. For some of the mixtures, disease severity and virulence were lower than expected from infection by the same genotypes alone, suggesting that competition between genotypes was reducing their aggressiveness and virulence. One host line was fully resistant, but there were differences in resistance in the other lines. The two host lines that became infected differed slightly in tolerance, but mixed-genotype infections had no effect on host tolerance.     

Disease spread of non-specialised fungal pathogens from inoculated point sources in intraspecific mixed stands of cereal cultivars

Spread of Septoria nodorum from inoculated point sources was examined in pure stands and mixtures of two spring wheat cultivars Kolibri and Maris Butler. Gradients in disease were observed soon after inoculation; the presence of the more resistant cultivar (Maris Butler) in the mixtures retarded the outward spread of disease compared with the susceptible pure stand (Kolibri). Regression analysis suggested that gradients in incidence were influenced by nearness to the source whereas gradients in severity were not. Spread of disease was also examined in pure stands and mixtures for the host/pathogen combinations, winter wheat (cvs Maris Huntsman and Maris Ranger)/S. nodorum and winter barley (cvs Maris Otter and Hoppel)/Rhynchosporium secalis. In contrast to the spring wheat experiment, no gradients were observed; explanations for the uniform distribution of disease were (a) extensive spread prior to the period of assessment, (b) no physical barrier to dispersal due to the juvenile growth stage of the crop and (c) exhaustion of the point-source.     

Crop Growth, Disease and Yield Components of Rusted Phaseolus Beans in Ethiopia

Crop growth and disease epidemics in sprayed and non-sprayed bean plots, artificially infected with rust (Uromyces appendiculatus) 3 weeks after emergence. were assessed weekly in two cultivars, at two locations for two seasons. Disease intensity was regulated by the application of a fungicide at 5 spray frequencies. Fungicide application influenced leaf area index (LAI) and reduced rust intensity. The fungicide had no significant effect on other diseases and dead leaf area. Fungicide application increased seed yield (SY) by increased numbers of pods per plant (PP). Rust severity was strongly correlated with pustule density but the overall relationships among rust assessment parameters depended on cultivar and location. Seed yield and pods per plant were highly correlated with LAI. The relationships between LAI and seeds per pod or seed weight depended on cultivar and location. Overall rust assessment parameters (rust severity and pustule density) showed close, negative relationships with seed yield. seed weight and pods per plant but not with seeds per pod. The relationships obtained in the partially resistant line 6-R-395 were less definite than those in the susceptible line Mexican 142. The yield parameters seed yield and pods per plant, showed strong positive relationships.     

Yield response of various wheat genotypes to inoculation withAzospirillum brasilense

Summary Twenty different wheat genotypes representing a wide genetic variability, were tested for their yield and yield components response to inoculation withAzospirillum brasilense, at two levels of N fertilization. Only two cultivars responded by a significant yield increase of 7.4 and 8.0 per cent — both at the higher N level. The response reflected an increase in the number of grains per plant added as additional spikes. The importance of the host plant genotype for a successful wheat-bacteria association is discussed.     

Selection of parents for crossing based on genotyping and phenotyping for stripe rust (<Emphasis Type="Italic">Puccinia striiformis</Emphasis>) resistance and agronomic traits in bread wheat breeding

Bread wheat (Triticum aestivum L.) germplasm consisting of 45 genotypes were clustered phenotypically using ten morphological traits and Area Under Disease Progress Curve (AUDPC) as measure of stripe rust resistance. The clustering was ratified by using twenty three molecular markers (SSR, EST and STS) linked to stripe rust (Puccinia striiformis f. sp. tritici) resistant QTLs. The aim was to asses the extent of genetic variability among the genotypes in order to select the parents for crossing between the resistant and susceptible genotypes with respect to stripe rust. The Euclidian dissimilarity values resulted from phenotypic data regarding morphological traits and AUDPC were used to construct a dendrogram for clustering the accessions. Using un-weighted pair group method with arithmetic means, another dendrogram resulted from the similarity coefficient values was used to distinguish the genotypes with respect to stripe rust. Clustering based on phenotypic data produced two major groups and five clusters (with Euclidian dissimilarity ranging from 2.44 to 16.16) whereas genotypic data yielded two major groups and four clusters (with percent similarity coefficient values ranging from 0.1 to 46.0) to separate the gene pool into highly resistant, resistant, moderately resistant, moderately susceptible and susceptible genotypes. With few exceptions, the outcome of both type of clustering was almost similar and resistant as well as susceptible genotypes came in the same clusters of molecular genotyping as yielded by phenotypic clustering. As a result seven genotypes (Bakhtawar-92, Frontana, Saleem 2000, Tatara, Inqilab-91, Fakhre Sarhad and Karwan) of diverse genetic background were selected for pyramiding stripe rust lesistant genes as well as some other agronomic traits after hybridization.     

Diallel analysis of the latent period of stripe rust in wheat

A half diallel was made amongst five wheat (Triticum aestivum L.) genotypes of which one was susceptible, while the others had adult-plant resistance, to stripe rust (Puccinia striiformis West.). The five parent and ten F₁ progeny were grown in the glasshouse and were inoculated with three rust pathotypes at the seedling stage. The latent period was measured on the first leaf. Two procedures were used to analyze the half diallel. Both methods showed that the average effects of alleles were of much greater importance than was dominance in conditioning resistance in response to two of the pathotypes, while for the third pathotype dominance was important. Resistance was conditioned by partial dominance for two pathotypes whereas for the third it was determined by full dominance. Broad-sense heritabilities range from 60–73% and the number of genes involved was different (from 1 to 4), depending on the pathotype.     

Studies of the incidence, severity and distribution of false rust (Synchytrium psophocarpi), leaf spot (Pseudocercospora psophocarpi) and powdery mildew (Oidium sp.) on winged bean (Psophocarpus tetragonolobus) in Papua New Guinea

The incidence, severity and distribution of false rust, (Synchytrium psophocarpi), leaf spot (Pseudocercospora psophocarpi) and powdery mildew (Oidium sp.) on winged beans (Psophocarpus tetragonolobus) were evaluated on pure lines and mixed stands grown in Papua New Guinea between 1976 and 1978. Almost all of the 148 pure lines were susceptible to false rust and leaf spot and the incidence of these diseases increased with plant age. Standard area diagrams were superior to disease keys in evaluating disease severity. Levels of false rust on 10 promising agronomic lines differed according to the locality where the lines were grown. Two Thai lines showed resistance to false rust but P. scandens was immune to false rust and leaf spot. Surveys of 51 sites containing mixed stands of winged beans in the Eastern Highlands, Western Highlands, East New Britain, East Sepik, Madang and Morobe Provinces of Papua New Guinea indicated that leaf spot and false rust were present in both the wet and dry seasons and the apparent absence of disease was probably due to time and size of samples. The diseases appeared to be present in all the traditional winged bean growing areas of Papua New Guinea. It was difficult to obtain evidence of differences in palatability of tissues infected with false rust. The value of surveys and disease assessment in relation to evaluation of crop losses of winged bean caused by these pathogens and selection for resistance is discussed.     

Genome-wide identification of QTL conferring high-temperature adult-plant (HTAP) resistance to stripe rust (<Emphasis Type="Italic">Puccinia striiformis</Emphasis> f. sp<Emphasis Type="Italic">. tritici</Emphasis>) in wheat

High-temperature adult-plant (HTAP) resistance to stripe rust (caused by Puccinia striiformis f. sp. tritici) is a durable type of resistance in wheat (Triticum aestivum L.). This study identified quantitative trait loci (QTL) conferring HTAP resistance to stripe rust in a population consisting of 169 F_8:10 recombinant inbred lines (RILs) derived from a cross between a susceptible cultivar Rio Blanco and a resistant germplasm IDO444. HTAP resistance was evaluated for both disease severity and infection type under natural infection over two years at two locations. The genetic linkage maps had an average density of 6.7 cM per marker across the genome and were constructed using 484 markers including 96 wheat microsatellite (SSR), 632 Diversity Arrays Technology (DArT) polymorphisms, two sequence-tagged-site (STS) from semi-dwarf genes Rht1 and Rht2, and two markers for low molecular-weight glutenin gene subunits. QTL analysis detected a total of eight QTL significantly associated with HTAP resistance to stripe rust with two on chromosome 2B, two on 3B and one on each of 1A, 4A, 4B and 5B. QTL on chromosomes 2B and 4A were the major loci derived from IDO444 and explained up to 47 and 42% of the phenotypic variation for disease severity and infection type, respectively. The remaining five QTL accounted for 7–10% of the trait variation. Of these minor QTL, the resistant alleles at the two QTL QYrrb.ui-3B.1 and QYrrb.ui-4B derived from Rio Blanco and reduced infection type only, while the resistant alleles at the other three QTL, QYrid.ui-1A, QYrid.ui-3B.2 and QYrid.ui-5B, all derived from IDO444 and reduced either infection type or disease severity. Markers linked to 2B and 4A QTL should be useful for selection of HTAP resistance to stripe rust.     

QTL mapping of stripe,leaf and stem rust resistance genes in a Kariega × Avocet S doubled haploid wheat population

Adult plant resistance to stripe (yellow) rust in the wheat cultivar Kariega has previously been ascribed to a major quantitative trait locus (QTL) on each of chromosomes 2B and 7D, along with a number of minor QTL. We have extended both the size of the cv. Kariega × cv. Avocet S mapping population, and the marker coverage within it, by assembling a set of Diversity Array Technology (DArT) markers. This has allowed for the analysis of the genetic basis of the adult plant and seedling resistances to stripe, leaf and stem rust present in the two mapping population parents. The stripe rust reactions of the segregating material were assessed in both field (three scoring dates) and greenhouse experiments. The chromosome 2B QTL became more important than the Lr34/Yr18 complex on chromosome 7D as the plants aged. As the infection progressed, the two QTL explained an increasing proportion of the variance for percentage leaf area infected. The cv. Kariega allele at the minor chromosome 4A QTL had a consistent effect on the severity of stripe rust infection and the overall plant reaction at the earlier scoring dates, but lost importance as the disease progressed. Several rust resistances were detected using an improved greenhouse-based test.     

Atmospheric teleconnection patterns associated with wheat stripe rust disease in North China

 Previous work has shown that long-term (>40 years) time series of wheat stripe rust disease from North China contain a signal related to the Southern Oscillation Index (SOI). However, no cause-and-effect relationships or direct links between SOI and disease have been established. Because mid-latitude teleconnection patterns form important links between global atmospheric variations and regional weather anomalies, such as the Southern Oscillation, studies of such patterns could result in a better physical and biological interpretation for the SOI-disease association observed previously. We used cross-spectral analyses to determine if and on which time scales severity of stripe rust in five regions of North China between 1952 and 1990 was associated with fluctuations in four Northern Hemisphere teleconnection patterns. These included the Pacific/North American, the Western Atlantic, the Western Pacific, and the Eurasian patterns. The analysis showed consistent and significant (P≤0.10) coherence relationships between the Western Atlantic (WA) pattern and stripe rust severity at a periodicity of 3.00 years. The phase relationships showed that, at the 3.00-year periodicity, the WA series and the disease series were out of phase by about half a period in all five regions. This phase difference indicated that peaks in the WA series, which are associated with below-normal winter temperatures over much of China, coincided with troughs in the disease series (i.e. low disease severity), presumably because of the negative effect of low winter temperatures on survival of the stripe rust pathogen. The analysis further showed that the WA series and the SOI series were highly significantly (P≤0.01) coherent at a periodicity of 4.33 years, which could explain the association between SOI and stripe rust severity reported previously. Received: 12 February 1997 / Accepted: 18 May 1998     

Disease progress of non-specialised fungal pathogens in intraspecific mixed stands of cereal cultivars. II. Field experiments

Epidemic development of Septoria nodorum was examined in pure stands and mixtures of two spring wheat cultivars, Kolibri and Maris Butler. In mixtures, disease development was reduced almost to that of the more resistant pure stand (Maris Butler). Disease levels, however, were low and no significant differences in grain yield per head were demonstrated. Disease development was further investigated for the host-pathogen combinations winter wheat/S. nodorum and winter barley/Rhynchosporium secalis. With both combinations the mixed stands again reduced disease levels to almost that of the more resistant pure stands. In general, mixed stands were more effective against R. secalis, although, with S. nodorum, disease levels were low. The yield response in mixed stands differed for the host cultivars and was not significantly changed by the presence of disease. The complexity of analysing such situations and the implications of these findings for mixtures of cultivars differing in resistance to non-specialised pathogens are discussed.