Inheritance and allelic relationships of fertility restoration genes for seven new sources of male-sterile cytoplasm in sunflower

The inheritance of fertility restoration of six mitomycin C and streptomycin‐induced cytoplasmic male‐sterile (cms) mutants and one cms line derived from Native American cultivar PI 432513 in sunflower was evaluated. These seven new cms sources were also compared with the commercially used cms PET1 (Helianthus petiolaris Nutt.) cytoplasm, using USDA inbred lines with restoration genes (Rf₁) specific for cms PET1 and new restoration lines identified for cms PI 432513. Restoration genes for cms PI 432513 were found in ‘Armavir’, VNIIMK, P21 and male‐fertile (MF) plants of PI 432513. F₂ and F₃ segregation ratios of crosses between cms PI 432513 and these restoration sources indicated a single dominant gene controlled fertility restoration. Progenies of cms PI 432513 testcrossed with F₁’s of half‐diallel crosses among the respective four homozygous restoration lines and RHA 274 suggested that the restoration genes of RHA 274, VNIIMK, P21 and PI 432513 were at the same locus. Restoration genes from VNIIMK, P21 and PI 432513 satisfactorily restored pollen stainability in the heterozygous condition. A very weak expression of the Rf gene in ‘Armavir’ was observed in the heterozygous condition. Fertility restoration capability of these genes for the six mutant cms HA 89 and cms HA 89 (in PET1 cytoplasm) was observed. The mutant cms HA 89 lines were also restored completely by RHA 266, RHA 274, RHA 280 and RHA 296, and F₂’s segregation ratios indicated single dominant gene control, implying a common cytoplasmic male sterility in all lines. F₁’s of half‐diallel crosses among RHA 266, RHA 273, RHA 274, RHA 280 and RHA 296 were testcrossed onto the cms lines, and their all MF progenies among lines, except RHA 280, confirmed that fertility restoration was controlled by a single Rf₁ gene locus. The restoration gene in confection line RHA 280, namely Rf₃, was at a different locus than Rf₁ and was equally capable of restoring all the cms lines. Cms HA 89 mutants and cms PI 432513 are in H. annuus cytoplasm, and are agronomically equal in hybrid performance to the cms PET1 used in commercial sunflower hybrids. These new cms lines will provide immediate alternative cms sources for reducing the genetic vulnerability resulting from the exclusive use of the single cms source PET1 in sunflower hybrid production.     

Colocation of downy mildew (Plasmopara halstedii) resistance genes in sunflower (Helianthus annuus L.)

Summary The Pl6 locus in the inbred sunflower (Helianthus annuus L.) line HA335 giving resistance to French races of downy mildew (Plasmopara halstedii (Farl.) Berl. & de Toni. was localized by molecular techniques. A bulked segregant analysis was made on the F2 progeny from a cross between this line and H52, a downy mildew susceptible line. The resistance gene in HA335 was found to have the same linked RFLP marker loci as those determined for Pl1 (resistance to race 1 in the line RHA266) on linkage group 1 of the consensus RFLP map of the cultivated sunflower. Pl1 and Pl6 thus appear either to be allelic or closely linked. The implications for sunflower breeding are discussed.     

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.     

Inheritance of resistance to sunflower downy mildew races 1, 2 and 3 in cultivated sunflower

Sunflower downy mildew (SDM) caused by Plasmopara halstedii, is a major disease of sunflower. Eleven resistance genes have been identified, but allelic relationships among these genes are not clear. This study examined the inheritance and allelic relationships of genes conferring resistance to SDM races 1, 2 and 3 (virulence phenotypes 100, 300 and 700, respectively) and confirmed a twelfth resistance gene. Three USDA Plant Introductions, AMES 3235, PI 497250, and PI 497938, and three released lines, RHA 266, RHA 274 and DM‐2 were studied. RHA 266 has only the Pl₁ gene for race 1 resistance. Digenic inheritance of resistance was found in AMES 3235, PI 497250, and RHA 274. These lines have the Pl₁ and Pl₁₂ genes, conferring resistance to race 1, and the Pl₂ and Pl₁₁ genes, conferring resistance to race 2. DM‐2 and PI 497938 have Pl₁₂ (but not Pl₁ for resistance to race 1, the Pl₁₂ gene (but not the Pl₂) for resistance to race 2, and Pl₅ for resistance to race 3. These resistance genes will serve as a foundation for future gene designations and genetic diversity studies of resistance to SDM.     

Characterization for agronomic use of cytoplasmic male-sterility in sunflower (Helianthus annuus L.) introduced from H. resinosus Small

A backcrossing programme was carried out both to assess the stability of a cytoplasmic male‐sterility (CMS) source from Helianthus resinosus, designated RES1, and to incorporate it into inbred sunflower lines (HA89, RHA271, RHA801). All the progenies, grown in different environments, were completely male‐sterile. This suggests that the expression of this cytoplasm is stable. Female‐fertility of lines HA89, RHA271 and RHA801 carrying CMS RES1 were compared with those of the corresponding fertile inbred lines. There were no differences in the number of seeds per head. This indicates that female‐fertility is not affected by RES1 cytoplasm. Cytological studies showed that meiosis proceeds normally until the tetrad stage; consequently, the absence of pollen is caused by alterations that take place during postmeiotic stages. With the aim of identifying male‐fertility restorer genotypes, crosses were made between HA89 (CMS RES1) plants and different annual diploid and perennial hexaploid Helianthus species. All the diploid germplasm evaluated behaved as a CMS RES1 maintainer. However, the hexaploid species, H. resinosus, H. x laetiflorus, H. pauciflorus and H. tuberosus, restored pollen fertility in CMS RES1 plants.     

Generation mean analysis of resistance to downy mildew in adult muskmelon plants

Downy mildew (Pseudoperonospora cubensis) is the most devastating disease in muskmelon (Cucumis melo). A generation mean analysis study was designed to determine the types of gene action and estimate the heritability for resistance to downy mildew in four selected crosses of muskmelon. Generation mean analysis revealed that genetic dominance may be of greater importance for expression of resistance to downy mildew in both greenhouse and field experiments and in all the crosses. The F₁ mean was significantly lesser than the mid-parent value and skewed towards resistant parent in all the crosses. Negative sign associated with gene effects indicated, in those crosses, that disease level could be decreased in relation to midparent. All the crosses expressed significant and positive additive (d) gene effects. Dominance (h) and dominance × dominance (l) gene effects had opposite sign in all crosses and both experiments, which implied duplicate type of gene action. High mid-parent heterosis in all the crosses indicated strong dominance effects (as combination of parental alleles) for resistance to downy mildew. In all the crosses, both resistant and susceptible parent contributed one or more dominant/partially dominant factors for resistance. Estimates of broad-sense heritability were high and relatively consistent in both experiments. The two different screening experiments showed that fixable gene effects (d + i) were lower than the non-fixable gene effects (h + l) in all the crosses indicating greater role of non-additive effects in the inheritance of resistance to downy mildew. Resistance to downy appeared to be controlled mainly by dominance effects, therefore the inbred lines IIHR 121 and IIHR 122 could be used strategically to exploit heterotic effects.     

Involvement of higher order interactions addressing complex polygenetically controlled inheritance of downy mildew [Sclerospora graminicola (Sacc.) Schrot] resistance in pearl millet [Pennisetum glaucum (L.) R.Br.]

Inheritance of downy mildew [Sclerospora graminicola (Sacc.) Schrot]resistance was studied using generation mean analysis in pearl millet [Pennisetum glaucum (L.) R.Br.]. Eleven basic generations, namely, P₁, P₂, F₁, F₂, B₁, B₂, B₁F₂, B₂F₂, L₁, L₂ and L₃ of three crosses involving six diverse lines for downy mildew incidence were evaluated under artificial epiphytotic conditions over two environments. The downy mildew incidence was best fitting for digenic, trigenic and tetragenic ratios when fitted into classical Mendelian ratios demonstrating involvement of two or more genes. Digenic and trigenic interaction models were adequate in the case of crosses I and III respectively, to account for the total variability in generation means. Unlike severity, comparative estimates of gene effects over two environments were mostly consistent in all crosses for prevalence. Most of the epistatic and major gene effects were found significant in all crosses for both the disease traits. Non-allelic interactions particularly at three-gene loci viz., w (additive × additive × additive) and y (additive × dominance × dominance) in cross II and all trigenic interactions in cross III were predominant. Duplicate dominance (cross I) and complementary epistasis (crosses II and III) were observed for both the traits revealing inconsistency of gene effects over crosses. The gd₁ (interaction of additive gene effect with e₁) and gh₁(interaction of dominant gene effect with e₁) were significant in crosses I and II, indicating interaction of additive and dominance gene effects with environments. Thus a breeding method that can mop up the resistant genes to form superior gene constellations interacting in a favorable manner against pathotype I would be more suitable to accelerate the pace of resistance improvement. This revised version was published online in August 2006 with corrections to the Cover Date.     

Resistance to Sunflower Rust and its Inheritance in Wild Sunflower Species

Resistance to the prevailing races of sunflower rust, Puccinia hehanthi Schw., is lacking in the commercial hybrids (Helianthus annuus L.). The objective of this study was to identify new sources of resistance to the four North American rust races in wild Helianthus species, and to determine their mode of inheritance. Seventy-eight accessions of H. annuus L., H. argophyllus Torrey and Gray, and H. petiolans Nutt. were evaluated in the greenhouse. Resistance to races 1, 2, 3, and 4 was observed in 25, 28, 15, and 26% of the plants, respectively, and 10% of the plants were resistant to all four races. Seven accessions that had a high percentage of resistant plants to all the four races were selected and one resistant plant from each accession was crossed with susceptible inbred line HA89. Three to four F₁ plants resistant to all four races from each cross were backcrossed with HA89. F₁ plants from PI-413118 × HA89 and PI 413175 × HA89 were resistant to all four races. The PI 413023 × HA89 F₁ plants were 100 % resistant to races 3 and 4 and segregated in a 3: 1 resistant (R) to susceptible (S) ratio to races 1 and 2. The other four F₁ combinations segregated 3R: IS ratios to all four races. Bc₁F₁ progenies revealed that plants from PI 413048, PI 413037, PI 413038, and PI 413171 used in the crosses possessed two dominant genes in heterozygous condition for resistance to each of the four races, whereas plants from PI 413023 possessed two dominant genes in heterozygous condition for resistance to each of races 1 and 2, and one dominant resistance gene in homozygous condition for each of races 3 and 4. Plants from PI 413118 and PI 413175 carried a single dominant gene in homozygous condition for resistance against each of the four races.     

Inheritance of downy mildew resistance in Indian cauliflower (group III)

Summary Inheritance of downy mildew (Peronospora parasitica) resistance in three resistant x susceptible crosses, one susceptible x susceptible and one resistant x resistant cross were studied in Indian cauliflower (Group III) over the two years (1990 and 1991). No significant difference was observed between the years for various estimates and hence pooled data are presented. Downy mildew resistance in crosses cc×HR 5-4 and 3-5-1-1×244 (R×S) is governed by single dominant gene PPA3 but in cross cc×244 (R×S), recessive epistasis was observed. The resistance level was not improved in both the cc×3-5-1-1 (R×R) and 244×267-6-9 (S×S) crosses. Exploitation of downy mildew resistance from cc and 3-5-1-1 in F₁ hybrid is explained in detail.     

Sources of Resistance to Powdery Mildew in Barley Lines Derived from Hordeum spontaneum Collected in Israel

Fourty two barley lines direved from the F₇ of crosses between barley cultivars and different accessions of Hordeum spontaneum collected in Israel and 30 lines or varieties with known genes for resistance to powdery mildew were included m this study. Eleven European and three Israeli powdery mildew cultures, possessing virulence genes corresponding to known resistance genes, were used to make comparisons between the varieties with known resistance genes and H. spontaneum derived lines. The reaction pattern of 39 H. spontaneum derived lines was clearly different from the reaction pattern o; any of the known genes for mildew resistance included in this study. Only two cases were observed in which the reaction pattern of H. spontaneum derived lines agreed with reaction patterns of known genes for mildew resistance viz. Ml-a9 and Ml-p. Trie Mildew resistance of one line apparently traces back to uncontrolled outcrossing with a Ml-a.6+Ml-g resistant cultivar. Since the majority of the 42 host genotypes tested showed distinctive variation in resistant reaction types against different mildew cultures, this study docs not support the assumption that differences in resistant infection types against distinct mildew cultures are sufficient to indicate the presence of supplementary genes for resistance in a given genotype of the host. The results justify the conclusion that the natural population of H. spontaneum in Israel forms a large gene pool for mildew resistance which is not yet used m cultivated barley.     

Interspecific amphiploid‐derived alloplasmic male sterility with defective anthers,narrow disc florets and small ray flowers in sunflower

The cytoplasmic male‐sterility (CMS)/fertility‐restoration system is important for hybrid sunflower (Helianthus annuus L.) seed production. The objective of this study was to characterize two novel alloplasmic CMSs, designated CMS GRO1 and CMS MAX3, with defective anthers, narrow disc florets with no swollen corolla, and short, narrow ray flowers derived from two tetraploid amphiploids (AMPs). Among 26 tested lines, only AMP Helianthus cusickii/P 21 and HA 410 failed to restore male‐fertility. Segregation of CMS, male‐fertile plants and plants with reduced male‐fertility was observed both in the testcross progeny of a six line half‐diallel cross of F₁s with CMS MAX3 and in an F₂ population of CMS GRO1 × RHA 274. Male‐fertility restoration was controlled by at least two dominant genes. Detailed analysis of the mitochondrial genes may provide insight into the differences between these CMSs and other CMS lines. The new CMSs will facilitate the studies of the incompatibility between cytoplasmic and nuclear genes, especially for the alloplasmic CMS involving perennial species, and also provide unique ornamental flower types and CMS sources for hybrid sunflower breeding.     

Screening of melon (Cucumis melo L.) germplasm for multiple disease resistance

Summary Melon germplasm was screened for cucumber green mottle mosaic virus (CGMMV), powdery mildew (Sphaerotheca fuliginea), downy mildew (Pseudoperonospora cubensis) and Fusarium wilt (Fusarium oxysporum f. sp. melonis) resistance under artificial conditions except downy mildew for which screening was done under natural epiphytotic conditions. High level resistance to all the four diseases was not recorded in any of the collections tested. Nevertheless, ertheless, resistance to three diseases was located in three germplasm. Wild Cucumis species C. figarei exhibited absolute resistance to CGMMV and Fusarium wilt and high level resistance to downy mildew. Phoot or snapmelon (Cucumis melo var. momordica) — a non-dessert from of Indian origin—was highly resistant to downy mildew and resistant to CGMMV and medium resistant to Fusarium wilt. Iroquois was resistant to powdery mildew and medium resistant to downy mildew and CGMMV.     

Genetic variability for resistance to sorghum downy mildew (Peronosclerospora sorghi) and Rajasthan downy mildew (P. heteropogoni) in the tropical/sub-tropical Asian maize germplasm

Downy mildews cause considerable damage to maize (Zea mays L.) worldwide, particularly in the tropical Asia. We have evaluated a set of 42 tropical/sub-tropical maize inbred lines developed in different countries in Asia (India, Thailand and Philippines), and Mexico, for analysing the genetic variability for resistance to sorghum downy mildew [Peronosclerospora sorghi; SDM] and Rajasthan downy mildew [P. heteropogoni; RDM]. Experiments were carried out in replicated trials under artificial infection in field conditions against SDM and RDM at Mandya in Karnataka, India, and Udaipur in Rajasthan, India, respectively, during 1999 and 2000. The study resulted in identification of five inbred lines offering consistent and strong resistance to both SDM and RDM, while several inbred lines revealed resistance only to RDM. It was also revealed that the SDM-resistant inbreds are invariably resistant to RDM, while the RDM-resistant inbreds might show differential responses to the SDM. The maize inbred lines identified in this study with broad-spectrum resistance to downy mildews could be potentially useful for basic and applied research work on downy mildews in tropical Asia. This revised version was published online in August 2006 with corrections to the Cover Date.     

Inheritance of resistance to Pseudoperonospora cubensis Rost. In cucumber (Cucumis sativus L.)

Summary Since June 1973 Pseudoperonospora cubensis (Berk & Curt) Rost., which causes downy mildew in cucumber, occurs in the Netherlands. The resistance against this disease appears to be based on one recessive gene in linkage with the dominant gene D for dull green fruit skin colour. It is demonstrated that this recessive gene is also linked with one of the genes for resistance to powdery mildew present in the variety Ashley.The powdery mildew resistant lines tested are also resistant against downy mildew, the linkage with the gene D having been broken.Stationed at Breeding Station Pannevis B.V., De Lier, the Netherlands.     

Genetic markers linked to Striga gesnerioides resistance for the improvement of Ghanaian cowpea (Vigna unguiculata) cultivars

The parasitic plant, Striga gesnerioides (Willd.) Vatke, is one of the most important constraints of cowpea production and food security in West Africa. Currently, few Striga resistant cowpea varieties have been developed that are well‐adapted to the dry savannah regions of Ghana. While genes conferring resistance to Striga races SG1, SG3 and SG5 have been mapped, the genetic locus of resistance to the race of Striga found in Ghana (SG‐GH) has not been characterized. Here, we report identification of genetic markers linked to SG‐GH resistance and define the relationship of this locus to SG3 resistance in recombinant inbred line populations generated from crosses between Striga resistant IT97K‐499‐35 and the Striga susceptible varieties Apagbaala and SARC‐LO2. The populations were genotyped with five genetic markers associated with SG3 and SG5 Striga resistance genes and a genetic map was developed. Genes conferring resistance to SG‐GH and SG3 mapped 4.2 cM from each other on chromosome Vu11. The identification of genetic markers linked to SG‐GH resistance will facilitate the marker‐assisted development of high‐quality Striga resistant cowpea varieties in Ghana.     

Quantitative trait loci with additive and epistatic effects underlying resistance to two HG types of soybean cyst nematode

The resistance of soybean (Glycine max L. Merr.) cultivars varies with the different races of the soybean cyst nematode (SCN), Heterodera glycines, referred to as HG types (biotypes). Resistant cultivars with durable resistance are emphasized in recent years. The aim here was to identify quantitative trait loci (QTLs) for resistance to two SCN HG types (HG type 2.5.7, race 1; and HG type 1.2.3.5.7, race 4) in resistant cultivar ‘L‐10’ and to analyse the additive and epistatic effects of the identified QTLs. A total of 140 F₅‐derived F₁₀ recombinant inbred lines (F_5:10 RILs) were advanced via single‐seed‐descent from the cross between ‘L‐10’ (broadly resistant to SCN) and “Heinong 37” (SCN‐susceptible). For SCN HG type 2.5.7 and HG type 1.2.3.5.7 resistance, three and six QTLs for resistance to SCN HG type 2.5.7 and HG type 1.2.3.5.7 were identified, respectively, most of which could explain <10% of the phenotypic variation. Among these QTLs, five were identified over 2 years, while the other QTLs were detected in either 2009 or 2010. QSCN1‐2, located near the SSR marker Sat_069 of linkage group D1b (Chromosome, 2), was responsible for the largest proportion of phenotypic variation (16.01% in 2009 and 18.94% in 2010), suggested that it could effectively be used as a candidate QTL for the marker‐assisted selection (MAS) of soybean lines resistant to SCN. Additionally, for SCN HG type 2.5.7 and HG type 1.2.3.5.7 resistance, two and four QTLs showed an additive effect (a), respectively. One epistatic pair of QTLs (QSCN1‐1‐QSCN1‐3) for SCN HG type 2.5.7 resistance and eight epistatic pairs of QTLs for SCN HG type 1.2.3.5.7 resistance were found to have significant aa effects, among which one pair of QTLs (QSCN4‐4 and QSCN4‐5) contributed a large proportion of aa effects (3%). The results indicated that additive and epistatic effects could significantly affect SCN resistance. Therefore, both of a and aa effects should be considered in MAS programmes.     

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

Quantitative resistance to sunflower downy mildew was studied on inbred lines and hybrids not carrying efficient major gene resistance, in field trials in one to four sites over 3 years. Hybrids from factorial crosses showed that inheritance is under additive control and comparison with reactions of parental inbred lines gave narrow sense heritabilities of 27–57%. Analysis of a polymorphic recombinant inbred line population without efficient major gene resistance indicated that two highly significant Quantitative Trait Loci (QTL) explained 42% of variation in field reaction to downy mildew. These QTL were mapped on linkage groups 8 and 10, and do not appear related to any of the known major resistance gene clusters. Possible bases of this type of resistance and its use in breeding are discussed.     

Screening for resistance to broomrape (Orobanche cernua) in cultivated sunflower

Racial evolution of sunflower broomrape (Orobanche cernua) has been very rapid in Spain during recent years, in which resistance has been overcome several times and there has been an important increase in areas infested with this parasitic angiosperm. In order to find resistance to a highly virulent population of sunflower broomrape that could be used directly in breeding programmes, three different sets of cultivated plant material composed of 429 entries were tested by artificial inoculation. All evaluated inbred lines from Moden, Canada, were fully susceptible. Out of the 240 P.I. accessions tested, only 10 segregated for resistance to broomrape, the rest being susceptible. From the 160 USDA breeding lines evaluated, 5% were resistant and 19% segregated for resistance to O. cernua. These lines traced back mainly to crosses of RHA 274 and RHA 801 with Russian, Turkish and Romanian hybrids. The origin of P.I. accessions that segregated for resistance were primarily derived from the former USSR and from Romania.     

Quantitative resistance to downy mildew (Plasmopara halstedii) in sunflower (Helianthus annuus)

Downy mildew of sunflower, caused by the Oomycete, Plasmopara halstedii is at present controlled by major resistance genes. However, the pathogen has shown a considerable capacity for changes in virulence and these resistance genes are overcome only a few years after they have been introduced into new sunflower varieties. This paper presents research for quantitative, non-race-specific resistance independent of major genes. The reaction of cultivated sunflower genotypes to field attack by downy mildew was studied over 4 years in several environments and in the presence of the two most common races in France: 703 and 710. An experimental protocol with pre-emergence irrigation was developed, making it possible to observe downy mildew reaction whatever the weather conditions. Significant levels of partial resistance were observed in about 50 inbred sunflower lines among the 800 observed. These results suggest that it should be possible to select for non-race-specific downy mildew resistance and to include it in modern varieties. However, since this non-specific resistance is partial, it may be necessary to combine it with major gene resistance. Possible strategies are discussed to obtain durable resistance to downy mildew.     

Two genes and linked RAPD markers involved in resistance to Fusarium oxysporum f. sp. Ciceris race 0 in chickpea

The inheritance of resistance to fusarium wilt race 0 of chickpea and linked random amplified polymorphic DNA (RAPD) markers were studied in two F₆:₇ recombinant inbred line (RIL) populations. These RILs were developed from the crosses CA2156 × JG62 (susceptible × resistant) and CA2139 × JG62 (resistant × resistant), and were sown in a field infected with fusarium wilt race 0 in Beja (Tunisia) over 2 years. A1:1 resistant to susceptible ratio was found in the RIL population from the CA2156 × JG62 cross, indicating that a single gene with two alleles controlled resistance. In the second RIL population (CA2139 × JG62) a 3:1 resistant to susceptible ratio indicated that two genes were present and that either gene was sufficient to confer resistance. Linkage analysis showed a RAPD marker, OPJ20600, linked to resistance in both RIL populations, which is present in the resistant parent JG62.