Effect of early infection on pathotype frequencies in barley powdery mildew (Blumeria graminis f.sp. hordei) populations in field plots

A field experiment with barley powdery mildew ( Blumeria graminis f.sp. hordei ) was designed in order to study how the time of arrival of inoculum in the field influenced pathotype frequencies in the resulting populations. Three isolates belonging to pathotypes that were absent or rare in the local aerial inoculum were used to inoculate field plots of winter barley cv. Plaisant. Two successive inoculations with different combinations of the three isolates were performed with an approximately two-generation delay, and frequencies of inoculated pathotypes were assessed four and nine generations after the first inoculation. Pathotypes of the first inoculated isolates generally persisted throughout the period of sampling; this is described as an 'early arrival' effect. During the epidemics the inoculated isolates were not replaced by isolates from the natural airborne inoculum. Pathotype frequencies depended mainly on the time of arrival of inoculum in the plot, but frequencies also depended on the isolate that had been inoculated. The most frequent isolate, GL1, belonged to the clonal lineage dominant in powdery mildew populations on winter barley in the north of France. These results confirmed that the composition of a powdery mildew population in a field is largely determined by the composition of the initial inoculum.     

Variation in aggressiveness is detected among Puccinia triticina isolates of the same pathotype and clonal lineage in the adult plant stage

Puccinia triticina reproduces asexually in France and thus individual genotype is the unit of selection. A strong link has been observed between genotype identities (as assessed by microsatellite markers) and pathotypes (pools of individuals with the same combination of qualitative virulence factors). Here, we tested whether differences in quantitative traits of aggressiveness could be detected within those clonal lineages by comparing isolates of identical pathotype and microsatellite profile. Pairs of isolates belonging to different pathotypes were compared for their latent period, lesion size and spore production capacity on adult plants under greenhouse conditions, with a high number of replicates. Isolates of the same pathotype showed remarkably similar values for the measured traits, except in three situations: differences were obtained within two pathotypes for latent period and within one pathotype for sporulation capacity. One of these differences was tested again and confirmed. This indicates that the average aggressiveness level of a leaf rust pathotype may increase without any change in its virulence factors or microsatellite profile.     

鲁豫皖交界地区四个小麦禾谷孢囊线虫群体致病型鉴定

为了明确鲁豫皖交界地区小麦禾谷孢囊线虫的致病类型,利用23个国际鉴别寄主品种和1个当地小麦品种温麦19对采自山东菏泽、安徽颍上、河南商丘和淮阳的4个小麦禾谷孢囊线虫群体致病型进行了鉴定。4个群体的致病型不同于国际上已命名的16个小麦禾谷孢囊线虫致病型。13个A组鉴别寄主对山东菏泽与安徽颍上群体具有相同的抗性反应,这2个线虫群体属于同一致病型;河南商丘群体与上述2个群体的毒性相似,但大麦寄主KVL191对山东菏泽群体和安徽颍上群体表现为抗病,而对河南商丘群体表现感病;河南淮阳群体的致病谱较宽,明显不同于其它3个群体。对照小麦品种温麦19对4个小麦禾谷孢囊线虫群体均表现感病。     

Distribution of pathotypes with regard to host cultivars in French wheat leaf rust populations

ABSTRACT Isolates of wheat leaf rust collected from durum and bread wheat cultivars in France during 1999-2002 were analyzed for virulence on 18 Thatcher lines with single genes for leaf rust resistance (Lr genes). Sampling focused on the five most widely grown bread wheat cultivars (two susceptible and three resistant) to allow statistical comparison of diversity indexes between the cultivars. Leaf rust populations from durum and bread wheats were different. The diversity of the bread wheat leaf rust pathotypes, as measured by the Shannon index, ranged from 2.43 to 2.76 over the 4 years. Diversity for wheat leaf rust resistance was limited in the host since we postulated only seven seedling resistance genes in the 35 cultivars most widely grown during 1999-2002. Leaf rust populations were strongly differentiated for virulence within bread wheat cultivars, and diversity was higher on those that were resistant, mainly due to a more even distribution of virulence phenotypes than on susceptible cultivars. The pathogen population on the susceptible cv. Soissons was largely dominated by a single pathotype (073100), whereas all other pathotypes virulent on cv. Soissons either decreased in frequency or remained at a low frequency during the period studied. Several pathotypes including the most complex one were found only on resistant cultivars, even though most of them were virulent on the susceptible cv. Soissons. Specific interactions were necessary, but not always sufficient, to account for pathotype distribution and frequencies on the cultivars, suggesting that selection for virulence to host resistance genes is balanced by other selective forces including selection for aggressiveness.     

Is virulence phenotype evolution driven exclusively by Lr gene deployment in French Puccinia triticina populations?

Puccinia triticina is a highly damaging wheat pathogen. The efficacy of leaf rust control by genetic resistance is mitigated by the adaptive capacity of the pathogen, expressed as changes in its virulence combinations (pathotypes). An extensive P. triticina population survey has been carried out in France over the last 30 years, describing the evolutionary dynamics of this pathogen in response to cultivar deployment. We analysed the data set for the 2006–2016 period to determine the relationship between the Lr genes in the cultivars and virulence in the pathotypes. Rust populations were dominated by a small number of pathotypes, with variations in most of the virulence frequencies related to the corresponding Lr gene frequencies in the cultivated landscape. Furthermore, the emergence and spread of a new virulence matched the introduction and use of the corresponding Lr gene (Lr28), confirming that the deployment of qualitative resistance genes is an essential driver of evolution in P. triticina populations. However, principal component analysis (PCA) revealed that certain pathotype–cultivar associations cannot be explained solely by the distribution of Lr genes in the landscape. This conclusion is supported by the predominance of a few pathotypes on some cultivars, with the persistence of several other compatible pathotypes at low frequencies. Specific interactions are not, therefore, sufficient to explain the distribution of virulence in rust populations. The hypothesis that quantitative interactions between P. triticina populations and bread wheat cultivars—based on differences in aggressiveness—is also a driver of changes in pathotype frequencies deserves further investigation.     

Seasonal changes in pathotype complexity in French populations of barley powdery mildew

Airborne conidia of Erysiphe graminis f.sp. hordei were sampled in three regions and a single locality in the northern part of France for 2 years. Sampling was carried out in early spring, in late spring and in autumn, in order to separate the effects of winter barley cultivars, carrying few specific resistance alleles, and of spring barley cultivars, carrying diverse resistance alleles, on the structure of the pathogen population. Although complex pathotypes with three to 10 virulences were selected by spring cultivars, simple pathotypes, including a pathotype with the single unnecessary virulence allele Va22 , which formed a clear majority of the samples, remained dominant in early spring, when winter but not spring cultivars were growing. In early spring, simple pathotypes were more prevalent in the north, where the winter cultivars represented 90% of the barley acreage, than in the east, where winter cultivars represented 65%. In the west, the frequency of simple pathotypes was limited compared to the north, possibly because of the resistance allele Mlg in winter cultivars. The high frequency of simple pathotypes in early spring could be explained by a differential adaptation between simple and complex pathotypes or by delayed epidemics on spring cultivars compared to winter cultivars.     

Improved Selection with Newly Identified RAPD Markers Linked to Resistance Gene to Four Pathotypes of Colletotrichum lindemuthianum in Common Bean

ABSTRACT Three F(2) populations derived from crosses between the resistant cultivar AB 136 and the susceptible cultivar Michelite (MiA), and one F(2) population derived from a cross between AB 136 and Mexico 222 (MeA), were used to identify markers linked to anthracnose resistance genes present in cultivar AB 136. Primer OPZ04 produced a DNA band (OPZ04(560)) linked in coupling phase to the resistance gene for pathotype 89 (8.5 +/- 0.025 cM) in one population derived from the cross MiA. In the same population, primer OPZ09 produced one band (OPZ09(950)) linked in repulsion phase (20.4 +/- 0.014 cM) to the same resistance gene. The simultaneous use of markers in coupling and in repulsion phases allowed the identification of the three genotypic classes. In the other two populations from cross MiA, OPZ04(560) was linked in coupling phase to resistance genes for pathotypes 73 (2.9 +/- 0.012 cM) and 81 (2.8 +/- 0.017 cM). In population MeA, OPZ04(560) was linked in coupling phase (7.5 +/- 0.033 cM) to resistance to pathotype 64. These data suggest that a single gene or complex locus of linked resistance genes present in cultivar AB 136 confers resistance to all four pathotypes of C. lindemuthianum.     

Changes in specific virulence in Polish populations ofPhytophthora infestans: 1985–1991

Ninety-five isolates ofPhytophthora infestans collected throughout Poland during 1985–1991 and characterized for multilocus genotypes based on mating type, allozymes and DNA fingerprint, were analyzed for specific virulence to differential potato cultivars carrying ten major resistance genes. The multilocus analysis led to three groupings. The first group contained 22 isolates of a clonal lineage (PO-1) that is postulated to have been present in Europe during most of the twentieth century, but PO-1 isolates were recovered in Poland only during 1985–1988. This group contained, on average, virulence to 5.5 specific resistance genes per isolate. The second group consisted of 30 isolates in a clonal lineage (PO-4) that had not been detected before 1988. PO-4 isolates had virulence to a mean of 6.5 resistance genes per isolate. The third group was composed of 43 isolates representing 38 multilocus genotypes also not detected before 1988. These diverse genotypes had virulence to an average of 6.7 specific resistance genes per isolate. More than half (53%) of the PO-4 isolates shared a single pathotype. The group of 43 isolates was dominated by two pathotypes: the most common one (47% of the isolates) was the same pathotype that dominated PO-4 isolates; the next most common one (21%) differed from the most common one by the absence of virulence to resistance gene R5. The recent immigrant isolates (not detected before 1988) generally had virulence to a greater number of specific resistance genes than did isolates in the previous population [detected before 1988 (PO-1)]. Recent immigrant populations were dominated by one or two pathotypes, so their pathotypic diversity values were somewhat less than that of the previous population.     

Virulence and Molecular Diversity in Cochliobolus sativus

ABSTRACT Spot blotch, caused by the fungal pathogen Cochliobolus sativus, is an important disease of barley in many production areas of the world. To assess genetic diversity in this pathogen, a worldwide collection of C. sativus isolates was evaluated for virulence on barley and DNA polymorphism. Three pathotypes (0, 1, and 2) were identified among the 22 isolates tested in this study and the 36 isolates characterized previously on three barley differentials (ND5883, Bowman, and NDB112) that differ in their resistance to C. sativus. Pathotype 2, which exhibits high virulence on cv. Bowman, was only found in North Dakota, whereas the other two pathotypes occurred in many other regions of the world. Genetic diversity of the 58 C. sativus isolates, together with isolates of three related pathogenic Cochliobolus spp. (C. heterostrophus, C. carbonum, and C. victoriae) was analyzed using amplified fragment length polymorphism (AFLP) markers. A total of 577 polymorphic AFLP markers were recorded among the 70 isolates of the four Cochliobolus spp. using eight primer combinations. Cluster analysis revealed distinct groups corresponding to the four different species, except in one case where race 0 of C. carbonum was placed in an outgroup that may belong to a different species. In C. sativus, 95 polymorphic AFLP markers were detected with the eight primer pairs used, and each isolate exhibited a unique AFLP pattern. Allelic diversity in the pathotype 2 group was lower (0.10) than in the pathotype 0 (0.23) and pathotype 1 (0.15) groups, indicating that pathotype 2 may have arisen more recently. Cluster analysis did not reveal a close correlation between pathotypes and AFLP groups, although two AFLP markers unique to pathotype 2 isolates were identified. This low correlation suggests that genetic exchange may have occurred through parasexual recombination in the fungal population. Some isolates collected from different regions of the world were clustered into the same AFLP group, suggesting that migration of the fungal pathogen around these regions has occurred.     

Problems of potato cyst nematode pathotypes in the Czech Republic

During 1983–1989 about 1,200 infested plots in the Czech republic were investigated for the occurrence of new pathotypes of the potato cyst nematodes (PCN) Globoderu rostochiensis and G. pallida. Only eight populations of the pest were suspected of containing another pathotype than Ro1. Glasshouse pot trials were performed with those eight natural populations, two populations of Ro1, 10 potato cultivars resistant to Ro1 and 10 cultivars resistant to more pathotypes, G. pallida included. The multiplication rates (Pf/Pi) showed that four of the PCN populations were probably mixtures, with pathotype Ro1 prevailing. Three populations were suspected of containing G. pallida , but their morphological characters corresponded only with G. rostochiensis. One population corresponded with pathotype Ro5. The extent of the occurrence of pathotypes or virulence groups different from Ro1 can be estimated at 0.3–0.4% of infested plots.     

Physiological specialization and pathotype distribution of Puccinia recondita in western Europe, 1995

A pathogenicity survey of Puccinia recondita f.sp. tritici ( Prt ) was conducted in western Europe in 1995. Random urediospore isolates (850) of Prt were collected from the air by means of a jet spore sampler in wheat-growing regions of Austria, Belgium, France, Germany, northern Italy, Switzerland and the UK. Pathogenicity of the isolates was determined in tests of detached primary leaf segments maintained on water agar supplemented with benzimidazole (35 p.p.m.). The differential genotypes used were Thatcher, 20 near-isogenic Thatcher lines each with a single leaf rust resistance gene, and five cultivars/lines with additional resistance genes. All isolates were avirulent for the genes Lr9, Lr19, Lr21, Lr24, Lr25 and Lr29 , and both virulence and avirulence were detected for the remaining 19 genes. Fifty-three pathotypes were identified, four of which predominated (64% of isolates) and were widespread throughout western Europe. Three of the four predominant pathotypes were also identified in collections of wheat leaf rust collected in Poland, Hungary, Estonia and Finland. One pathotype, which comprised 35% of isolates in the south of France, was not detected in any other region. This pathotype was indistinguishable from several isolates obtained from Morocco, which suggested that it may have originated from northern Africa. Comparisons with previously published data suggested that the four predominant pathotypes were very similar and possibly the same as pathotypes present in the former Czechoslovakia for up to 20 years. The results obtained provide evidence of migration of Prt over considerable distances in western Europe, stressing the need for a co-ordinated approach for genetical control of the disease in this region.     

The effect of fungicide dose on the composition of laboratory populations of barley powdery mildew

The effect of the use of different doses of the fungicide fenpropimorph on populations of barley powdery mildew Blumeria ( Erysiphe ) graminis f. sp. hordei was investigated in a laboratory selection experiment. A sample from the Danish aerial population of powdery mildew was split into populations, and these were kept separately for 31 generations on susceptible barley seedlings treated with fungicide at two concentrations, as well as on a control. Samples from these populations were tested for their resistance to fenpropimorph and their virulence spectra. There was a large amount of environmental variation in the ED₅₀ values used to measure fungicide resistance. In both treated populations, the average level of fungicide resistance increased, this increase being faster and greater in the population treated with the high dose. The diversity of pathotypes of the treated populations decreased, with the decline being more rapid in the population treated with the high dose, where one pathotype dominated the population after 31 generations. This pathotype was apparently not the fittest in the population treated with the low dose. This implies that knowledge of ED₅₀ is not sufficient to predict pathotype evolution under different fungicide treatments. The dominant pathotype in the high-dose treatment may not have been clonal, as there was evidence of two levels of fungicide resistance. The large environmental variation observed in estimated ED₅₀ values for resistance towards fenpropimorph may help to explain why this resistance has evolved at a slower rate than resistance towards other fungicides.     

Low pathotype diversity in a recombinant Puccinia striiformis population through convergent selection at the eastern Himalayan centre of diversity (Nepal)

Worldwide Puccinia striiformis f. sp. tritici (Pst) epidemics have been reported to be driven by few genetic lineages, while a high diversity is evident at the Pst Himalayan centre of diversity. This study investigated the relationship between pathotype diversity and genetic structure in Nepal, the eastern Himalayan region, which has been largely unexplored. Despite the high genetic diversity and recombinant structure detected through microsatellite genotyping, characterization of virulence phenotypes for 62 isolates identified only eight pathotypes, with two pathotypes predominant over all the populations. This is in contrast to the Pakistani and Chinese recombinant populations, where high pathotype diversity is associated with genetic diversity. The most prevalent Nepali pathotype was not a unique clonal lineage, but was represented by seven multilocus genotypes from four distinct genetic subgroups, suggesting strong directional selection on virulence genes, resulting in convergent pathotypes in distinct genetic groups. This convergent selection is discussed in comparison with clonal French and recombinant Pakistani populations. Additionally, the Nepali Pst population carried virulence to 17 out of 24 tested yellow rust resistance genes (Yr), with the absence of virulence to Victo and Early Premium and resistance genes Yr5, Yr10, Yr15, Yr24 and Yr26. Virulence to Yr2, Yr7, Yr27 and YrSu were fixed in all isolates, in line with the deployment of these resistance genes in Nepal. The results reflect the influence of resistance gene deployment on selection of virulence and pathotypes in a recombinant pathogen population, which must be considered in the context of durable resistance gene deployment.     

Studies on the origin,spread, and evolution of an important group ofPuccinia recondita f. sp.tritici pathotypes in Australasia

Wheat brown rust pathotype (pt) 104-2,3,(6),(7), 11 was first detected in Australasia in Victoria during 1984. Although it appeared similar to a pre-existing pathotype, 104-2,3,6,(7), detailed greenhouse test revealed nine pathogenic differences between the two rusts. Six differences involved contrasting virulence/avirulence for the resistance genes/specificitiesLr12, Lr27+Lr31 andLr16, and three uncharacterised genes, present in the wheat cultivars Gaza and Harrier, and in triticale cultivar Lasko. Differences in partial virulence between the pathotypes were found for the genesLr2a, Lr13 andLr26. A comparison of the phenotypes for 13 isozyme systems in the two pathotypes revealed two differences, including aPgm2 allele in pt 104-2,3,(6),(7),11 not found in other contemporary AustralasianPuccinia recondita f. sp.tritici pathotypes. On the basis of these differences, it was concluded that pt 104-2,3,(6),(7),11 was introduced into the Australasian region before or during 1984.Seven variants of pt 104-2,3,(6),(7),11, that differed by single virulences, were detected during 1984–1992. Pt 104-2,3,(6),(7),11 and a derivative pathotype with virulence forLr20 underwent rapid increases in frequency, largely displacing pathotypes which predominated before 1984. Although first detected in eastern Australia, both pathotypes spread to New Zealand, and the derivative pathotype appeared in Western Australia. The rapid spread and increase of these pathotypes could not be explained by host selection. Pt 104-2,3,(6),(7),11 and derivatives may therefore be more aggressive than other contemporary Australasian pathotypes.Abbreviations NSW New South Wales - Prt Puccinia recondita f. sp.tritici - Qld Queensland - SA South Australia - WA Western Australia     

云南省水稻细菌性条斑病菌的致病型划分和水稻抗性资源的鉴定

 为明确云南省水稻细菌性条斑病菌(Xanthomonas oryzae pv. oryzicola, Xoc)的致病力分化以及不同类型水稻品种对Xoc的抗感特性,通过针刺接种法将云南省8个稻区采集的86株Xoc菌株,接种于6个携带不同抗性基因的水稻鉴别品种(IRBB4、IRBB5、IRBB14、IRBB18、IRBB21和IR24)。根据这些菌株在鉴别品种上的毒力差异进行了UPGMA聚类分析,将其划分为9个致病型(Ⅰ型 ~ Ⅸ 型)。其中,Ⅰ型为优势菌群,分布频率为50.5%。对不同稻区的优势菌群进行分析,发现云南省各稻区Xoc的致病型呈多样性分布,以强毒力的Ⅰ型为高频率致病型。选用 Ⅰ 型、Ⅱ 型和 Ⅵ 型代表菌株对云南省的80个主栽和区试水稻品种进行抗性评价,对3个致病型表现抗性的材料比例分别为30.0%、35.0%和57.5%。筛选出9个对3种致病型都表现为抗性的品种,其中“Deyou16”和“Changgui2”表现为高抗。研究结果可为云南省防治水稻细菌性条斑病的水稻区域性布局和抗性品种的利用提供理论依据。     

Virulence spectrum in populations of the barley powdery mildew pathogen, Erysiphe graminis f. sp. hordei in Tunisia and Morocco in 1992

The virulence of 57 single colonies of Erysiphe graminis f. sp. hordei from Tunisia and Morocco was investigated using the Pallas near-isogenic differential set. Virulence patterns in general were similar in both countries. Va8, Va10+Du2, V41/145, VLa and VRu2 appear to be common in the region. The resistance alleles Mla7 and Mla9, in combination with other resistances, were highly effective against the isolates tested. No virulence against mlo and Mla9 + k was detected in Tunisia. No virulence on Mla7 and Mla9 was detected in Morocco. The frequency of virulence against several resistances was significantly higher in Morocco than in Tunisia. On the other hand, virulence against other resistances was higher in Tunisia than in Morocco. Three isolates from Morocco infected mlo to an extent greater than previously described. All pathotypes were unique. An attempt is made to interpret the results by comparison with pathotype evolution in Europe.     

Development of simple sequence repeat markers for the classification of the clubroot pathogen <Emphasis Type="Italic">Plasmodiophora brassicae</Emphasis>

Clubroot disease caused by Plasmodiophora brassicae is one of the most serious diseases in cruciferous crops. To classify isolates, we developed simple sequence repeat (SSR) markers for P. brassicae. Twenty-four Japanese isolates were used in this study: 12 isolates of an unknown pathotype from the Kyoto Prefecture, as well as 12 isolates of known pathotypes, including three single-spore lines. From the 12 isolates from Kyoto Prefecture, 11 were classified into either pathotype 2 (three isolates) or 4 (eight isolates). We designed 23 SSR markers based on the P. brassicae genome, of which 11 markers from intergenic regions showed polymorphisms in the 24 isolates. Many haploid isolates belonging to pathotypes 2 and 4 were monomorphic, and typical alleles were detected in some isolates not belonging to pathotype 4. Two bands were detected for eight SSR loci in five isolates, indicating that different genotypes were mixed in these isolates. We constructed a phylogram based on the 11 polymorphic SSRs. Pathotypes 2 and 4 formed a cluster, from which pathotypes 3 and 1 were successively placed. These results strongly suggest a close genetic relationship between isolates in pathotypes 2 and 4, consistent with our finding that isolates in these two pathotypes were found at one collection site. In combination with pathotype classification and other marker systems, the SSR markers can be used for more detailed analyses to improve the control of clubroot disease.     

Suggestions for Determination and Terminology of Pathotypes and Genes for Resistance in Cyst-forming Nematodes, especially Heterodera avenae1

A short review of differentiation into pathotypes is given. Use of the word ≪ pathotype ≫ is recommended when a very clear difference is established between virulence of nematode populations. Our present knowledge makes it possible to differentiate between 10 pathotypes of Heterodera avenae. It is suggested that the pathotypes are given numbers, and corresponding terms should be used for genes for resistance in plants, e.g. a gene Hal on the barley chromosome gives resistance to nematode pathotypes 11,21,31,41 etc. Some proposals for improvement of pathotype identification are given.     

Intraspecific Relationship of Plasmopara halstedii Isolates Differing in Pathogenicity and Geographic Origin Based on ITS Sequence Data

Sequence parts of the internal transcribed spacer (ITS) region of nuclear ribosomal DNA were analysed to screen for the intraspecific variability of a non-coding genomic region in 15 Plasmopara halstedii populations of different pathotype and geographic origin. Samples revealed uniformity in a ca. 790 Bp fragment comprising of the ITS-1, 5.8S and front parts of the ITS-2. In contrast, clear differences were found in a ca. 810 Bp fragment of the ITS-2 thus allowing differentiation between populations of pathotype 100, 310 and 330 and a group of populations representing pathotypes 700, 701, 703, 710 and 730. Samples of pathotypes 700 to730 originated from Slovakia, France, and Germany, but were uniform in both ITS sequence parts, thus indicating very recent origin of these highly aggressive physiological races. The potential use of ITS sequences for pathotype differentiation and phylogenetic studies in P. halstedii is discussed.     

Relation between pathogenicity and genetic variation within <Emphasis Type="Italic">Plasmodiophora brassicae</Emphasis>

The relation between diversity of pathogenicity on clubroot-resistant (CR) cultivars of Chinese cabbage (Brassica rapa subsp. pekinensis) bred in Japan and DNA polymorphisms in 17 populations of Plasmodiophora brassicae from cruciferous plants was examined by inoculation tests and random amplified polymorphic DNA (RAPD) analysis using 18 arbitrary primers. Four pathotypes (A–D) were identified after inoculation of six CR cultivars of Chinese cabbage in the 17 populations from cruciferous crops. A relatively high level of genetic diversity was also detected among these populations in the RAPD analysis. Although the four pathotypes could not be clearly differentiated using the RAPD data, most populations of three pathotypes had a consistent location on the dendrogram. All pathotype B (virulent on five cultivars except Utage 70) and D (avirulent on all cultivars) populations, which were common in incompatible interactions with cv. Utage 70, were located in a single subcluster. All five pathotype C populations (virulent only on cv. Utage 70) except for one population grouped in another single subcluster. Because four pathotype A populations (virulent on all six cultivars, races 4 and 9) fell in different subclusters, the populations may be genetically polyphyletic. Populations from cruciferous weed Cardamine flexuosa differed remarkably from those from cruciferous crops in pathogenicity on common cultivars of Chinese cabbage and turnip and C. flexuosa, but they grouped in a single cluster with all race 9 populations from crops. Race 9 populations from crops may thus be closely related to populations from the weed rather than to races 1 and 4 from crops.