Seed transmission of<Emphasis Type="Italic">Fusarium oxysporum</Emphasis> f.sp.<Emphasis Type="Italic">lactucae</Emphasis>

Twenty-seven seed samples belonging to the lettuce cultivars most frequently grown in Lombardy (northwestern Italy), in an area severely affected by Fusarium wilt of lettuce, were assayed for the presence ofFusarium oxysporum on a Fusarium-selective medium. Isolations were carried out on subsamples of seeds (500 to 1500) belonging to the same seed lots used for sowing, and either unwashed or disinfected in 1% sodium hypochloride. The pathogenicity of the isolates ofF. oxysporum obtained was tested in four trials carried out on lettuce cultivars of the butterhead type, very susceptible to Fusarium wilt. Nine of the 27 samples of seeds obtained from commercial seed lots used for sowing in fields affected by Fusarium wilt were contaminated byF. oxysporum. Among the 16 isolates ofF. oxysporum obtained, only one was isolated from disinfected seeds. Three of the isolates were pathogenic on the tested cultivars of lettuce, exhibiting a level of pathogenicity similar to that of the isolates ofF. oxysporum f.sp.lactucae obtained from infected wilted plants in Italy, USA and Taiwan, used as comparison. The results obtained indicate that lettuce seeds are a potential source of inoculum for Fusarium wilt of lettuce. The possibility of isolatingF. oxysporum f.sp.lactucae, although from a low percent of seeds, supports the hypothesis that the rapid spread of Fusarium wilt of lettuce observed recently in Italy is due to the use of infected propagation material. Measures for prevention and control of the disease are discussed. http://www.phytoparasitica.org posting Dec. 16, 2003.     

The use of GFP<Emphasis Type="Italic">-</Emphasis>transformed isolates to study infection of banana with <Emphasis Type="Italic">Fusarium oxysporum</Emphasis> f. sp. <Emphasis Type="Italic">cubense</Emphasis> race 4

Fusarium oxysporum f. sp. cubense (Foc) is the causal pathogen of Fusarium wilt of banana. To understand infection of banana roots by Foc race 4, we developed a green fluorescent protein (GFP)-tagged transformant and studied pathogenesis using fluorescence microscopy and confocal laser scanning microscopy. The transformation was efficient, and GFP expression was stable for at least six subcultures with fluorescence clearly visible in both hyphae and spores. The transformed Foc isolate also retained its pathogenicity and growth pattern, which was similar to that of the wild type. The study showed that: (i) Foc race 4 was capable of invading the epidermal cells of banana roots directly; (ii) potential invasion sites include epidermal cells of root caps and elongation zone, and natural wounds in the lateral root base; (iii) in banana roots, fungal hyphae were able to penetrate cell walls directly to grow inside and outside cells; and (iv) fungal spores were produced in the root system and rhizome. To better understand the interaction between Foc race 4 and bananas, nine banana cultivars were inoculated with the GFP-transformed pathogen. Root exudates from these cultivars were collected and their effect on conidia of the GFP-tagged Foc race 4 was determined. Our results showed that roots of the Foc race 4-susceptible banana plants were well colonized with the pathogen, but not those of the Foc race 4-resistant cultivars. Root exudates from highly resistant cultivars inhibited the germination and growth of the Fusarium wilt pathogen; those of moderately resistant cultivars reduced spore germination and hyphal growth, whereas the susceptible cultivars did not affect fungal germination and growth. The results of this work demonstrated that GFP-tagged Foc race 4 isolates are an effective tool to study plant–fungus interactions that could potentially be used for evaluating resistance in banana to Foc race 4 by means of root colonization studies. Banana root exudates could potentially also be used to identify cultivars in the Chinese Banana Germplasm Collection with resistance to the Fusarium wilt pathogen.     

Microbial community responses associated with the development of <Emphasis Type="Italic">Fusarium oxysporum</Emphasis> f. sp. <Emphasis Type="Italic">cucumerinum</Emphasis> after 24-epibrassinolide applications to shoots and roots in cucumber

Fusarium wilt, caused by Fusarium oxysporum f. sp. cucumerinum (FO), is one of the major diseases in cucumber (Cucumis sativus) production. Root and foliar applications of 24-epibrassinolide (EBL), an immobile phytohormone with antistress activity, were evaluated for their effects on the incidence of Fusarium wilt and changes in the microbial population and community in roots of cucumber plants. EBL pre-treatment to either roots or shoots significantly reduced disease severity followed by an improved plant growth regardless of the treatment methods applied. EBL applications decreased the Fusarium population on root surfaces and in nutrient solution, but increased the population of fungi and actinobacteria on root surfaces. PCR-DGGE analysis showed that FO-inoculation had significant effects on the bacterial community on root surfaces as expressed by a decreased diversity index and evenness index, but EBL applications alleviated these changes. Moreover, several kinds of decomposing bacteria and growth-promoting bacteria were identified from root surfaces of FO-inoculated plants and EBL-pre-treated plants, respectively. Overall, these results show that the microbial community on root surfaces was affected by a complex interaction between phytohormone-induced resistance and plant pathogens.     

Enhancement of germination of spores ofVerticillium dahliae andFusarium oxysporum f.sp.vasinfectum in vascular fluid from cotton plants infected with the root-knot nematode

Root-knot nematode [RKN] (Meloidogyne incognita) can increase the severity of Verticillium (V. dahliae) and Fusarium (F. oxysporum f.sp.vasinfectum) wilt diseases in cotton (Gossypium hirsutum). This study was conducted to determine some of the physiological responses caused by nematode invasion that might decrease resistance to vascular wilt diseases. The effect of RKN was investigated on spore germination and protein, carbohydrate and peroxidase content in the xylem fluids extracted from nematode-infected plants. Two cotton cultivars were used with different levels of resistance to both of the wilt pathogens. Spore germination was greater in the xylem fluids from nematode-infected plants than from nematode-free plants. The effect on spore germination was greater in theFusarium-resistant cultivar (51%). Analysis of these fluids showed a decrease in total protein and carbohydrate levels for both wilt-resistant cultivars, and an increase in peroxidase concentration. Fluids from nematode-free plants of theVerticillium-resistant cultivar contained 46% more peroxidase than theFusarium-resistant cultivar. The results provide further evidence that the effect of RKN on vascular wilt resistance is systemic and not only local. Changes in metabolites in the xylem pass from the root to the stem, accelerating disease development.     

芝麻枯萎病病原菌致病力室内鉴定方法

 芝麻枯萎病(Sesame Fusarium wilt, SFW)是由尖孢镰刀菌芝麻专化型(Fusarium oxysporum Schl. f. sp. sesami (Zap.), FOS)引起的一种土传真菌病害, 是世界芝麻生产上的主要病害之一。为测定FOS致病力, 本文选用郑芝98N09等4个芝麻品种, 在苗期对15个FOS菌株的致病力强弱进行了室内鉴定和评价。结果表明, 采用1×10⁶个/mL分生孢子悬浮液与无菌蛭石和无菌土壤按V1∶V2∶V6比例混合接菌(即最终接菌浓度为1.4×10⁵孢子/g土壤), 在接菌后第7 d幼苗开始出现枯萎病症状, 调查菌株致病性的最佳时间为接菌后第25 d~28 d;在供试15个FOS菌株中, 对4个品种均表现为强致病力的菌株有8个(DI>50), 均表现为弱致病力的菌株有5个(DI<20);不同芝麻品种对不同菌株的抗性有一定差异。该方法可应用于芝麻枯萎病病原菌致病性测定和芝麻种质抗枯萎病特性评价, 并为后续的机理研究提供了技术支持。     

Comparison of pathogenicity of the Fusarium crown rot (FCR) complex (<Emphasis Type="Italic">F. culmorum</Emphasis>, <Emphasis Type="Italic">F. pseudograminearum</Emphasis> and <Emphasis Type="Italic">F. graminearum)</Emphasis> on hard red spring and durum wheat

Fusarium species involved in the Fusarium crown rot (FCR) complex affect wheat in every stage of development from seedling to grain fill. This study was designed to compare the aggressiveness of the FCR complex members including F. culmorum, F. pseudograminearum and F. graminearum in causing seedling blight, decreased plant vigour and crown rot. To assess their relative pathogenicity, two hard red spring wheat cultivars and two durum wheat cultivars were inoculated in the field with five isolates from each of the three species for two years. Significant differences in patterns of pathogenicity were identified. In particular, F. culmorum caused greater seedling blight while F. pseudograminearum and F. graminearum caused greater crown rot. Greatest yield reductions were caused by F. pseudograminearum. Cultivar differences were identified with respect to seedling disease and late season crown rot. No interactions were identified between cultivar performance and isolates or species with which they were challenged.     

Characterization of <Emphasis Type="Italic">Fusarium oxysporum</Emphasis> f. sp. <Emphasis Type="Italic">melongenae</Emphasis> isolates from eggplant in Turkey by pathogenicity,VCG and RAPD analysis

Fusarium wilt is an economically important fungal disease of common eggplant (Solanum melongena) cultivated in the eastern Mediterranean region of Turkey. Seventy-four isolates of Fusarium oxysporum isolated from diseased eggplant displaying typical Fusarium wilt symptoms were screened for pathogenicity on the highly susceptible cv. ‘Pala’. All the isolates tested were pathogenic to eggplant and designated as Fusarium oxysporum f. sp. melongenae (Fomg). Genetic diversity among a core set of 20 Fomg isolates that were selected based upon geographic locations, were characterized by using pathogenicity, vegetative compatibility grouping (VCG), and random amplified polymorphic DNA (RAPD) analysis. The area under the disease progress curve (AUDPC) was calculated for each Fomg isolate until 21 days after inoculation (DAI). The most virulent isolate was identified as Fomg10 based on AUDPC, disease severity and vascular discoloration measurements at 21 DAI. At this date, a good correlation was observed between disease severity and AUDPC values for all isolates (r = 0.73). UPGMA (unweighted pair group method with arithmetic average) cluster analysis of RAPD data using Dice’s coefficient of similarity differentiated all the Fomg isolates tested, and indicated considerable genetic variation among Fomg isolates, but isolates from the same geographic region were grouped together. There was no direct correlation between clustering in the RAPD dendrogram and pathogenicity testing of Fomg isolates. Twenty isolates of Fomg were assigned to VCG 0320.     

Pathogenic variation and molecular characterization of <Emphasis Type="Italic">Fusarium</Emphasis> species isolated from wilted Welsh onion in Japan

Thirty-two isolates of Fusarium species were obtained from wilted Welsh onion (Allium fistulosum) grown on nine farms from six regions in Japan and identified as F. oxysporum (18 isolates), F. verticillioides (7 isolates), and F. solani (7 isolates). The pathogenicity of 32 isolates was tested on five commercial cultivars of Welsh onion and two cultivars of bulb onion in a seedling assay in a greenhouse. The Fusarium isolates varied in the degree of disease severity on the cultivars. Five F. oxysporum isolates (08, 15, 17, 22, and 30) had a higher virulence on the cultivars than the other isolates. The host range of these five isolates was limited to Allium species. Molecular characterization of Fusarium isolates was performed using polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP) analysis of the internal transcribed spacer (ITS) regions of ribosomal DNA. The 32 isolates were grouped into eight types (four types for F. oxysporum, one for F. verticillioides, and three for F. solani). Restriction patterns of the ITS region were not related to pathogenicity. However, the haplotypes obtained with five enzymes (RsaI, HinfI, HaeIII, ScrFI, and MspI) and the phylogenetic analysis permitted the discernment of the three Fusarium species. The PCR-RFLP analysis should provide a rapid, simple method for differentiating Fusaruim species isolated from wilted Welsh onion in Japan.     

Validation of a QTL for resistance to ascochyta blight linked to resistance to fusarium wilt race 5 in chickpea (<Emphasis Type="Italic">Cicer arietinum</Emphasis> L.)

Ascochyta blight caused by Ascochyta rabiei and fusarium wilt caused by Fusarium oxysporum. f. sp. ciceris are the two most serious diseases of chickpea (Cicer arietinum). Quantitative trait loci (QTL) or genes for ascochyta blight resistance and a cluster of resistance genes for several fusarium wilt races (foc1, foc3, foc4 and foc5) located on LG2 of the chickpea map have been reported independently. In order to validate these results and study the linkage relationship between the loci that confer resistance to blight and wilt, an intraspecific chickpea recombinant inbred lines (RIL) population that segregates for resistance to both diseases was studied. A new LG2 was established using sequence tagged microsatellite sites (STMS) markers selected from other chickpea maps. Resistance to race 5 of F. oxysporum (foc5) was inherited as a single gene and mapped to LG2, flanked by the STMS markers TA110 (6.5 cM apart) and TA59 (8.9 cM apart). A QTL for resistance to ascochyta blight (QTL_AR3) was also detected on LG2 using evaluation data obtained separately in two cropping seasons. This genomic region, where QTL_AR3 is located, was highly saturated with STMS markers. STMS TA194 appeared tightly linked to QTL_AR3 and was flanked by the STMS markers TR58 and TS82 (6.5 cM apart). The genetic distance between foc5 and QTL_AR3 peak was around 24 cM including six markers within this interval. The markers linked to both loci could facilitate the pyramiding of resistance genes for both diseases through MAS.     

兰州百合枯萎病病原菌鉴定及罹病组织超微结构观察

为明确引起甘肃省兰州百合主产区百合枯萎病的致病镰孢菌种类,对从百合主产区枯萎病罹病植株上分离纯化的4株镰孢菌株进行形态学鉴定、分子生物学鉴定以及致病性测定,同时利用电子显微镜对尖孢镰孢菌Fusarium oxysporum侵入百合鳞片后的细胞超微结构进行观察。结果表明:4株镰孢菌菌株经鉴定分别为尖孢镰孢菌、茄病镰孢菌F. solani、三线镰孢菌F. tricinctum和燕麦镰孢菌F. avenaceum。4株镰孢菌菌株的致病力由强到弱的顺序依次是尖孢镰孢菌、燕麦镰孢菌、茄病镰孢菌、三线镰孢菌;尖孢镰孢菌侵入后,鳞片细胞壁、细胞质膜和细胞核结构被破坏,细胞核附近出现大量线粒体,细胞中淀粉粒数量减少。表明尖孢镰孢菌是兰州百合枯萎病防治的重点防控对象。     

Benomyl-resistant Fusarium-isolates in ecological studies on the biological control of fusarium wilt in carnation

Ecological properties and stability of benomyl resistance of three benomyl-resistant mutants of nonpathogenicFusarium-isolates antagonistic to fusarium wilt in carnation, and three benomyl-resistant mutants of a pathogenic isolate ofFusarium oxysporum f.sp.dianthi were evaluatedin vitro and in glasshouse experiments. The benomyl resistance of the nonpathogenic mutants was stable under all conditions tested, also after a 1000-fold increase of the population in sterilized soil. Mutants of the pathogen were stable during allin vitro tests, but after proliferation in carnation stems only part of the population was benomyl resistant.Compared to the wild type, mutants of the pathogen were less pathogenic, also if thein vitro propeties were similar. Colonization of carnation by benomyl-resistant nonpathogenicFusarium in the presence of the pathogen showed that the antagonistic effect correlated with the presence of the nonpathogenic isolates within the carnation stem. The wild types and two of the mutant nonpathogenicFusarium-isolates controlled fusarium wilt in the susceptible cultivar Lena for 50% or more.UV-induced benomyl resistance appeared to be a valuable marker to distinguish between differentFusarium isolates and to study the population dynamics, but intensive screening of the mutants is a prerequisite since alterations in antagonism and pathogenicity can occur.     

Pathogenic variability in Ethiopian isolates of Fusarium oxysporum f. sp. ciceris and reaction of chickpea improved varieties to the isolates

Twenty-four isolates of Fusarium oxysporum f. sp. ciceris were isolated from wilted chickpea plants obtained from different districts and ‘wilt sickplots’ of central Ethiopia to assess variability in pathogenecity of the populations. Each isolate was tested on 10 different chickpea lines and eight improved chickpea varieties. Isolates showed highly significant variation in wilt severity on the differential lines and improved varieties. Based on the reaction types induced on differential lines, isolates were grouped into four corresponding races. Of the 24 isolates, F13, F20 and F22 were the most virulent. Isolates of race 3 were found in all of the districts and ‘wilt sickplots’ studied. Improved chickpea varieties also showed differential reactions to the isolates. All varieties were resistant to isolates of race 3, while varieties Arerti and DZ-10-4 were resistant to all isolates tested, showing the lowest mean wilt severity. Varieties DZ-10-11 and Maryie were susceptible to isolates F13, F20 and F22 and showed the highest mean wilt severity. Identification of races can be useful in breeding chickpea varieties resistant to wilt. The differential reactions of the improved varieties against different races might be important in managing chickpea wilt through gene deployment.     

Fusarium rot of melon is caused by several Fusarium species

Fusarium rot of melon, caused by species of the genus Fusarium, has become an important postharvest disease for many Brazilian producers. Due to the delayed onset of symptoms, this disease is often only detected when fruits arrive at the importing country, thus generating economic loss for the exportation of the fruit. This study was developed with the aim of investigating which Fusarium species cause fruit rot in melon and to evaluate any differences in aggressiveness and development of symptoms. Species were identified through phylogenetic analysis of two loci and morphological markers. The 28 isolates obtained from diseased melon fruits of different commercial cultivars were identified as Fusarium falciforme (FSSC), F. sulawesiense, F. pernambucanum (FIESC), and F. kalimantanense (FOSC). Three isolates belong to a new phylogenetic lineage within the F. fujikuroi species complex (FFSC). All isolates were tested for pathogenicity, and first symptoms of rot in Canary melon were observed 2 days after inoculation. Isolates of F. falciforme and F. sulawesiense were shown to be more aggressive. Our results extend information on Fusarium species that cause fruit rot in melon and support the development of management strategies, as there is currently no efficient control for this disease. To our knowledge, this is the first report of the occurrence of species of the FSSC, FOSC, and FFSC from muskmelon fruits in Brazil.     

Occurrence of Fusarium oxysporum f. sp. melonis Race 1 in Japan

In 1994, Fusarium wilt of melon cultivars which are resistant to races 0 and 2 of Fusarium oxysporum f. sp. melonis was observed in southern area of the Lake Biwa region, Shiga prefecture. In commercial fields, mature plants of cv. Amus which were grafted onto cv. Enken Daigi 2, and of cv. FR Amus showed yellowing, wilting and finally death before harvesting of fruits. Diseased plants had vascular and root discolorations, and their stem sections yielded typical colonies of F. oxysporum. When the Shiga strains were tested for their pathogenicity to 12 species of cucurbits, they caused wilts only on melon. Using race differential cultivars of melon, the Shiga strains were classified as race 1 of F. oxysporum f. sp. melonis, which has not been reported in Japan. To further characterize their pathogenicity, the strains were used to inoculate 46 additional cultivars of melon, oriental melon and oriental pickling melon. All the race 1 strains were pathogenic to the cultivars tested, and their host range was apparently different from those of strains belonging to other races (races 0, 2 and 1,2y). DNA fingerprinting with a repetitive DNA sequence, FOLR3, differentiated race 1 strains from strains of races 0 and 2, but not from race 1,2y strains. Received 2 July 1999/ Accepted in revised form 30 September 1999     

Fusarium commune associated with wilt and root rot disease in rice

Wilt and root rot disease in plants has been caused mainly by Fusarium species. Previous studies reported that members of the Fusarium oxysporum species complex (FOSC) were usually associated with this disease, but there has been no report of it being caused in rice by specific Fusarium species. However, in this study, Fusarium commune was identified and characterized as a causal agent of wilt and root rot disease of rice. Four Fusarium isolates (BD005R, BD014R, BD019R, and BD020R) were obtained from different parts (root, stem, and seeds) of diseased rice plants. In morphological studies, these isolates produced key characteristics of F. commune, such as long and slender monophialides, polyphialides, and abundant chlamydospores. In molecular studies, the isolates were identified as F. commune based on sequences of the translation elongation factor 1-α (TEF1) gene that had 99.7%–100% sequence identity with the reference strain F. commune NRRL 28058. The phylogenetic tree showed that all four isolates belonged to the F. commune clade. A mating type test determined that three isolates carried MAT1-2. Their teleomorph stage was still unknown. Pathogenicity assays showed that all the isolates produced wilt and root rot symptoms and the isolate BD019R was observed as the most virulent among the isolates. To our knowledge, this is the first report of F. commune causing wilt and root rot disease on rice.     

Fusarium wilt of banana: biology,epidemiology and management

Fusarium oxysporum is a soil borne hyphomycete that causes vascular wilts in several crop plants. A variety of remedial measures such as the use of fungicides, soil amendments and biological antagonists have proved insufficient in controlling F. oxysporum. Ever since it was first reported in banana crop, the only effective control strategy known is planting of resistant cultivars. However, presumably due to the high mutation rates and rapid co-evolution with its host, Fusarium wilt has surmounted host defense barriers and has already begun infecting even the resistant Cavendish varieties that dominate export markets worldwide. Transgenic banana plants showing enhanced resistance to Fusarium wilt have been developed in recent past, but they remain largely confined to the laboratory. The importance of banana as source of food and income in developing countries world over and the need to develop Fusarium wilt tolerant cultivars by novel biotechnological approaches is detailed herein. In this communication, we review the biology and management of Fusarium wilt in banana with the aim of providing the baseline of information to encourage much needed research on integrated management of this destructive banana crop disease problem.     

Integrated management of Meloidogyne incognita‐Fusarium oxysporum f. sp. ciceri wilt disease complex in chickpea

A field experiment was conducted to study the management of a root‐knot nematode, Meloidogyne incognita (Kofoid and White) Chitw. (Tylenchida: Heteroderidae)‐wilt inducing fungi, Fusarium oxysporum Schlecht. emend. Snyd & Hans. f. sp. ciceri (Padwick) Snyd. & Hans. (Moniliales: Tuberculariaceae) wilt disease complex in chickpea (Cicer arietinum L.). cv. Annegiri. The results indicated that integration of soil solarization (for 6 weeks), VA mycorrhizal fungus (VAM), Glomus fasciculatum inoculation (12g/hill) and seed treatment with carbosulfan (3% w/w) was highly effective in reducing population levels of both pathogens, root‐knotdisease and wilt incidence and in increasing chickpea grain yield significantly. However, seed treatment with carbendazim (0.25% w/w) together with carbosulfan (3% w/w) was not only effective in reducing the wilt disease complex but also economic with an incremental cost: benefit ratio of 1: 2.4.     

An Experiment to Assess Losses Caused by Frit-fly (Oscinella frit L) Shoot Attack and the Application of Phorate in a Crop of Sweet Corn (Zea mays L)

Wilt caused by the fungus Fusarium oxysporum f. sp. ciceris adversely affects the productivity of cultivated chickpea. For the management of this disease, seed and soil application formulations developed from another fungus, Trichoderma species, were evaluated. In pot experiments, T. harzianum-based formulations Pusa 5SD for seed dressing and Pusa Biopellet (PBP) 10G and Pusa Biogranule (PBG) 5 for soil application, and T. viride-based formulations Pusa 5SD for seed dressing and PBP 4G and PBG 4 for soil application, were found to be highly effective against the disease. A combination of PBP 4G (T. viride) for soil application and Pusa 5SD (T. harzianum) for seed treatment together with a fungicide, carboxin, provided the highest seed germination, shoot and root lengths and grain yield with the lowest incidence of wilt in chickpea under field conditions. Individually, soil application of PBP 4G, and seed treatment with Pusa 5SD were effective in reducing the incidence of wilt and increasing the grain yield of chickpea, but their effectiveness was greater when applied as a combination. Thus, combined application of the formulations of two different species of Trichoderma in two modes of application is recommended for the management of chickpea wilt.     

Screening of cultivated and wild adzuki bean for resistance to race 3 of <Emphasis Type="Italic">Cadophora gregata</Emphasis> f. sp. <Emphasis Type="Italic">adzukicola</Emphasis>, cause of brown stem rot

Two diseases of adzuki bean, brown stem rot (BSR, caused by Cadophora gregata f. sp. adzukicola) and adzuki bean Fusarium wilt (AFW, caused by Fusarium oxysporum f. sp. adzukicola), are serious problems in Hokkaido and have been controlled using cultivars with multiple resistance. However, because a new race of BSR, designated race 3, was identified, sources of parental adzuki bean for resistance to race 3 were needed. Therefore, we examined 67 cultivars and lines of cultivated and wild adzuki bean maintained at the Tokachi Agricultural Experiment Station using a root-dip inoculation method. Consequently, nine adzuki bean cultivars, one wild adzuki bean accession and 30 lines (including two lines resistant to all the three races of BSR and AFW) were confirmed to be resistant or tolerant to race 3 of BSR, and we found a cultivar Akamame as well as a wild adzuki bean Acc2515 to be a new source for a resistance gene to the race 3. This cultivar also holds promise as a source of resistance against other races of BSR and AFW.