Deployment Of Resistance Genes Research Articles

Pathotypes of Pyricularia grisea in Rice Fields of Central and Southern China.

Blast, caused by the fungal pathogen Pyricularia grisea, is the most devastating disease of rice worldwide. Knowledge of the pathotype composition of the pathogen in rice fields is essential for rational deployment of resistance genes in rice breeding programs. In this study, we assayed the pathotypes of the pathogen populations using samples recently collected from 13 major rice-growing provinces of central and southern China. In all, 792 single spore isolates were tested for pathogenicity reactions using 13 host differentials consisting of six indica and seven japonica near-isogenic lines (NILs). The compositions of the pathogen populations were complex; 48 pathotypes were identified with the indica NILs, 82 pathotypes were detected with the japonica NILs, and a total of 344 pathotypes were identified with both indica and japonica NILs. There were large differences in distribution of the pathotypes among the different rice-growing areas. Even neighbor provinces seemed to differ sharply in types and frequencies of the most prevalent pathotypes. There was also a large difference in the frequencies of the isolates producing compatible reactions on the NILs, indicating the difference in frequencies of avirulence genes in the pathogen populations. The data provided very useful information for formulating strategies for improving blast resistance in rice breeding programs.

Biotype composition of Hessian fly (Diptera: Cecidomyiidae) populations from the southeastern, midwestern, and northwestern United States and virulence to resistance genes in wheat.

Twenty-three Hessian fly, Mayetiola destructor (Say), populations collected in the southeastern (Alabama and Mississippi), midwestern (Indiana), and northwestern (Idaho and Washington) United States from 1995 to 1999 were evaluated for biotype composition based on response to Hessian fly resistance genes H3, H5, H6, and H7H8 in wheat, Triticum aestivum L. Biotypes L and O, combined, made up at least 60% of all Alabama populations. Biotype L was predominant in the northern third of Alabama and biotype O in the southern two-thirds of the state. Based on biotype data, wheat cultivars with H7H8 resistance should be highly effective in central and southern Alabama. Fifty-four percent of the Mississippi population consisted of biotype L, and the remaining virulent biotypes (B, D, E, G, J, and O) ranged in frequency from 1 to 17%. The Mississippi population also contained 4% of the avirulent biotype GP. Only biotypes D and L were found in Indiana populations, but biotype L was predominant. Hessian fly populations from Idaho and Washington contained one or more of the virulent biotypes D-H, J, and L-O; however, only biotypes E, F, and G occurred at frequencies > 12%. The avirulent biotype GP made up 25-57% of Idaho and Washington populations, a much higher percentage than found in populations from the eastern United States. Although the highest level of virulence in Idaho and Washington populations was found to resistance genes H3 and H6, the frequency of biotype GP would indicate that the currently deployed gene H3 would provide a moderate to high level of resistance, depending on location. Nine of the populations, plus populations collected from the mid-Atlantic state area in 1989 and 1996, also were tested against the wheat cultivar 'INW9811' that carries H13 resistance to Hessian fly biotype L and two Purdue wheat lines with unidentified genes for resistance. The H13 resistance in INW9811 was highly effective against all populations tested from the eastern and northwestern U.S. wheat production areas, except Maryland and Virginia. Population studies also indicated that wheat line CI 17960-1-1-2-4-2-10 likely carries the H13 resistance gene, based on the similarity of its response and that of INW9811 to eight fly populations. Continued monitoring of biotype frequency in Hessian fly populations is required for optimal deployment and management of resistance genes in all wheat production areas.

Resistance gene deployment strategies in cereal hybrids using marker-assisted selection: gene pyramiding, three-way hybrids, and synthetic parent populations.

Marker-assisted selection (MAS) for resistance genes (R-genes), identified using molecular markers and quantitative trait loci (QTL) analysis, is now possible in many crops. MAS can be used to pyramid several R-genes into a single host genotype. However, this may not provide durable genetic resistance because the pathogen is exposed to a full homozygous pyramid during hybrid seed production and to a full heterozygous pyramid in the resultant hybrid. Alternative gene deployment strategies that generate genetic variability were analysed, for hybrid cereal cultivars of pearl millet, maize, sorghum and rice, using maintainer lines (B-lines) with two smaller complementary pyramids. An F1 seed parent, produced on two such B-lines, can be used to produce a three-way hybrid. All target loci are heterozygous for resistance alleles in the F1 seed parent, and the pathogen is exposed in the hybrid to a host population that is heterogeneous and heterozygous for alleles at the resistance loci targeted by MAS. Alternatively, single-cross hybrids can be made on seed parents that are maintained by two B-lines that differ for the complementary resistance gene pyramids. In a cross-pollinated crop, the B-lines are allowed to intermate to produce a synthetic B-line. In an inbreeding crop, the B-lines are equivalent to a two-component multiline variety. In inbreeding crops, because there is no intermating between the B-line components, the resultant synthetic seed parents have a higher frequency of genotypes with resistance alleles (R-alleles) at several resistance loci. However, in both cross-pollinated and inbreeding crops the genotypic structure in the hybrids is almost the same. All alternatives to a single-cross hybrid having a full pyramid produce hybrid cultivars having lower frequencies of resistance alleles. The frequency of genotypes having R-alleles at several loci increases greatly in both seed parent and hybrid when the overall frequency of R-alleles in the maintainer lines increases. This is simply done by adding a maintainer line that has a full pyramid or by the component lines having overlapping pyramids.

Epidemiology in Mixed Host Populations

ABSTRACT Although plant disease epidemiology has focused on populations in which all host plants have the same genotype, mixtures of host genotypes are more typical of natural populations and offer promising options for deployment of resistance genes in agriculture. In this review, we discuss Leonard's classic model of the effects of host genotype diversity on disease and its predictions of disease level based on the proportion of susceptible host tissue. As a refinement to Leonard's model, the spatial structure of host and pathogen population can be taken into account by considering factors such as autoinfection, interaction between host size and pathogen dispersal gradients, lesion expansion, and host carrying capacity for disease. The genetic composition of the host population also can be taken into account by considering differences in race-specific resistance among host genotypes, compensation, plant competition, and competitive interactions among pathogen genotypes. The magnitude of host-diversity effects for particular host-pathogen systems can be predicted by considering how the inherent characteristics of a system causes it to differ from the assumptions of the classic model. Because of the limited number of studies comparing host-diversity effects in different systems, it is difficult at this point to make more than qualitative predictions. Environmental conditions and management decisions also influence host-diversity effects on disease through their effect on factors such as host density and epidemic length and intensity.

Deployment of disease resistance genes by plant transformation – a ‘mix and match’ approach

Breeding for disease resistance has often resulted in the evolution of a matching virulence within the pathogen population, leading to an apparent ‘breakdown’ of resistance. In general, plant breeders have responded by introducing new genes for resistance, with similar consequences. This has led to ‘boom–bust’ cycles, where varieties possessing effective resistance are grown on an expanding acreage (boom) until matching virulence evolves and spreads within the pathogen population (bust). A variety of resistance genes have recently been identified and characterized in model systems. Together with the development of efficient plant transformation systems these genes offer an alternative means to introduce specific resistance into a crop improvement programme. However, unless the resistance genes are deployed with care, the boom–bust cycle is likely to be perpetuated.

Changes in the Racial Composition of Phytophthora sojae in Australia Between 1979 and 1996.

Surveys of commercial soybean fields, disease nurseries, and trial plots of soybean were conducted throughout eastern Australia between 1979 and 1996, and 694 isolates of Phytophthora sojae were collected and classified into races. Fourteen races, 1, 2, 4, 10, 15, and 25, and eight new races, 46 to 53, were identified, but only races 1, 4, 15, 25, 46, and 53 were found in commercial fields. Races 1 and 15 were the only races found in commercial fields in the soybean-growing areas of Australia up until 1989, with race 1 being the dominant race. Race 4 was found in central New South Wales in 1989 on cultivars with the Rps1a gene, and it is now the dominant race in central and southern New South Wales. Races 46 and 53 have only been found once, in southern New South Wales, and race 25 was identified in the same region in 1994 on a cultivar with the Rps1k gene. Only races 1 and 15 have been found in the northern soybean-growing regions, with the latter dominating, which coincides with the widespread use of cultivars with the Rps2 gene. Changes in the race structure of the P. sojae population from commercial fields in Australia follow the deployment of specific resistance genes.

Genetics of resistance to wheat leaf rust.

Leaf rust (caused by Puccinia recondita f. sp. tritici) is the most widespread and regularly occurring rust on wheat. Genetic resistance is the most economical method of reducing yield losses due to leaf rust. To date, 46 leaf rust resistance genes have been designated and mapped in wheat. Resistance gene expression is dependent on the genetics of host-parasite interaction, temperature conditions, plant developmental stage, and interaction between resistance genes with suppressors or other resistance genes in the wheat genomes. Genes expressed in seedling plants have not provided long-lasting effective leaf rust resistance. Adult-plant resistance genes Lr13 and Lr34 singly and together have provided the most durable resistance to leaf rust in wheat throughout the world. Continued efforts to isolate, characterize, and map leaf rust resistance genes is essential given the ability of the leaf rust fungus to overcome deployed resistance genes.

Diversity among Finnish net blotch isolates and resistance in barley

Seedlings of a differential barley (Hordeum vulgare L.) series (21 genotypes) and of six check genotypes were used in the greenhouse to assess variation in virulence among 20 single-spore isolates of the net blotch pathogen, Pyrenophora teres Drechs. f. teres Smedeg., collected from various sites in Finland. The experiment was run twice and symptom expression was recorded on the first three leaves. Analysis of second leaf symptom scores from Run 1 indicated differences in virulence between isolates, all of which were pathogenic, and differential resistance among the barleys. The virulence of P. teres isolates appeared to be conditioned by the host barley from which the isolate derived; the average virulence of isolates collected from a susceptible host was greater than that of isolates collected from a less susceptible host. Results from Run 2 were similar regarding resistance within the barleys, but variation in virulence among the P. teres isolates was not consistent with that from Run 1. CI 9819 carries duplicate genes for resistance to P. teres (Rpt1b and Rpt1c), and CI 7548 possesses Rpt3d. Both genotypes were highly resistant to all isolates. The Rptla gene of Tifang (CI 4407) conferred resistance to all isolates in Run 2, but only to about half of the isolates in Run 1. The checks, including two of the symptomatically most resistant Nordic barley genotypes, were universally susceptible during these stringent tests. No selective pressure has been placed on Finnish isolates of P. teres through previous deployment of major resistance genes, and it is speculated that any variation in virulence among isolates is likely to be due to a combination of evolutionary forces including, natural selection, random genetic drift and gene flow.

Molecular Approaches to Manipulation of Disease Resistance Genes

Various molecular approaches are greatly increasing our ability to characterize and manipulate disease resistance genes in plants. Molecular markers are allowing the dissection of monogenic and quantitative resistance. The utility of marker-aided selection in breeding for disease resistance is being evaluated. Markers will soon enhance the effective deployment of resistance genes to provide more stable resistance. Several resistance genes have been charac­ terized at the molecular level. This has led to the identification of several mechanistic classes and provides the opportunity for transgenic disease control strategies. As the mechanisms of detennining specificity and generating vari­ ation at resistance loci are understood in greater detail. they will be manipulated to generate new specificities. probably using microbial selection systems. Disease resistance strategies utilizing classical resistance genes will comple­ ment the use of novel trans genes.

Genetic Diversity of Xanthomonas oryzae pv. oryzae in Asia

Restriction fragment length polymorphism and virulence analyses were used to evaluate the population structure of Xanthomonas oryzae pv. oryzae, the rice bacterial blight pathogen, from several rice-growing countries in Asia. Two DNA sequences from X. oryzae pv. oryzae, IS1112, an insertion sequence, and avrXa10, a member of a family of avirulence genes, were used as probes to analyze the genomes of 308 strains of X. oryzae pv. oryzae collected from China, India, Indonesia, Korea, Malaysia, Nepal, and the Philippines. On the basis of the consensus of three clustering statistics, the collection formed five clusters. Genetic distances within the five clusters ranged from 0.16 to 0.51, and distances between clusters ranged from 0.48 to 0.64. Three of the five clusters consisted of strains from a single country. Strains within two clusters, however, were found in more than one country, suggesting patterns of movement of the pathogen. The pathotype of X. oryzae pv. oryzae was determined for 226 strains by inoculating five rice differential cultivars. More than one pathotype was associated with each cluster; however, some pathotypes were associated with only one cluster. Most strains from South Asia (Nepal and India) were virulent to cultivars containing the bacterial blight resistance gene xa-5, while most strains from other countries were avirulent to xa-5. The regional differentiation of clusters of X. oryzae pv. oryzae in Asia and the association of some pathotypes of X. oryzae pv. oryzae with single clusters suggested that strategies that target regional resistance breeding and gene deployment are feasible.

Biotype Status of Hessian Fly (Diptera: Cecidomyiidae) Populations from the Eastern United States and Their Response to 14 Hessian Fly Resistance Genes

Twenty-eight Hessian fly, Mayetiola destructor (Say), populations from 15 states in soft winter wheat area of the eastern United States were evaluated for biotype composition and response to 14 Hessian fly resistance genes. The biotype composition of each Hessian fly population was estimated by confining individual gravid females on the wheat differentials ‘Seneca’ (H7H8), ‘Monon’ (H3), ‘Abe’ or ‘Magnum’ (H5), and ‘Caldwell’ (H6). The response of Hessian fly populations to resistance genes not deployed in the eastern United States was evaluated in replicated tests with wheat containing H9, H10, H12, H13, H14H15, H17, H18, and H19+ genes and two Purdue germplasm lines with unknown sources of resistance (’ACC 1566-1-1-IX’ and ‘Giorgio 331-39-10’). Based on biotype determinations, H7H8 was resistant or moderately resistant to populations from the South and Southeast, H3 and H5 were resistant to populations from East Texas, H6 was susceptible to all but the ML-TX population, and no deployed genes were resistant to populations from most locations in the Midwest and in the Mid-Atlantic region and the Northeast. Predominant biotypes in Florida, Georgia, and South Carolina were E, G, and O, respectively. Biotype L was predominant in populations from the Midwest and some states of the Mid-Atlantic region. Texas populations were predominantly GP and A. The most resistant undeployed genes or gene combinations were H9, H13, H14H15, and H17, but Hessian fly virulence was found to all genes except H17. Continued monitoring of biotype frequency in Hessian fly populations is required for optimal deployment and management of resistance genes.

Wheat stem rust in Australia dash 1969-1985

Pathogenic changes in Puccinia graminis f. sp. tritici (P. g. tritici) in Australia during the period 1969-1985 are reviewed. As found in a previous study covering the period 1954-1968, the frequencies of individual pathotypes were influenced by the combinations of resistance genes present in the most common wheat cultivars. From 1969 to 1978, pathotype composition and diversity varied between the four regions comprising the Australian wheat-growing area. However, from 1978 to 1985, pathotype 343-1,2,3,5,6 predominated in all regions. It is suggested that this pathotype was repeatedly derived from pathotype 326-1,2,3,5,6 at widely separated locations, and that these events facilitated its widespread increase. Another significant event was the specialization of P. g. tritici on triticale. Survey data suggest that pathotype 34-2,12, which rendered cultivar Coorong susceptible, developed from pathotype 34-2, possibly during 1979. A second pathotype which attacked cultivar Satu (designated 34-2,12,13) was detected in 1984. The deployment of different resistant genes in triticale and wheat has resulted in distinct populations within P. g. tritici adapted to each crop. Future Cultivar development for triticale should be aimed at maintaining this divergence.

Effectiveness of Deploying Single Gene Resistances in Wheat for Controlling Damage by the Hessian Fly (Diptera: Cecidomyiidae)

Discussions and simulation models have been used by others to develop strategies for deploying genes in wheat ( Triticum spp. ) for resistance to the Hessian fly, Mayetiola destructor (Say). Effects of deploying genes one at a time in sequence and redeploying a gene after a 10-yr absence from the commercial crop in Indiana during 1955–1989 are presented. Three genes, H3, H6 , and H5 , were deployed in 1955, 1962, and 1971, respectively, and H6 was redeployed in 1981. A reduction in infestation levels of Hessian fly occurred for 6–8 yr after each deployment. Redeployment of H6 , after a 10-yr absence in commercial use, was highly effective. Other methods of deploying resistance genes in conjunction with cultural practices should be considered for control of Hessian fly. When considering the pyramiding of genes for resistance, one must evaluate the time required to develop new productive cultivars. Efficient methods to determine that all of the pyramided genes are present in a line are important for the successful use of pyramided genes. Wheat cultivars are seldom grown for more than 8–10 yr, but the same pyramided genes might be used in subsequent cultivars.

Genetic Model for Wheat/Hessian Fly (Diptera: Cecidomyiidae) Interaction: Strategies for Deployment of Resistance Genes in Wheat Cultivars

A genetic model describing the interaction between the wheat plant, Triticum aestivum L., and the Hessian fly, Mayetiola destructor (Say), is presented. The model is based on multiplicative interlocus interaction and gene-for-gene interaction between host and insect. It can be used to predict changes in allelic frequency in the insect and in the level of potential crop damage caused by deployment of host resistance genes, given genotypic fitnesses and initial allelic frequencies. Given reliable estimates of genetic parameters, the model can be used to make predictions where there are unequal genetic effects and frequencies and a much wider range of resistance-gene deployment patterns.

Recommendations on strategies for the deployment of resistance genes

Recommendations on strategies for the deployment of resistance genes