Phytophthora infestans , the agent of potato and tomato late blight disease, produces a 10-kD extracellular protein, INF1 elicitin. INF1 induces a hypersensitive response in a restricted number of plants, particularly those of the genus Nicotiana. In virulence assays with different P. infestans isolates, five Nicotiana species displayed resistance responses. In all of the interactions, after inoculation with P. infestans zoospores, penetration of an epidermal cell was observed, followed by localized necrosis typical of a hypersensitive response. To determine whether INF1 functions as an avirulence factor in these interactions, we adopted a gene-silencing strategy to inhibit INF1 production. Several transformants deficient in inf1 mRNA and INF1 protein were obtained. These strains remained pathogenic on host plants. However, in contrast to the wild-type and control transformant strains, INF1-deficient strains induced disease lesions when inoculated on N. benthamiana . These results demonstrate that the elicitin INF1 functions as an avirulence factor in the interaction between N. benthamiana and P. infestans . INTRODUCTIONMicrobial plant pathogens often exhibit high degrees of specialization and can only infect a limited number of plant species (Agrios, 1988). Pathogen specialization results when a complex set of preformed and induced mechanisms is put into motion to defend a plant against invading pathogens. In some interactions, preformed physical barriers and antimicrobial compounds in the plant help to ward off pathogens (Osbourn, 1996a(Osbourn, , 1996b. In other interactions, perception by the plant of signal molecules, namely, elicitors, produced by the avirulent pathogen leads to the induction of effective defense responses, including a programmed cell death response termed the hypersensitive response (HR) (Lamb et al., 1989; Dixon and Harrison, 1990; Ebel and Scheel, 1992; Baker et al., 1997;Morel and Dangl, 1997). This model has been genetically defined by Flor's gene-for-gene hypothesis (Flor, 1956(Flor, , 1971. According to this hypothesis, a resistance reaction is determined by the simultaneous expression of a pathogen avirulence ( Avr ) gene with the corresponding plant resistance ( R ) gene (Staskawicz et al., 1995).In recent years, the gene-for-gene hypothesis has received tremendous experimental support through the identification and functional characterization of both Avr and R genes. A number of Avr genes from fungi, bacteria, and viruses were shown to encode specific elicitor proteins. This was demonstrated directly by infiltration of Avr proteins into plant leaves or indirectly by expression of Avr genes in plant cells containing the corresponding R gene (Culver and Dawson, 1991; de Wit, 1995; Alfano and Collmer, 1996;Knogge, 1996; Bonas and van den Ackerveken, 1997; van den Ackerveken and Bonas, 1997). Elicitor treatment or Avr gene expression triggers the HR and related defense responses in plants that mimic the response induced by avirulent pathogens (Hahlbrock et al., 1995; Hammond-Kosa...
Most species of the genus Phytophthora produce 10-kDa extracellular protein elicitors, collectively termed elicitins. Elicitins induce hypersensitive response in a restricted number of plants, particularly in the genus Nicotiana within the Solanaceae family. A cDNA encoding INF1, the major secreted elicitin of Phytophthora infestans, a pathogen of solanaceous plants, was isolated and characterized. The expression of the corresponding inf1 gene during the disease cycle of P. infestans was analyzed. inf1 was shown to be expressed in mycelium grown in various culture media, whereas it was not expressed in sporangiospores, zoospores, cysts, and germinating cysts. In planta, during infection of potato, particularly during the biotrophic stage, expression of inf1 was down-regulated compared to in vitro. The highest levels of expression of inf1 were observed in in vitro grown mycelium and in late stages of infection when profuse sporulation and leaf necrosis occur. The potential role of INF1 as an elicitor in interactions between P. infestans and Solanum species was investigated. Nineteen lines, representing nine solanaceous species with various levels of resistance to P. infestans, were tested for response to an Escherichia coli expressed INF1. Within the genus Solanum, resistance to P. infestans did not appear to be mediated by a defense response elicited by INF1. However, INF1 recognition could be a component of nonhost resistance of tobacco to P. infestans.
Transformation of the diploid oomycete plant pathogen Phytophthora infestans with antisense, sense, and promoter-less constructs of the coding sequence of the elicitin gene inf1 resulted in transcriptional silencing of both the transgenes and the endogenous gene. Since heterokaryons obtained by somatic fusion of an inf1-silenced transgenic strain and a wild-type strain displayed stable gene silencing, inf1 silencing is dominant and acts in trans. Inf1 remained silenced in nontransgenic homokaryotic progeny from the silenced heterokaryons, thereby demonstrating that the presence of transgenes is not essential for maintaining the silenced status of the endogenous inf1 gene. These findings support a model reminiscent of paramutation and involving a trans-acting factor that is capable of transferring a silencing signal between nuclei.
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