小麦对白粉菌的抗性和感病性的诱导

 用小麦白粉菌(Erysiphe graminis f.sp.tritici)的无毒和有毒菌株,以3种接种顺序:1.先接无毒菌株,间隔一定时间后,再接有毒菌株;2.无毒和有毒菌株同时接;3.和"1"相反,接种具有不同抗性基因的小麦品种的幼苗或叶段,以研究小麦对白粉菌的诱导抗性和诱导感病性。结果表明:先接无毒菌株诱导的抗性最强;后接无毒菌株或无毒、有毒菌株同时接,当诱导菌(inducer)的密度很高时,也可以诱导出抗性,但较微弱;诱导的抗性随诱导菌的密度增高而增强;一经诱导,抗性可在被诱导部位维持到第6天;所使用的3种接种顺序都可以诱导小麦对白粉菌产生感病性,但较之诱导的抗性弱得多;在抗病性的诱导中,存在着小麦品种与诱导菌,或诱导菌与挑战菌(challenger)之间的相互作用;一个无毒菌株可以通过诱导而获得在抗病品种上生长的毒性,但当缺少有毒菌株存在时,获得毒性即消失;没有发现所诱导的抗性具有系统传导性。     

Altered Phenotypic Response to Peronospora parasitica in Brassica juncea Seedlings Following Prior Inoculation with an Avirulent or Virulent Isolate of Albugo candida

A study was made of the biological interactions between an isolate of Peronospora parasitica compatible with Brassica juncea and two isolates of Albugo candida either incompatible or compatible with the host species. Prior inoculation with the incompatible isolate of A. candida induced resistance to subsequently inoculated P. parasitica. The degree of resistance was proportional to the zoosporangia concentration of the incompatible isolate and induced resistance was more marked in the cotyledon receiving the inducing inoculum compared to the opposite cotyledon and subsequently emerging true leaves that had not been pre-inoculated. Induction of resistance was also observed if the incompatible isolate of A. candida and P. parasitica were co-inoculated simultaneously. However, the effect was greater the longer the interval between inoculations, up to a period of 4 days. When the incompatible isolate of A. candida was inoculated 4h after P. parasitica, there was no marked effect on resistance to the latter. In contrast, prior inoculation with the compatible isolate of A. candida increased susceptibility to P. parasitica inoculated subsequently. However, pre- or co-inoculation with P. parasitica suppressed the development of the compatible isolate of A. candida. A spectrum of responses was observed when one cotyledon was inoculated simultaneously with both the incompatible and compatible isolates of A. candida and followed subsequently with P. parasitica after different time intervals. In such combinations, a transition was observed in the host response to P. parasitica from induced resistance/reduced susceptibility, which increased up to 24h following a simultaneous inoculation with incompatible + compatible isolates of A. candida to an almost neutral reaction after 72h to induced susceptibility after 96h. This range of altered responses appeared to reflect the outcome of the differing kinetics and counter-effects of resistance and susceptibility induction.     

Reaction of three cotton (Gossypium hirsutum) cultivars to single and mixed isolates of Xanthomonas campestris pv. malvacearum

Cotyledons of resistant and susceptible cotton ( Gossypium hirsutum ) cultivars were inoculated artificially with four isolates of Xanthomonas campestris pv. malvacearum from Africa (H V1, HV3, H V7 and Sudan) and USA race 18. Each isolate was used as single inoculum and in combinations of mixed inoculum. Disease reactions were graded 15 days after inoculation on a scale of I (immunity) to 10 (fully susceptible). Inoculation with HVI alone produced the highest disease grade of all inoculum treatments. When HVI was included in a mixed inoculum, disease grades were greatly reduced compared to HVI alone. Disease grades were highest for the inoculum mixture of the most virulent isolates HVI and race 18, but were still reduced compared to those from HVI alone. The efficiency of selection when screening cotton germplasm for bacterial blight resistance may be reduced if the highly virulent HVI isolate is included in an inoculum mixture composed of non-virulent isolates.     

Induction of Systemic Resistance to Albugo candida in Brassica juncea by Pre- or Coinoculation with an Incompatible Isolate

ABSTRACT In an investigation of the interaction between two isolates of Albugo candida that were compatible (CO) and incompatible (IN) on a Brassica juncea accession, the IN isolate induced both local and systemic protection of cotyledons and true leaves against the CO isolate. The extent of the protection was proportional to the zoosporangia concentration used in the inducing (IN) inoculation. Protection was greatest locally on cotyledons and least on true leaves (the most remote tissue from the point of the inducing inoculation). Protection occurred when the two isolates were inoculated together but was greater when the interval between the IN and CO isolate inoculations was longer. The IN isolate induced only slight protection when it was inoculated after the CO isolate. No induced susceptibility to the IN isolate occurred with any treatment. There was some evidence of competition between CO and IN zoospores for infection sites (stomata). The occurrence of systemic protection and changes detected in phenylalanine ammonia lyase and total soluble peroxidase activities in inoculated cotyledons, particularly after the inducing (IN) inoculation, suggested that host-mediated factors also may be involved in the interaction between the two isolates.     

The importance of non‐penetrated papillae formation in the resistance response of triticale to powdery mildew (Blumeria graminis)

Triticale is the intergeneric hybrid between wheat and rye. With the expansion of the triticale growing area, powdery mildew has emerged and become a significant disease on this new host. Recent research demonstrated that this ‘new’ powdery mildew on triticale has emerged through a host range expansion of powdery mildew of wheat. Moreover, isolates sampled from triticale still infect their previous host, wheat, but isolates sampled from wheat hardly infect triticale. Race‐specific and adult‐plant resistance have been identified in triticale cultivars. The main objective of this study was to characterize the cellular basis of powdery mildew resistance in triticale. Commonalities with resistance responses in other cereals such as wheat, barley and oat are discussed. A detailed comparative histological study of various resistance responses during cross‐inoculation of either virulent or avirulent wheat and triticale isolates on both hosts was carried out. The present data provide evidence that for incompatible interactions, the formation of non‐penetrated papillae is the predominant resistance response, while the hypersensitive response (HR) acts as a second line of defence, to cut the fungus off from nutrients, if penetration resistance fails. It is not clear yet what causes the slower growth and reduced colony size of triticale isolates when inoculated on wheat. Possibly, post‐penetration resistance mechanisms, other than HR, are switched on during these (semi‐) compatible interactions. Molecular studies on gene expression and gene function of defence‐related genes might reveal further insights into the genetic basis of these resistance responses.     

Effects of preinoculation with an avirulent isolate of Pyricularia grisea on infection and development of leaf blast lesions caused by virulent isolates on near-isogenic lines of Sasanishiki rice

Eight near-isogenic Sasanishiki rice lines with different genes for complete resistance to rice blast were inoculated with an avirulent isolate 72 h before inoculation with a virulent isolate of Pyricularia grisea to clarify the mechanisms of induced resistance in the leaf blades. Subsequent lesions on the leaf blades were classified as brown spots (b type), as observed on Sasanishiki BL no. 8 with resistance gene Pii, or no symptoms (HR type), as observed on Sasanishiki BL no. 4 with the gene Piz-t and on the six other lines. Lesion expansion was significantly reduced in Sasanishiki BL no. 8 compared with that in Sasanishiki BL no. 4 when the leaf blades were preinoculated with a high concentration of a conidial suspension of the avirulent isolate. Moreover, after preinoculation with the conidial suspension of the avirulent isolate in silicon rings on the leaf blades, induced resistance was expressed only in areas close to the inoculation sites. These resistant areas were larger in Sasanishiki BL no. 8 than in BL no. 4. Hyphal growth was markedly inhibited in the epidermal cells of Sasanishiki BL no. 8, whereas inhibition was weak in those of Sasanishiki BL no. 4. In the epidermal cells of leaf blades of Sasanishiki BL no. 8 preinoculated with the avirulent isolate, the frequency of hyphal penetration of the virulent isolate in the presence of host cell browning decreased, as did the frequency of invading hyphae after inoculation with virulent isolates. The results indicate that induced resistance may play a role in the suppression of lesion development in the Sasanishiki near-isogenic lines and that the lines differ in the extent of suppression.     

大豆和大豆疫霉菌共培养体系的构建

 本研究旨在建立一个适于分析大豆遭受大豆疫霉菌侵染后基因表达研究的互作体系。在比较了15个携带不同抗病基因的大豆基因型的组织培养情况和7个大豆疫霉菌分离物及其游动孢子单孢系的毒力变化的基础上,构建了大豆悬浮细胞和大豆疫霉菌游动孢子的共培养体系,进而分析了共培养过程中大豆细胞的活力和防御基因表达情况。结果表明,不同亲和力菌株的游动孢子引起的大豆细胞活力变化十分相似;非亲和菌株的游动孢子可诱导寄主防御基因的表达发生变化。此系统为深入开展大豆抗性机制研究提供了良好的平台。     

Induction of a defense response in strawberry mediated by an avirulent strain of <Emphasis Type="Italic">Colletotrichum</Emphasis>

In the strawberry crop area of Tucumán (north-west Argentina) the three species of Colletotrichum causing anthracnose disease (C. acutatum, C. fragariae and C. gloeosporioides) were detected. Among all isolates characterized, one of them identified as C. acutatum (M11) and another as C. fragariae (F7) were selected due to their conspicuous interaction with the strawberry cultivar Pájaro. Whereas isolate M11 produced a strong compatible interaction in cv. Pájaro with clear disease symptoms (DSR = 5.0), the isolate F7 brought about a typical incompatible interaction (DSR = 1.0). When plants of cv. Pájaro were inoculated with F7 prior to the inoculation with M11, the former avirulent strain prevented the growth of the latter virulent pathogen. Experimental evidence indicated that the time elapsed between the first inoculation with the avirulent pathogen and the second inoculation with the virulent one was crucial to inhibit the growth of the latter. The growth of F7 on the plant without provoking damage and the fact that there was no in vitro antagonistic effect between the pathogens, suggests that the avirulent strain triggers a plant defensive response against M11. The defense response was further confirmed by the detection of an early oxidative burst occurring within 4 h after the first inoculation and by the observation of anatomical changes associated with defense mechanisms that lasted 50 days after the inoculation with F7. Results obtained support the hypothesis that the plant resistance against the virulent strain M11 is elicited by one or more diffusible(s) compound(s) produced by the avirulent strain F7.     

Programmed cell death pathways induced by early plant‐virus infection are determined by isolate virulence and stage of infection

Programmed cell death (PCD) pathways caused by Turnip mosaic virus (TuMV) infection before symptom appearance were studied by light microscopy and electrolyte leakage following sap inoculation of Brassica carinata (Ethiopian mustard) TZ‐SMN‐44‐6 plants. Leaf responses to inoculation with avirulent (TuMV‐avir) and virulent (TuMV‐vir) isolates, and mock‐inoculation, were compared at 2, 20 and 52 h after inoculation (hai). The phenotypes induced were localized resistance (TuMV‐avir) and systemic susceptibility (TuMV‐vir). No visible TuMV symptoms were recorded in any inoculated plants during the 2–52 hai sampling period, but appeared as chlorotic spots in inoculated leaves at 5 days after inoculation. With TuMV‐vir alone, they were followed by systemic infection (mosaic). Dead cell number, deformation, percentage area and percentage integrated intensity, and conductivity of electrolyte leakage data, were analysed to examine their possible roles in stimulating cell death pathways. At 2 hai, dead cell number and percentage area were significantly greater for TuMV‐avir than TuMV‐vir infection or mock‐inoculation. Overall, isolate TuMV‐vir caused significantly greater cell deformation than TuMV‐avir, whereas wounding by mock‐inoculation had negligible effects. By 52 hai, isolate TuMV‐avir caused significantly greater electrolyte leakage than isolate TuMV‐vir or mock‐inoculation. This suggests both isolates triggered morphological changes consistent with apoptotic‐like PCD and necrosis‐like PCD that depended upon isolate virulence and stage of infection, respectively. These findings highlight how quantification of dead cell deformation and electrolyte leakage offer a new understanding of compatible and incompatible plant responses to early virus infection in plants.     

Protection of Citrus Rootstocks Against Phytophthora spp. with a Hypovirulent Isolate of Phytophthora nicotianae

ABSTRACT Phytophthora root rot of citrus in Florida is caused by Phytophthora nicotianae and P. palmivora. A naturally occurring isolate of P. nicotianae (Pn117) was characterized as hypovirulent on citrus roots. Pn117 infected and colonized fibrous roots, but caused significantly less disease than the virulent isolates P. nicotianae Pn198 and P. palmivora Pp99. Coincident inoculation of rootstock seedlings of Cleopatra mandarin (Citrus reticulata) or Swingle citrumelo (C. paradisi x Poncirus trifoliata) with the hypovirulent Pn117 and the virulent isolates Pn198 and Pp99 did not reduce the severity of disease caused by the virulent Phytophthora spp. When either rootstock was inoculated with the hypovirulent Pn117 for 3 days prior to inoculation with virulent isolates, preinoculated seedlings had significantly less disease and greater root weight compared with seedlings inoculated with the virulent isolates alone. Recovery of the different colony types of Phytophthora spp. from roots of sweet orange (C. sinensis) or Swingle citrumelo was evaluated on semiselective medium after sequential inoculations with the hypovirulent Pn117 and virulent Pp99. Pn117 was isolated from roots at the same level as the Pp99 at 3 days post inoculation. Preinoculation of Pn117 for 3 days followed by inoculation with Pp99 resulted in greater recovery of the hypovirulent isolate and lower recovery of the virulent compared with coincident inoculation.     

稻梨孢菌与其他寄主梨孢菌在水稻植株上的交互作用

作者研究了1个稻梨孢菌株与4个稻以外寄主梨孢菌株在混合接种和间隔接种条件下,在水稻植株上的交互作用。结果表明:弱致病菌与强致病菌间交互作用较强,非致病菌与强致病菌间较弱。弱致病菌可明显降低强致病菌的致病性,非致病菌对强致病菌的作用,多数组合表现为减轻病害,但有少数组合表现为促进发病。弱致病菌先接种;2天后接种强致病菌,比两者混合接种具有更强烈的互作效应,病斑数减少达35.7％—38.1％。先接种菌的孢子浓度对后接种菌的致病性也有影响。     

A qualitative host-pathogen interaction in the Theobroma cacao-Moniliophthora perniciosa pathosystem

The aim of this study was to test whether resistance of clones of Theobroma cacao (cocoa) varied between isolates of Moniliophthora (formerly Crinipellis ) perniciosa , the cause of witches' broom disease. Developing buds of vegetatively propagated T. cacao grown in greenhouses in the UK were inoculated with 16 000 spores of M. perniciosa per meristem in water, under conditions where water condensed on the inoculated shoot for at least 12 h after inoculation. The proportion of successful inoculations varied between clones and was inversely correlated with time to symptom production or broom formation. A specific interaction was demonstrated among three single-spore isolates of M. perniciosa and the clone Scavina 6 (SCA 6) and a variety of susceptible clones. Isolates Castenhal-I and APC3 were equally likely to infect SCA 6 and the other clones, but isolate Gran Couva A9 never infected SCA 6, although it was as virulent on the other clones. The interaction was maintained when the wetness period was extended to 70 h. Offspring of SCA 6 × Amelonado matings were all susceptible to both Castenhal-I and GC-A5, with no evidence of greater variability in susceptibility to GC-A5 than Castanhal-I. This suggests recessive inheritance of a single homozygous factor conferring resistance to GC-A5, from SCA 6. The progenies were slightly more susceptible to Castanhal-I than GC-A5. The implications for managing the disease are discussed.     

稻梨孢菌与其他寄主梨孢菌在水稻植株上的交互作用

作者研究了1个稻梨孢菌株与4个稻以外寄主梨孢菌株在混合接种和间隔接种条件下,在水稻植株上的交互作用。结果表明:弱致病菌与强致病菌间交互作用较强,非致病菌与强致病菌间较弱。弱致病菌可明显降低强致病菌的致病性,非致病菌对强致病菌的作用,多数组合表现为减轻病害,但有少数组合表现为促进发病。弱致病菌先接种;2天后接种强致病菌,比两者混合接种具有更强烈的互作效应,病斑数减少达35.7%-38.1%。先接种菌的孢子浓度对后接种菌的致病性也有影响。     

Plant defence reactions against fusarium wilt in chickpea induced by incompatible race 0 of Fusarium oxysporum f.sp. ciceris and nonhost isolates of F. oxysporum

Germinated seeds of 'kabuli' chickpea cv. ICCV 4 were inoculated with a conidial suspension of the incompatible race 0 of Fusarium oxysporum f.sp. ciceris (Foc) or of nonhost F. oxysporum resistance 'inducers', and 3 days later were challenged by root dip with a conidial suspension of highly virulent Foc race 5. Prior inoculation with inducers delayed the onset of symptoms and/or significantly reduced the final amount of fusarium wilt caused by race 5. However, the extent of disease suppression varied with the nature of the inducing agent; the nonhost isolates of F. oxysporum were more effective at disease suppression than the incompatible Foc race 0. Inoculation with the inducers gave rise to synthesis of maackiain and medicarpin phytoalexins in inoculated seedlings; these did not accumulate in plant tissues but were released into the inoculum suspension. Inoculation with inducers also resulted in accumulation of chitinase, β-1,3-glucanase and peroxidase activities in plant roots. These defence-related responses were induced more consistently and intensely by nonhost isolates of F. oxysporum than by incompatible Foc race 0. The phytoalexins and, to a lesser extent, the antifungal hydrolases, were also induced after challenge inoculation with Foc race 5. However, in this case the defence responses were induced in both preinduced and noninduced plants infected by the pathogen. It is concluded that the suppression of fusarium wilt in this study possibly involved an inhibitory effect on the pathogen of preinduced plant defences, rather than an increase in the expression of defence mechanisms of preinduced plants following a subsequent challenge inoculation.     

Colonization of roots of subterranean clover cultivars by virulent and avirulent races of Phytophthora clandestina

The colonization of the roots of four cultivars of subterranean clover by isolates representing four races of Phytophthora clandestina was studied. There was a highly significant race × cultivar interaction in the growth of inoculated tap roots and the degree of colonization of roots by the pathogen. While all races were able to infect the roots of all cultivars tested, roots of the susceptible cultivars were colonized more rapidly and extensively than those of the resistant cultivars. In compatible combinations, fungal colonization extended for a few centimetres in the tap root and lateral roots in the moderately susceptible cultivars Trikkala and Meteora, or throughout the whole root system leading to the death of the host in the very susceptible cultivar Woogenellup. In contrast, limited fungal colonization of the tap root and lack of extension of the fungus into lateral roots was typical of incompatible combinations. In all cultivars, lateral roots were as susceptible to infection as tap roots. The number of lateral roots of Woogenellup was significantly reduced by infection. However, neither the rate of lateral root formation nor the total number of lateral roots of Seaton Park, Meteora and Trikkala was reduced by infection with virulent or avirulent races of the pathogen.     

Japanese pear black spot and apple alternaria blotch

The pathogenicity and taxonomy of 15 isolates of Alternaria spp. from pear and apple were compared. Only isolates from Asian pear ( Pyrus pyrifolia ) from Italy and Korea were virulent on leaves and young fruits of the susceptible Asian pear cv. Nijisseiki. Their conidial morphology was typical of A. gaisen (= A. kikuchiana ). Only isolates of A. mali from USA were virulent on susceptible American apple cvs Indo and Red Gold. No virulence was demonstrated in any isolate/host combination among isolates from stem infections of Asian and European pear ( Pyrus communis ), leaf spots of European apple, and ripe fruit rots of Chinese pear ( Pyrus ussuriensis ), European pear and apple. These non-virulent isolates could be readily distinguished from A. gaisen and A. mali by their pattern of branching of conidial chains, the branching associated with A. alternata sensu stricto being most common among non-virulent isolates. This limited survey implies that A. gaisen is only virulent to Asian pear and the toxigenic form of A. mali to certain American apple cultivars; also that A. gaisen is not established outside eastern Asia or the toxigenic form of A. mali outside eastern Asia and parts of USA.     

Differences in virulence of <Emphasis Type="Italic">Phytophthora capsici</Emphasis> isolates from a worldwide collection on host fruits

Phytophthora capsici causes root, crown, and fruit rot of vegetable and tropical hosts. Cucumber, zucchini, tomato, and pepper fruits were inoculated using 6-mm-diameter agar plugs of P. capsici, incubated in clear plastic boxes at room temperature (25 ± 2°C and 100% relative humidity), and virulence was estimated by measuring the lesion diameter, pathogen growth diameter, and pathogen sporulation density three (cucumber, zucchini) or four (tomato, pepper) days later. When isolates were grouped by genetic cluster, significant differences in virulence were observed on cucumber and zucchini, with isolates belonging to genetic cluster five causing larger lesions than isolates from genetic cluster six. On tomato, no significant differences were observed for isolates grouped by genetic cluster, but isolates from vegetable crops were generally more virulent than isolates from tropical hosts. Isolates from fabaceous hosts sporulated better on cucumber fruits than isolates from solanaceous hosts. Isolates from vegetable hosts sporulated better on zucchini than isolates from tropical hosts. No significant differences in lesion diameter were noted on pepper when isolates were grouped by host family of origin or genetic cluster, but differences in pathogen sporulation were apparent by host family. Our findings suggest that isolate characteristics such as host family of origin and genetic cluster membership may be used to guide initial isolate selection for cucurbit fruit resistance screening. Final isolate selection should incorporate the phenotypic and genetic diversity of P. capsici, including isolates with differing virulence to the host organ of interest.     

Protective Effects Against Erwinia amylovora Induced by hrp Mutants in the Whole Plant are Only Partially Mimicked in Cultivated Cells

A virulent strain of Erwinia amylovora, the causal agent of fire blight of Maloideae, and two of its non-virulent hrp mutants (a secretory and a regulatory mutant) were inoculated into apple cell suspensions either alone or in mixed inoculations. In single inoculations, death of 4- to 5-day-old apple cells occurred only when the concentration of the virulent strain of E. amylovora reached a threshold inoculum concentration of 10⁴CFUml^–1, while high concentrations of the hrp mutants were unable to kill apple cells. When mixed inoculated with the virulent parental strain, both hrp mutants protected apple cells from death caused by the virulent strain. The protective effect was associated with a decrease in the population level of the virulent strain and it was dependent on the non-virulent to virulent inoculum concentration suggesting a competition between the non-virulent mutant and the virulent strain. However, no differential protective ability between the two types of mutants could be noticed, contrary to previous results obtained with apple seedlings or apple flowers in which the regulatory mutant was significantly more effective than the secretory mutant. In conclusion, inoculation of apple cell cultures with E. amylovora does not seem to be a model suitable for investigating mechanisms leading to protection.     

Characterization of pathotypes among isolates of Xanthomonas axonopodis pv. manihotis in Colombia

Cassava bacterial blight, caused by Xanthomonas axonopodis pv. manihotis ( Xam ) is a destructive disease occurring in most cassava growing-areas. Although Colombian isolates of Xam differ in DNA polymorphism and pathogenicity, no suitable host differentials have been identified to demonstrate physiological specialization. A set of 26 Xam isolates from three edaphoclimatic zones (ECZs) in Colombia was selected for inoculation on a set of 17 potential cassava differentials. Leaf inoculation and stem puncture were used in order to detect possible specific interactions between cultivars and isolates. Cultivar × isolate interaction was highly significant ( P  < 0·001) after stem inoculation, but not after leaf inoculation. The stem inoculation technique was selected as a method for resistance screening of cassava cultivars for bacterial blight resistance. A highly significant interaction was also detected when cultivar behaviour was rated as area under the disease progress curve (AUDPC) after stem inoculation. Different pathotypes were defined among the 26 isolates and differential cultivars were proposed to define the pathotypic composition of Xam populations in three ECZs in Colombia. The results should help to improve selection of sources of resistance to cassava bacterial blight.     

Evidence for a Minor Gene-for-Minor Gene Interaction Explaining Nonhypersensitive Polygenic Partial Disease Resistance

ABSTRACT Partial resistance is a quantitative type of resistance that, by definition of Parlevliet, is not based on hypersensitivity. It is largely pathotype nonspecific, although some minor isolate-specific responses have been reported. In order to elucidate the isolate specificity of individual genes for partial resistance, three barley recombinant inbred line mapping populations were analyzed for resistance to the leaf rust fungus Puccinia hordei. The mapping populations were inoculated with one isolate avirulent and two isolates virulent to resistance gene Rph7g. Six significant quantitative trait loci (QTLs) were detected. Of these, two (Rphq3 and Rphq11) were detected with only the avirulent isolate (1.2.1.) and one (Rphq18) only with both virulent isolates (CO-04 and 28.1). The effectiveness of these QTLs was tested with 14 isolates, using a tester set of genotypes containing alleles for resistance or susceptibility for these QTLs. QTL Rphq18 was effective to only two isolates, CO-04 and 28.1, whereas Rphq3 and Rphq11 were ineffective to CO-04 and 28.1 but effective to all other isolates, except one. This resulted in a significant Person's differential interaction, which is a hallmark of a gene-for-gene interaction. The minor gene-for-minor gene interaction is not based on hypersensitivity and there is no evidence that the resistance is based on genes belonging to the nucleotide-binding leucine-rich repeat class.