Pathogenic and molecular diversity in highly virulent populations of the parasitic weed Orobanche cumana (sunflower broomrape) from Europe

The parasitic weed Orobanche cumana (sunflower broomrape) constrains sunflower production in eastern and southern Europe and in the Middle East. Although genetic resistance is the most effective control method, new parasite races evolve overcoming sunflower resistance. In this work, highly virulent populations of O. cumana were analysed for pathogenicity and genetic diversity. The virulence of 11 populations from Hungary, Romania, Spain and Turkey was assessed and compared after infection of sunflower inbred lines to differentiate races of the parasite under glasshouse conditions. Molecular diversity among and within 27 parasite populations was studied by RAPD‐PCR, UPGMA and amova analyses. Highly virulent race F was identified in Hungary, Spain and Turkey. The most virulent race (G) was also found in Turkey. The molecular analysis among highly virulent populations of O. cumana identified four molecular clusters, respectively, grouping populations from Central Spain, Hungary, South Spain and Turkey. The genetic homogeneity within parasite populations was confirmed, since no molecular divergences were found within them. This work constitutes the first geographical study of O. cumana together with pathogenicity and molecular traits inherent to each geographical group, and provides useful information for possible phylogenetic analyses of O. cumana. In addition, molecular markers associated with geographical origin could be developed and used as diagnostic tools to track new broomrape introductions into areas free of virulent races where they might represent a threat to sunflower production.     

Indigenous highly virulent accessions of the sunflower root parasitic weed Orobanche cumana

Orobanche cumana (broomrape) is a severe constraint to sunflower production in southern and eastern Europe and the Middle East. Races A to E of this parasitic weed controlled by genes Or₁ to Or₅ have been described. A study of 38 seed accessions of O. cumana collected from different locations in Spain between 1983 and 2003 investigated the effect of long‐term storage in the laboratory on germination and infectivity and assessed their virulence on a number of sunflower cultivars. Only 18 infected the susceptible cultivar B117. Infectivity was maintained for up to 17 years of storage, but with a greatly decreased vigour as compared with that of recently collected seed. The 12 oldest viable accessions overcame the resistance of the gene Or₅ (in resistant line NR5). Seven out of them, in particular those collected in 1988 and 1989, were identified as race F. Three accessions were identified as race E allegedly holding components of higher virulence. Our results show evidence of the occurrence of race F prior to the use of sunflower hybrids resistant to race E, suggesting the former as indigenous to the country. This finding suggests the necessity of a continuous breeding of sunflower for resistance to O. cumana. The effectiveness and sustainability of genetic resistance must rely on the knowledge of the diverse virulence characteristics of O. cumana accessions.     

The role of peroxidase in the resistance of sunflower against Orobanche cumana in Russia

We present the results of a histochemical study comparing seedlings of races C and D of Orobanche cumana Wallr. (syn. O. cernua Loefl.) attacking sunflower (Helianthus annuus L.) in southern Russia. Three groups of O. cumana seedlings were distinguished according to the peroxidase content of the cells in the radicles: (1) those with neither extracellular nor intracellubr peroxidase and whose radicles have a smooth apex (these were classified as non-infective): (2) those with a high peroxidase content of the nuclei and the cytoplasm layer adjacent to the cell wall, as well as excretion of peroxidase from the apex of the radicles: (3) those with a similarly high peroxidase activity in the parasite cells, but without extracellular excretion. The apices of the radicles of the last two groups are swollen. It is suggested that these belong to O. cumana races C and D respectively. The extracellular peroxidase in O. cumana race C reacts with phenolic compounds, which are lignin precursors of the host, resulting in host resistance due to the formation of lignin layers in sunflowers possessing the Or3 gene for resistance. The absence of extracellular peroxidase in O. cumana race D prevents lignin formation and enables the parasite to attach to the host vascular system. Comparison of these data with the information on the earlier O. cumana races A and B, and older sunflower cultivars, points to a crucial role of peroxidase in the process of breeding new sunflower cultivars and the evolution of new O. cumana races.     

Pathogenic diversity within field populations of Orobanche cumana and different reactions on sunflower genotypes

Different races of the parasitic Orobanche cumana (sunflower broomrape) have been reported in Spain, race F being the most virulent. Full resistance in sunflower to races A–E is achieved with each of the single major genes Or₁ to Or₅ respectively. However, parasitised hybrids allegedly resistant to race F were observed in early 2002. The purpose of this study was to verify broomrape incidences (BI) on resistant sunflower genotypes, to assess the mixture of races within field populations and to test for partial resistance to race F in the sunflower hybrids showing a low degree of attack (DA) by the weed. Tests were conducted under field conditions in two locations of southern Spain. While no significant differences were found for yield and BI between locations, the DA on the cultivars depended on the location. With high infection levels and significantly lower yield in susceptible controls, marked differences in BI and DA were found within resistant cultivars, but all of them showed similar crop yield. When artificially inoculated with several populations of race F, line P96 and mainly line L86, were consistently slightly infected, suggesting they were inbred lines responsible for horizontal resistance in infested fields. L86 was extremely susceptible to race E populations, which is unusual as sunflower resistance to one race provided resistance to all the previously described races of O. cumana. No different virulences were detected within two groups of subpopulations (races E and F) inoculated onto resistant sunflower genotypes. However, race F subpopulations showed significant differences in aggressiveness, which seems to be related to horizontal (multigenic) resistance of the crop to the parasitic weed.     

Identification,characterisation and discriminatory power of microsatellite markers in the parasitic weed Orobanche cumana

Orobanche cumana is an obligate root parasite of sunflower. It represents a major agricultural problem in many countries of southern and eastern Europe. Information on O. cumana population genetics, structure and dynamics is scarce, particularly due to the lack of suitable molecular markers for such studies. The objective of this study was to identify and characterise simple sequence repeat (SSR) markers for O. cumana. Four thousand two hundred SSR‐containing candidate sequences were obtained from O. cumana using next‐generation sequencing, from which 298 SSR primer pairs were designed and 217 of them used for validation. Seventy nine SSR primers produced reproducible, high quality amplicons of the expected size that were polymorphic among 18 O. cumana populations from different geographical locations and hosts (sunflower, wild hosts from the Compositae family). The number of alleles per locus ranged from 2 to 10, with an average polymorphism information content value of 0.37. The O. cumana SSR markers were highly transferable to the closely related species Orobanche cernua. SSR markers showed high resolving power; UPGMA cluster analysis allowed proper classification of Orobanche spp. samples into species (O. cumana and O. cernua), geographical origin and host. The functional SSR markers reported in this study constitute a valuable tool for genetic analyses in O. cumana and related species and will contribute insights into the biology and genetics of this parasitic weed.     

Genetic diversity of Orobanche cumana populations from Spain assessed using SSR markers

Orobanche cumana (sunflower broomrape) is found in Spain as an allochthonous species parasitising exclusively sunflower. For many years, it was distributed in the Guadalquivir Valley and Cuenca province, but in recent years, it has spread to new areas. The objective of this research was to study genetic diversity of O. cumana populations from Spain using robust co‐dominant molecular markers. Cluster analysis on a set of 50 populations using 15 microsatellite markers revealed the existence of two distant gene pools, one in Cuenca province and another one in the Guadalquivir Valley. Within each gene pool, both inter‐ and intrapopulation variability were extremely low. This population structure probably reflects a founder effect, with the two genetically distant gene pools deriving from separate introduction events. Different races occurred within the same gene pool, suggesting that current races might have evolved through mutation from a common genetic background. Most of the populations from new areas were identical to the populations from the Guadalquivir Valley. Only a few populations showed larger intrapopulation variation. In these cases, our results suggested the co‐existence of both gene pools within the same population, as well as the occurrence of genetic recombination between them. Genetic recombination between distant gene pools is an important mechanism for creating new variation, which might also have an effect on race evolution. These results will contribute to the establishment of improved crop breeding and management strategies for O. cumana control.     

Inheritance of virulence in Fusarium circinatum,the cause of pitch canker in pines

Wildtype strains of Fusarium circinatum, the causal agent of pitch canker, were crossed to obtain an F₁ generation. Progeny of this cross were tested for virulence by inoculating Pinus radiata seedlings, and were found to induce a wide range of lesion lengths. Two strains from the F₁ generation that induced long lesions (= high virulence) were used as parents to produce an F₂ generation, followed by a second round of selection for high virulence to obtain an F₃ generation. Mean lesion lengths were not significantly different between the three generations (P ≥ 0.196). A parallel set of crosses was performed to select for low virulence by using progeny in the F₁ and F₂ generations that induced short lesions as parents for F₂ and F₃ generations, respectively. In this case, both rounds of selection resulted in a significant reduction in mean lesion length, from 33.8 ± 0.8 mm in the F₁ generation, to 19.7 ± 0.7 and 12.9 ± 0.7 mm in the F₂ and F₃ generations, respectively. Thus it is apparent that F. circinatum retains the genetic capacity for avirulence to pines, which could reflect a lack of strong selection for virulence in nature. Progeny of a cross between high and low virulence parents manifested nearly continuous variation in lesion lengths, consistent with virulence being a quantitatively inherited trait. Based on this cross, broad‐sense heritability (H²) was determined to be 0.74, which suggests that virulence is under strong genetic control.     

Widespread distribution of Xanthomonas perforans and limited presence of X. gardneri in Brazil

Tomato bacterial spot is caused by Xanthomonas euvesicatoria, X. vesicatoria, X. perforans and X. gardneri. In order to determine the distribution, frequency of occurrence, and diversity of these species in the Brazilian commercial tomato fields, a survey was conducted between 2009 and 2012. In this period, 204 strains were obtained from 33 counties (22 with processing tomatoes and 11 with fresh‐market tomatoes). Pathogenicity tests, BOX‐PCR, PCR with species‐specific primers, and sequence analysis of the avirulence gene avrXv3 were performed in order to identify the strains at species and race level. Xanthomonas perforans predominated among the strains (92%) and was present in most counties. In addition, this species was prevalent in most areas of both fresh‐market tomatoes (63.6% of counties surveyed) and processing tomatoes (95.4% of counties surveyed). Fifteen strains (7.5%) were identified as X. gardneri, which was found mostly in fresh‐market fields located at regions with altitude higher than 900 m, and only one strain of X. euvesicatoria (0.5%) was found in a processing tomato field. High genetic diversity was observed within X. perforans, with 137 BOX‐PCR haplotypes. Race T3 prevailed (97.5%), but reported here for the first time is the occurrence of five strains identified as race T4 in fresh‐market fields in the state of São Paulo. The race T4 phenotype of these strains resulted from the presence of an 859 bp insertion in the avirulence gene avrXv3. This insertion is related to amino acid sequences of a transposase found in X. gardneri, and to amino acid sequences of X. campestris.     

Pathogenic variability of Plasmopara halstedii infecting sunflower in the Czech Republic

Plasmopara halstedii was isolated from diseased sunflowers collected from eight locations in the Czech Republic from 2007 to 2014. Races of the pathogen were determined based on 84 isolates collected during the study. In total, eight races of P. halstedii were detected using a set of nine sunflower differential lines. Races 700, 704, 705, 710, 714 and 715 were proven by soil drench inoculation, and two additional races (730 and 770) proposed by the previously applied leaf disc inoculation method. Race 700 was the most dominant in the Czech P. halstedii populations, with race 710 being the second most frequent. Races 704 and 714 were found over three seasons, while other races were recorded only in one growing season (race 730 in 2010, and the new races 705 and 715 in 2014). A comprehensive study was further conducted for isolates collected in 2013–14 using an extended differential set consisting of 15 sunflower lines. According to the latter methodology which marks races with five‐digit virulence codes, races 70060, 70471, 70571, 71060, 71461 and 71571 were recorded. The growing complexity of P. halstedii pathogenicity exhibited by the ability to infect higher numbers of differential genotypes and resulting in determination of the new pathogen races (virulence profiles) 70571, 71461 and 71571 is alarming. Although the limited number of isolates studied cannot characterize the entire pathogen diversity in the Czech Republic, the trend towards more diverse virulence in P. halstedii populations is clearly demonstrated by the new records of races 704, 705, 714 and 715, all capable of overcoming the resistance gene Pl₆.     

向日葵上一种列当的发生分布及防治

从1979～1980年调查研究明确了吉林省白城地区寄生在向日葵上的恶性杂草——列当的种名是Orobanche coerulescens Steph(本文暂称白城列当)而不是向日葵列当(O.cumana)。白城列当在全地区8个县均有分布,发生密度较大的地块1平方米均有300棵左右,寄生株率96%以上,1株向日葵上最多寄生146棵。其垂直分布主要在土壤耕层里,是寄生于向日葵根部,以5～15厘米耕层寄生率最高达93%。在向日葵植株周围的水平分布以距向日葵5～20厘米寄生率达68.1%。被害的向日葵植株,表现株矮、茎细、盘小,甚至全株枯死。籽实减产38.3～70.3%,籽仁含油率降低3.3～17.9%,严重影响向日葵的生产,应引起各方面重视。提出因地制宜地采取选育抗性品种,合理轮作及提早铲除等综合防治措施,以便尽早地消灭在始发阶段。     

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

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

Race 2 of Verticillium dahliae infecting tomato in Japan can be split into two races with differential pathogenicity on resistant rootstocks

Verticillium dahliae infecting tomato can be differentiated into races 1 and 2 based on differential pathogenicity on tomato cultivars carrying resistance gene Ve1. Although no commercial cultivars resistant to race 2 are available, race 2‐resistant rootstock cultivars Aibou and Ganbarune‐Karis have been bred in Japan. Nevertheless, the resistance of these rootstocks appears to be unstable in commercial tomato fields. Pathogenicity assays conducted under controlled conditions revealed that these rootstock cultivars are resistant to some isolates of race 2; this resistance is controlled by a single dominant locus, denoted by V2, based on segregation of resistance in F₂ populations from selfed rootstock cultivars. However, some other isolates of race 2 can overcome this resistance. Therefore it is proposed that the current race 2 of V. dahliae should be divided into two races, i.e. ‘race 2’ (nonpathogenic on Aibou) and ‘race 3’ (pathogenic on Aibou). The distribution of these races was surveyed in 70 commercial tomato fields in Hida, Gifu Prefecture, Japan. Race 3 was found in 45 fields, indicating that race 3 had already spread throughout the region. On the other hand, 25 fields had only race 2, and thus race 2‐resistant rootstocks would be effective for disease management in these fields. Races 2 and 3 could not be identified by genomic Southern hybridization probed with a telomere sequence, nor with previously reported race‐specific PCR assays. Elucidation of race‐determining mechanisms and development of methods for quick race identification should be made in future studies.     

An avirulence gene to rice cultivar K60 is located on the 1.6-Mb chromosome in <Emphasis Type="Italic">Magnaporthe oryzae</Emphasis> isolate 84R-62B

A Japanese differential rice cultivar K60 was tested with 114 F₁ cultures of Magnaporthe oryzae from a cross between isolates 84R-62B and Y93-245c-2. Segregation patterns of avirulence and virulence in the progeny suggested that avirulence on cv. K60 was controlled by a single gene derived from 84R-62B and tentatively named AvrK60. In the F₁ population, AvrK60 cosegregated with avirulence gene AvrPik on a small 1.6-Mb chromosome of 84R-62B and with the 1.6-Mb chromosome itself. Therefore, we suggest that, along with AvrPik, AvrK60 is also located on the 1.6-Mb chromosome of 84R-62B.     

Resistance to Pseudomonas syringae in a collection of pea germplasm under field and controlled conditions

A total of 242 Pisum accessions were screened for resistance to Pseudomonas syringae pv. pisi under controlled conditions. Resistance was found to all races, including race 6 and the recently described race 8. Fifty‐eight accessions were further tested for resistance to P. syringae pv. syringae under controlled conditions, with some highly resistant accessions identified. Finally, a set of 41 accessions were evaluated for resistance to P. syringae pv. pisi and pv. syringae under spring‐ and winter‐sowing field conditions. R2, R3 and R4 race‐specific resistance genes to P. syringae pv. pisi protected pea plants in the field. Resistance sources to race 6 identified under controlled conditions were ineffective in the field. Frost effects were also evaluated in relation to disease response. Results strongly suggest that frost tolerance is effective in lowering the disease effects caused by P. syringae pv. pisi and pv. syringae under frost‐stress conditions, even in the absence of disease resistance genes, although the highest degree of this protection is reached when frost tolerance and disease‐resistance genes are combined in the same genetic background.     

A new race of Fusarium oxysporum f. sp. lactucae of lettuce

Fusarium oxysporum f. sp. lactucae, the causal agent of fusarium wilt of lettuce (Lactuca sativa), occurs in most countries in which lettuce is grown and causes serious economic losses. Three races (1, 2 and 3) of the pathogen have previously been identified on the basis of their ability to cause disease on differential lettuce cultivars, as well as by means of molecular tools developed to characterize different races of this pathogen. Only race 1 has been detected in Europe so far. In this study, two isolates of F. oxysporum, obtained from lettuce plants grown in the Netherlands showing symptoms of wilt, have been characterized by combining the study of pathogenicity with differential cultivars of lettuce and molecular assays to determine whether the isolates are different from the known races of F. oxysporum f. sp. lactucae. This study reports the presence of F. oxysporum f. sp. lactucae for the first time in the Netherlands. The causal pathogen has been identified, using the IRAP‐SCAR technique, as a new race of F. oxysporum f. sp. lactucae. Specific primers have been designed to identify this new race.     

Relationship between Avirulence Genes of the Same Family in Rice Blast Fungus Magnaporthe grisea

A genetic cross between rice-field isolates of Magnaporthe grisea produced progeny segregating for avirulence/ virulence on six rice cultivars among nine race differentials, while on three other cultivars, Shin 2 (Pik-s), Aichi Asahi (Pia) and Ishikari Shiroke (Pii), parental and progeny isolates were all virulent. Based on segregation ratios in 115 progeny isolates, avirulence on Kanto 51 (Pik), Yashiro-mochi (Pita), Fukunishiki (Piz) and Toride 1 (Piz-t) is under monogenic control. On Tsuyuake (Pik-m) and Pi No. 4 (Pita-2), however, a disproportionate ratio in the segregation was observed, suggesting that avirulence on these two cultivars is controlled by two or more genes. Assuming that the avirulence gene AvrPik-m consists of at least two genes, AvrPik-m1 and AvrPik-m2, each of which functions in the whole gene AvrPik-m, and that one of AvrPik-m1 and AvrPik-m2 is AvrPik, we could account for the disproportion in the avirulence/virulence segregation of the progeny. This hypothesis would also be consistently applied for avirulence gene AvrPita-2. There seem to be two types of the avirulence genes : AvrPik-m, that is comprised of the tightly linked genes, AvrPik-ml (=AvrPik) and AvrPik-m2, and AvrPita-2, that is comprised of the loosely linked genes AvrPita-2A (=AvrPita) and AvrPita-2B. As one possible explanation of the rice resistant reaction to blast, multiple specificity was suggested for the first time for the blast fungus. On the contrary, the avirulence genes AvrPiz and AvrPiz-t were inherited independently, despite the corresponding genes for resistance (Piz and Piz-t) being located at the same locus. The cross of rice blast isolates (races 447 and 337) produced only 25 kinds of races in the progeny, although theoretically about 64 kinds of races should be produced if six avirulence genes segregated independently. Because no progeny are with AvrPik (or AvrPita) and without AvrPik-m (or AvrPita-2), the number of races theoretically should be 36 at most. A number of strains, such as races 377 and 737, with a single avirulence gene were obtained from this cross. These strains may be valuable for analysis of resistance genes in rice plant. Received 19 August 2002/ Accepted in revised form 11 November 2002     

The tomato Pto gene confers resistance to Pseudomonas floridensis,an emergent plant pathogen with just nine type III effectors

A newly discovered bacterial species, Pseudomonas floridensis, has emerged as a pathogen of tomato in Florida. This study compares the virulence and other attributes of P. floridensis to Pseudomonas syringae pv. tomato, which causes bacterial speck disease of tomato. Pseudomonas floridensis reached lower population levels in leaves of tomato as compared to the P. syringae pv. tomato strains DC3000 and NYT1. Analysis of the genome sequence of the P. floridensis type strain GEV388 revealed that it has just nine type III effectors including AvrPtoB_GEV388, which is 66% identical to AvrPtoB in DC3000. Five of these effectors have been previously reported to be members of a ‘minimal effector repertoire’ required for full DC3000 virulence on Nicotiana benthamiana; however, GEV388 grew poorly on leaves of this plant species compared to the DC3000 minimal effector strain. The tomato Pto gene recognizes AvrPtoB in race 0 P. syringae pv. tomato strains, thereby conferring resistance to bacterial speck disease. Pto was also found to confer resistance to P. floridensis, indicating this gene will be useful in the protection of tomato against this newly emerged pathogen.     

Stability and variability of virulence of Phytophthora infestans assessed in a ring test across European laboratories

Determining virulence towards race‐specific resistance genes is a prerequisite to understanding the response of pathogen populations to resistant cultivars, and therefore to assess the durability of these resistance genes and the performance of resistance management strategies. In Phytophthora infestans, virulence testing began shortly after the introduction of R‐genes from Solanum demissum into S. tuberosum cultivars. However, the characteristics of R‐gene expression, the sensitivity of the phenotype to environmental and physiological parameters, and the diversity of experimental protocols make the comparison of data from different studies problematic. This prompted European teams working on P. infestans diversity to: (i) design a joint protocol, using detached leaflets from greenhouse‐grown plants of a shared set of differential cultivars inoculated with standardized suspensions of inoculum, and (ii) assess the performance of this protocol in a blind ring test involving 12 laboratories and 10 European isolates of the pathogen. A high level of consensus in the determination of virulence/avirulence to R1, R3, R4, R7, R8, R10 and R11 was achieved among the collaborators, showing that the protocol could be robustly applied across a range of laboratories. However, virulence to R2, R5 or R9 was detected more frequently in some laboratories, essentially from northern Europe; these genes are known to be highly sensitive to host and environmental conditions. The consensus determination was often markedly different from the original virulence phenotype of the isolates, suggesting virulence instability in stored P. infestans isolates. This indicates that creating reliable core collections of pathogen isolates with known virulences could be difficult.     

Induction of mutants of Fusarium oxysporum f. sp. lycopersici with altered virulence

Mutants ofFusarium oxysporum f. sp.lycopersici were obtained by UV irradiation. The mutants of race 1 and race 2 caused disease symptoms on plants with resistance genes against the corresponding wild type strains. Mutants of race 1 of the pathogen were stable, whereas mutants of race 2 lost the ability to cause disease symptoms in plants carrying the 1–2 resistance gene, after prolonged maintenenance on potato dextrose agar. Mutants of race 1 resembled race 2 in pathogenicity and they were vegetatively compatible with race 2, but no longer with race 1. These results suggest that the isolated strains with an altered virulence pattern have mutations in loci involved in avirulence.     

Allelism of theFcu-1 andFoc genes conferring resistance to fusarium wilt in cucumber

The inheritance of resistance toFusarium oxysporum f.sp.cucumerinum race 1 was determined in the cucumber cv. WIS-248 by analyzing segregation of F₁, F₂, and BC populations of crosses with the susceptible cv. Straight-8. Resistance was conferred by a single dominant gene. In an allelism test, it was proven that theFcu-1 gene, which confers resistance toF. oxysporum f.sp.cucumerinum races 1 and 2 in cucumber cv. SMR-18 and theFoc gene, which confers resistance toF. oxysporum f.sp.cucumerinum race 2 in cucumber cv. WIS-248, are indistinguishable.