Pathogenicity of the Rhynchosporium secalis population in the Western Cape province of South Africa

The virulence spectra of 50 Rhynchosporium secalis isolates from a population in the Western Cape province of South Africa were determined, and 21 races were detected when evaluated against 17 differential cultivars. The virulence spectrum of the R. secalis population shows considerable variation, and carries unnecessary virulence genes which is quite unexpected, since chiefly susceptible barley cultivars are grown in the south Western Cape. The two most prevalent races, namely races 4 and 7 had three and four virulence genes respectively. Both race 4 and 7 were virulent on the most susceptible cultivars, West China, Steudelli, C.I.8618 and C.I.2226. Considering the resistance genes reported for the cultivars Atlas 46, Turk, and C.I.3515 which showed no susceptible cultivar-pathogen interaction, it would appear that the Rh-Rh3-Rh4 complex is primarily involved in conferring resistance to the local R. secalis isolates This revised version was published online in July 2006 with corrections to the Cover Date.     

Resistance of nineteen cultivars of subterranean clover to four races of Phytophthora clandestina

Summary The resistance of 19 cultivars of subterranean clover was screened against 4 races of P. clandestina by mycelial inoculation of roots of 10-day-old seedlings growing in water agar and by growing seedlings in pasteurised potting mix containing infested vermiculite in controlled conditions. The cultivars showed differential resistance (vertical resistance) to races of the pathogen and can be divided into 4 resistance groups. Cultivars Clare, Esperance, Green Range, Junee, Mount Barker, Rosedale, Woogenellup and Yarloop were susceptible to all races. Cultivars Bacchus Marsh, Denmark, Enfield, Gosse, Goulburn, Karridale, Larisa, Leura and Trikkala were susceptible to races 1 and 3, but resistant to races 0 and 2. Cultivar Meteora was susceptible to races 2 and 3, but resistant to races 0 and 1. Cultivar Seaton Park (LF) was resistant to all races. Cultivars also varied in their race-non-specific (horizontal) resistance: cultivars that were susceptible to particular races usually varied in their degree of susceptibility to those races. In particular, Junee was more resistant to all four races than the other cultivars within its group. Similarly, cultivars Gosse, Larisa, Denmark and Trikkala were more resistant to races 1 and 3 than the other cultivars in their group. Races of the pathogen varied in their aggressiveness as well as in their virulence, as shown by the variation in aggressiveness of different isolates of race 0.     

Evaluation of resistance sources and inheritance of resistance in kidney bean to Indian virulences of Colletotrichum lindemuthianum

Summary Forty nine common bean lines comprising of exotic accessions and locally grown cultivars evaluated against Colletotrichum lindemuthianum exhibited differential resistance to its races in Himachal Pradesh, a north-western Himalayan state of India. Some exotic accessions like G 2333, Cornell 49242, PI 207262, Mexique 222, TO, Perry Marrow, Kaboon and Widusa were resistant to more than five Indian races, whereas two Indian accessions KRC-5 and Hans showed resistance to six and four races, respectively. However, nine accessions KRC-8, KR-40, KR-43, KR-81, KR-62-2, KR-90, KR-142, KR-148, and KR-216 were resistant to three races. Race specific resistance has been observed in different bean cultivars. Studies on inheritance of resistance in exotic accession G 2333 and Indian accession, KRC-5 showed that two independent dominant genes conferred resistance in G 2333 to race 3 and 515 and a single dominant gene controlled resistance in KRC-5 to race 775, indicating resistance from these sources is easily transferable to the locally adapted susceptible cultivars.     

Characterization of resistance of cassava genotypes to bacterial blight by evaluation of leaf and systemic symptoms in relation to yield in different ecozones

Twenty-two improved and local cassava genotypes were evaluated for their bacterial blight symptom types in reaction to infection by Xanthomonas axonopodis pv. manihotis under field conditions in the forest, forest savanna transition and wet savanna zones of Togo. High genotype × environment interactions in development of each symptom type were observed. Combining data on environments and genotypes, spot, blight and wilt symptoms were positively correlated. Analysing genotype reactions across environments, indications for independent mechanisms of resistance on leaf and stem level, varying by genotype, were found. Genotypes Main27 with resistance to spot and blight symptoms and TMS4(2)1425 with resistance to wilt symptoms are recommended to breeders to introgress their resistance characteristics. Significant negative correlations were generally observed between blight and wilt symptom development and root yield across ecozones, with blight being more important under lower, and wilt under higher inoculum pressure. Genotypes TMS30572, CVTM4, TMS92/0429 and TMS91/02316 showed low spot, blight and wilt symptoms combined with high root yield across ecozones.     

Fusarium wilt of chickpea: physiological specialization, genetics of resistance and resistance gene tagging

Chickpea wilt caused by Fusarium oxysporum f. sp. ciceris is one of the major yield limiting factors in chickpea. The disease causes 10–90% yield losses annually in chickpea. Eight physiological races of the pathogen (0, 1A, 1B/C, 2, 3, 4, 5 and 6) are reported so far whereas additional races are suspected from India. The distribution pattern of these races in different parts of the world indicates regional specificity for their occurrence leading to the perception that F. oxysporum f. sp. ciceris evolved independently in different regions. Pathogen isolates also exhibit differences in disease symptoms. Races 0 and 1B/C cause yellowing syndrome whereas 1A, 2, 3, 4, 5 and 6 lead to wilting syndrome. Genetics of resistance to two races (1B/C and 6) is yet to be determined, however, for other races resistance is governed either by monogenes or oligogenes. The individual genes of oligogenic resistance mechanism delay onset of disease symptoms, a phenomenon called as late wilting. Slow wilting, i.e., slow development of disease after onset of disease symptoms also occurs in reaction to pathogen; however, its genetics are not known. Mapping of wilt resistance genes in chickpea is difficult because of minimal polymorphism; however, it has been facilitated to great extent by the development of sequence tagged microsatellite site (STMS) markers that have revealed significant interspecific and intraspecific polymorphism. Markers linked to six genes governing resistance to six races (0, 1A, 2, 3, 4 and 5) of the pathogen have been identified and their position on chickpea linkage maps elucidated. These genes lie in two separate clusters on two different chickpea linkage groups. While the gene for resistance to race 0 is situated on LG 5 of Winter et al. (Theoretical and Applied Genetics 101:1155–1163, 2000) those governing resistance to races 1A, 2, 3, 4 and 5 spanned a region of 8.2 cM on LG 2. The cluster of five resistance genes was further subdivided into two sub clusters of 2.8 cM and 2.0 cM, respectively. Map-based cloning can be used to isolate the six genes mapped so far; however, the region containing these genes needs additional markers to facilitate their isolation. Cloning of wilt resistance genes is desirable to study their evolution, mechanisms of resistance and their exploitation in wilt resistance breeding and wilt management.     

Genetic diversity of Phaeoisariopsis griseola in the State of Minas Gerais, Brazil

Due to the importance of common bean angular leaf spotin the state of Minas Gerais-Brazil and to the greatvariability of the pathogen, Phaeoisariopsisgriseola, monitoring races becomes an important toolfor breeding programs aiming at genetic resistance.The pathogenic variability of 30 isolates of the P. griseola, collected from various locations in thestate of Minas Gerais, was studied using the followingcommon bean differential series (Don Timóteo,Bolón Bayo, Montcalm, G 5686, Amendoin, G 11796,BAT 332, PAN 72, Cornell 49-242, México 54, Florde Mayo and G 2858). The first trifoliate leaf wasinoculated with a 2 × 10⁴ conidia/mL. Plants weremaintained at 20–22 ^°C and 95% relativehumidity for 48 hours. Symptom evaluation wasperformed 15 days after inoculation. Thirteen raceswere identified demonstrating the wide geneticvariability of the pathogen in the state of MinasGerais. Race 63.63 was the most virulent, whereas race63.23 was the most frequent (10 of 30 isolates), beingwidely distributed among the regions studied. Thevirulence phenotype indicated that the races studiedbelonged to the Mesoamerican group, which wasconfirmed when the 30 isolates were compared to Andeanand Mesoamerican standards using RAPD markers.     

Resistance to bacterial blight (Pseudomonas syringae pv. pisi) in Spanish pea (Pisum sativum) landraces

Pea bacterial blight (Pseudomonas syringae pv. pisi) has long been known to be present in pea growing areas of Spain and to cause serious crop losses, although there is no published record of its occurrence. A collection of 16 isolates from a winter pea trial in Valladolid in 1991 which were shown in this study to be P.s. pv. pisi races 4 and 6 would appear to be the first published record of the disease in Spain. This occurrence of races 4 and 6 is the same as reported for winter-sown peas in the South of France. Ten Pisum sativum landraces from different geographical areas of Spain and considered to be representative of the traditional pea crop, were tested for resistance to seven races of P.s. pv. pisi. Seedlings of each landrace were stem inoculated with the type strain of each race in a glasshouse. Resistance exhibited by the different landraces mainly conformed to those previously described in pea cultivars indicating various combinations of the main resistance genes: R3, R2+4, R3+4 and R2+3+4. R3 was the most frequent R gene, being present in all landraces. R4 was present in four and R2 in three of the landraces tested. Variation for resistance within landraces was limited except for landrace accessions ZP-0102, ZP-0109 and ZP-0137 which also showed variation for morphological traits. The resistance responses of landrace ZP-0109 were difficult to interpret, but suggested a genetic mixture with some evidence of less well documented R genes, R5 and/or R6, and possibly some unknown resistance to race 6. This revised version was published online in July 2006 with corrections to the Cover Date.     

Race differentiation and search for sources of resistance to Rhynchosporium secalis in barley in Italy

Summary Barley in Italy has recently been seriously affected by Rhynchosporium secalis. The pathogenic variation of the fungus was studied and 17 races were differentiated on 13 barley cultivars carrying most of the currently known genes for resistance. RC 1, the most virulent and most frequent race, was virulent on 10 out of the 13 differentials and the remaining races proved to be less virulent variants of RC 1. Atlas (C.I. 4118), Atlas 46 (C.I. 7323) and Osiris (C.I. 1622) were the only three differentials resistant to all the analyzed single-spore isolates.Differential cultivars previously assumed to have identical resistance factors did not react in the same way to all the Italian races, thereby revealing either undisclosed differences in the genes described or the presence of additional unidentified ones.Our findings were compared with previous data about virulence of scald populations from different countries, on the basis of tests with common differentials: fundamental differences were found between the Italian population and those of other countries with regard to virulence patterns.The susceptible reactions to race RC 1 of most barley cultivars grown in Italy indicate the urgent need for resistance genes to be incorporated in the cultivated material. Seventy-one barley accessions, known as sources of resistance in different parts of the world, were screened for their behaviour to races RC 1 and RC 13. Twenty-two appeared resistant to both of them.     

The effect of pre- and post-inoculation temperature on resistance in certain cultivars of poplar to races of Melampsora larici-populina kleb.

Summary Four cultivars of Populus spp., compatible to varying degrees with four races of M. larici-populina Kleb., were raised in a controlled environment on a high (28°/20°C, day/night) and low (20°/10°C) temperature regime. Leaf discs cut from the plants were inoculated separately with four individual races of M. laricipopulina and subsequently incubated at either low (20°C) or high (25°C) temperature for 14 days when disease development on the discs was assessed using three parameters (Incubation period to flecking, uredia per leaf disc and uredospores per mm²). The degree of resistance in all cultivar/race combinations was high on cultivars cultured at a high temperature regime compared to those cultured on a low temperature regime. Analysis of variance demonstrated that the major components: pre-inoculation temperature regime, post-inoculation temperature regime, race and cultivar, and most second and third order interactions between these were highly significant (P<0.001) for most disease parameters. The variance of the temperature components and all interactions involving these were usually higher than those for the cultivar and race components and those interactions lacking temperature components.These results emphasize the importance of the temperature regime at which plants are raised and the temperature of incubation, following the inoculation in determining the relative degree of resistance of these cultivars of poplar to races of M. larici-populina. The implications of these results in the epidemiology of leaf rust and the stability of the host-parasite relationship are discussed.     

Genetics of host reactions to three races of the bacterial pustule pathogen in cowpea

Summary Inheritance of the brown hypersensitive resistant (BHR), non-hypersensitive resistant (R) and susceptible (S) host reactions produced by three races of the bacterial pustule pathogen (Xanthomonas campestris pv. vignaeunguiculatae) was studied in 45 F₁, F₂ and testcross progenies, using the infiltration inoculation method BHR reaction was dominant over R and S reactions, and R was recessive to S reaction. Two genes appeared to be involved in BHR reaction; one governing BHR reaction to race 1 and the other to races 1 and 2. Both were ineffective against race 3. R reaction, effective against all the races, appeared to be controlled by one, two or three recessive genes. One cowpea line had one BHR gene and two duplicate recessive R genes. Reaction expression in the segregants was clear and as expected with races 2 and 3 but was modified with race 1, possibly due to interactions between dominant or recessive alleles of the BHR genes and the homozygous recessive allele of the R genes. Gene symbols Bp-1 and Bp-2 are proposed for the BHR genes and bp-3, bp-4 and bp-5 for the recessive R genes. The genes present in each of the differential cowpea line are suggested.Contribution from the International Institute of Tropical Agriculture, Ibadan, Nigeria and Crop Development Division, Ministry of Agriculture, P.O. Box 9071, Dar es Salaam Tanzania.     

黄淮大豆主产区大豆胞囊线虫生理小种分布调查

大豆胞囊线虫(SCN)在黄淮地区普遍发生,调查小种分布情况,确定优势小种对抗病育种有重要意义。2012-2015年,取样调查黄淮地区6个省份土样,利用Riggs模式鉴定生理小种,绘制黄淮地区SCN生理小种分布图,并与文献报道结果对比,探讨黄淮地区SCN生理小种类型及其分布规律。结果表明,该病害在黄淮大豆主产区均有分布,在采集受SCN感染的322份土样中,112份被鉴定出生理小种类型,包括1号、2号、3号、4号、5号、6号和11号小种。其中,57份为2号小种,占样本总体的50.9%;26份土样为5号小种,占23.2%;11份土样为4号小种,占9.8%,1号、3号、6号和11号小种分别占总体的4.5%、5.4%、4.5%和1.8%。依据不同生理小种在各省发生频率由高到低的顺序,河南分布5号、2号、3号、11号小种;河北分布2号、5号、6号、3号、4号小种;安徽分布2号、5号、6号、3号小种;山西分布2号、4号、5号、1号、3号、11号小种;山东分布2号、3号、5号、1号、6号小种;江苏分布2号、5号、1号小种。以上结果表明,2号小种是目前黄淮海地区的优势小种,其次是5号小种,致病力最强的4号小种主要分布在山西省。在黄淮海地区,抗线虫育种目标应以抗2号生理小种为主,兼抗5号小种,部分地区应以兼抗2号和4号小种为主。在黄淮地区3号、6号和11号小种是新发现的小种。与2001-2003年调查结果比较,黄淮海地区大豆胞囊线虫生理小种组成及分布有一定的改变。     

The potential for breeding an improved lucerne-rhizobium symbiosis. 1. Assessment of genetic variation

Summary Six cultivars of lucerne (Medicago sativa L.) were grown in all possible combinations with eight strains of Rhizobium meliloti in order to assess genetic variation in symbiotic nitrogen fixation. Host genotype and Rhizobium genotype effects on nitrogen fixation were respectively 4.8% and 21.0% of total phenotypic variance. Genetic variation due to host cultivar × Rhizobium strain interactions accounted for a further 6.0% of phenotypic variance. The results indicated low heritability of general symbiotic effectiveness in the host with interaction effects being large enough to suggest that plant performance may be unpredictable with populations of R. meliloti in field soils. Joint regression analysis showed that about 50% of the interactive variation could be explained by generalized differences in the sensitivity of cultivars to alterations in the genotype of Rhizobium strains. One cultivar, Siriver, was relatively insentitive to changes in the Rhizobium genotype whilst still maintaining high average yield. The implications of the results for lucerne breeding are discussed.     

Inheritance of resistance to the chlorosis component of tan spot of wheat caused by Pyrenophora tritici-repentis, races 1 and 3

The fungus Pyrenophora tritici-repentis, causal agent of tan spot of wheat, produces two phenotypically distinct symptoms, tan necrosis and extensive chlorosis. The inheritance of resistance to chlorosis induced by P. tritici-repentis races 1 and 3 was studied in crosses between common wheat resistant genotypes Erik, Hadden, Red Chief, Glenlea, and 86ISMN 2137 and susceptible genotype 6B-365. Plants were inoculated under controlled environmental conditions at the two-leaf stage and disease rating was based on presence or absence of chlorosis. In all the resistant × susceptible crosses, F₁ plants were resistant and the segregation of the F₂ generation and F₃ families indicated that a single dominant gene controlled resistance. Lack of segregation in a partial diallel series of crosses among the resistant genotypes tested with race 3␣indicated that the resistant genotypes possessed␣the same resistance gene. This resistance gene was effective against chlorosis induced by P.␣tritici-repentis races 1 and 3.     

Influence of plant age,disease incubation time,and host plant resistance on Verticillium wilt symptom expression in lucerne (Medicago sativa)

Summary The most economical means to control Verticillium wilt in lucerne, caused by Verticillium albo-atrum, is through resistant cultivars. Most Verticillium wilt resistance evaluations use some form of root-cut-soak inoculation procedure. In Trial 1, three lucerne cultivars were inoculated at eight plant ages and plant symptoms were scored at ten successive weekly intervals. Trial 2 consisted of two cultivars inoculated at ten plant ages and symptoms were evaluated at eight weekly intervals. In an experiment to study the genetics of resistance to this pathogen, four plants showing no visual symptoms were selected from each of four germplasms. The selected populations were crossed in diallel fashion. The population crosses plus the parents were evaluated for resistance to V. albo-atrum. Results from Trial 1 and Trial 2 showed disease severity increased with increasing inoculation age and with increasing scoring age. Cultivar symptom expression interacted with inoculation age, but not with scoring age. The diallel analysis supported resistance to Verticillium wilt being conferred by additive gene action, as heterosis was not detected and general combining ability mean square was 3.75 times larger than the specific combining ability mean square.     

Identification of pathogenic races ofPhytophthora fragariae Hickman in Germany

Summary The fungusPhytophthora fragariae Hickman is the causal agent of red stele disease in strawberry (Fragaria × ananassa), this being a major disease in many areas with cool and moist conditions. Success of resistance breeding can be nullified by the appearence of specific races of the fungus capable of overcoming the introduced resistance. In some countries (USA, UK, Japan and Canada) races were identified by using a differential set of strawberry cultivars. The absence of an international standard differential set and the use of different test methods make a comparison of the identified races difficult or even lead to contradictory results for one variety/race combination. The aim of this study was to obtain information about the spectrum of pathogenic races in Germany as a basic contribution to research on resistance breeding against the fungus. The susceptibility of different strawberry cultivars to German isolates ofP. fragariae was evaluated. The inoculation was done by modifying a method described by Milholland et al. (1989). Rootedin vitro plants, four weeks after transferring them to the soil, were used for the investigation. Inoculation was done with a zoospore suspension of defined concentration. Up to now three German isolates, G-1, G-2, and G-3, can be separated by their ability to infect and produce oospores in the roots of the strawberry cultivars ‘Senga Sengana’, either ‘Saladin’, ‘Redgauntlet’ or ‘Climax’ andFragaria chiloensis clone ‘YaquinaB’. These results confirm the existence of pathogenic races ofP. fragariae in Germany and should be taken into consideration for resistance breeding against the fungus.     

Variations in pathogenicity patterns ofMycosphaerella graminicola withinTriticum spp. in Israel

Summary Pathogenicity patterns of 42Mycosphaerella graminicola (Septoria tritici) isolates secured from the major wheat growing regions in Israel, were assessed on seedlings of 16 bread and durum wheats. The spring bread wheat cultivar Titan (CI12615), the winter bread wheats Bezostaya 1, Kavkaz, NE7060 and the durum wheats Nursit 163 and Zenati Bouteille, all exhibited a high level of resistance. Significant cultivar × isolate interactions were recorded for isolates from the different regions in relation to the response of the 16 cultivars organized in 6 reponse classes to the test isolate ISR8036. Number of genes for resistance in the 16 cultivars was estimated, based on the assumption of a gene-for-gene relationship. Twelve complementary genes were hypothesized in the 42M. graminicola isolates × 16 wheat cultivars matrix. There was considerable variation in virulence frequencies between regions and between locations within the same region. The overall frequency of Bezostaya-Kavkaz virulence was low throughout the country (5–7%). The frequency of designated hypothesized virulence genes VST OLAF, VST COLOTANA, VST IAS 20 in Israel, was high in most locations. The reported analytical approach enables the identification of virulence hot-spots and assists in designing breeding for resistance strategies.     

Inheritance of resistance to two races of sunflower downy mildew (Plasmopara halstedii) in two Helianthus annuus L. lines

Sunflower downy mildew caused by Plasmopara halstedii is an important disease of sunflower capable of causing losses of more than 80% of production. Races 100, 300, 310, 330, 710, 703, 730 and770 of the fungus have been identified in Spain. Race 703, of high virulence, has been identified frequently in the northeast, while race 310 seems to occur over the south, the main sunflower growing region of the country. Oil sunflower lines RHA-274 and DM4 were studied for their resistance to races 310(RHA-274 and DM4) and 703 (DM4). In each cross, only one plant of the resistant parent was crossed to the inbred susceptible line HA-89 (or cmsHA-89).Plants from F₂ and backcross(BC₁F₁ to susceptible parent)generations were evaluated for fungal sporulation on true leaves and/or cotyledons. The resistant-to-susceptible ratios obtained in the F₂ and BC₁F₁ progenies from the crosses cmsHA-89 × RHA-274 and HA-89 × DM4suggested that one major gene in each line is responsible for resistance to race 703.The segregations of the progenies of the cross HA-89 × DM4 inoculated with race 703also fitted the ratios 1:1 and 3:1 (for BC₁F₁ and F₂, respectively)corresponding to control of resistance by a single dominant gene. In RHA-274, the gene for resistance to race 310 was designated Pl ₉, whereas Pl _v is tentatively proposed to designate the gene in DM4 responsible for resistance to races310 and 703. This revised version was published online in August 2006 with corrections to the Cover Date.     

Symbiotic variability in Vicia faba. 3. Genetic effects of host plant,rhizobium strain and of host × strain interaction

Summary Six Vicia faba populations were grown in all possible combinations with six Rhizobium leguminosarum strains in order to estimate the relative importance of the three genetic components of symbiotic variability. Additive genetic effects of host genotype and rhizobium genotype accounted for only 8.9% and 11.8% of the total phenotypic variation. Non-additive variation attriabutable to specific host genotype × rhizobium genotype interactions was by far the largest component of variation and accounted for 73.8% of phenotypic differences. Therefore the greatest improvement in symbiotic nitrogen fixation is likely to arise from simultaneous selection of both symbionts.     

Race-specific effects in major genic and polygenic resistance of barley to barley leaf rust in the field: identification and distinction

Summary Six partially resistant spring barley cultivars were exposed to four barley leaf rust (Puccinia hordei) races in the field and in the greenhouse. The 24 cultivar-race combinations were tested in field plots of 1.5×1.5 m² in two replications over two years. To reduce the interplot exchange of urediospores each plot was surrounded by winter rye.The level of barley leaf rust varied among cultivars, races and years. In both years the variance for cultivar-race interactions was highly significant and originating largely from the cultivar-race combinations Berac-22. Armelle-22, Armelle-A and Tyra-A. The Berac-22 interaction was towards higher, the other three interactions towards a lower level of barley leaf rust. The reduced rust levels of these three combinations were not due to interactions between the partial resistance of these cultivars and the aggressiveness of the races but to major genes for hypersensitivity not effective to the races 1-2-1 and F, common in Western Europe, but effective against the rare races 22 and A. This was revealed in the greenhouse experiments where all combinations had a susceptible infection type except Armelle-22, Armelle-A and Tyra-A, which showed low infection types in both the seedling and adult plant stages. The urediosori present in the field plots of these three combinations apparently arose from spores derived from other plots; this interplot interchange suggesting partial resistance.The interaction of Berac with race 22 truly was a small race-specific effect within the polygenic, partial resistance of barley to barley leaf rust like the one reported before between Julia and race 18.     

Race Specific Powdery Mildew Resistance in 31 Northwest European Wheat Cultivars

Race specific powdery mildew resistance in 23 winter wheat cultivars, eight spring wheat cultivars, and 14 lines/cultivars possessing known powdery mildew resistance genes, has been studied by analyzing host/pathogen interactions. The cultivars were tested as intact seedlings, and as detached primary leaf segments on water agar; both methods revealed reproducible and concordant results. The 45 cultivars/lines were divided into 24 resistance spectra according to the patterns of reaction to the powdery mildew isolates used. Of the 31 cultivars investigated, eight did not possess any of the resistance genes detected, and the remaining 23 were divided in 16 resistance spectra. The race specific resistance of nine cultivars was conferred by the single resistance genes Pm2, Pm4b, Pm5/Ml-i: or Pm6, while the race specific resistance of 14 cultivars was conferred by 2, 3, 4 or 5 genes in combination.