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When highly resistant wheat (Triticum aestivum L.) varieties are infected by an avirulent race of the stem rust fungus (Puccinia graminis Pers. f. sp. tritici Erics. and E. Henn.), penetrated host cells undergo rapid necrotization. This hypersensitive cell death is correlated with cellular lignification which efficiently restricts further fungal growth. Three competitive inhibitors of phenylalanine ammonia-lyase, the first enzyme of the general phenylpropanoid pathway and, thus, of lignin biosynthesis, namely a-aminooxyacetate, a-aminooxy-B-phenylpropionic acid, and (1-amino-2-phenylethyl)phosphonic acid, and two highly specific irreversible suicide inhibitors of the lignification-specific enzyme cinnamyl-alcohol dehydrogenase, namely N(O-aminophenyl)sulfinamoyl-tertiobutyl acetate and N(O-hydroxyphenyl)sulfinamoyltertiobutyl acetate, were applied to genetically resistant wheat plants prior to inoculation with stem rust. Treatment with any of these inhibitors decreased the frequency of lignified necrotic host cells and concomitantly led to increased fungal growth. The cinnamyl-alcohol dehydrogenase inhibitors were generally more effective than the phenylalanine ammonia-lyase inhibitors, occasionally allowing some sporulation to occur on the resistant wheat leaves. These results clearly point to a causal relationship between the formation of lignin precursors and the resistance of wheat to stem rust.The lignin content ofhigher plants has long been recognized as an important factor in the resistance response against potential pathogens. Lignin is extremely resistant to microbial degradation and thus constitutes one of the most effective barriers against pathogenic invasion (25). In addition to the role of lignin as a preformed resistance factor, induced lignification has been proposed as an active resistance mechanism of plants to fungi (15). However, it is still a matter of debate whether active lignification processes are causally involved in resistance (25).In the family Gramineae, active lignification appears to be of special importance in induced resistance mechanisms (28), possibly related to the near absence of phytoalexins in this family. In spite of intensive searches for infection-induced During the time of this study, B. M. M. received a Graduiertenstipendium des Landes Nordrhein Westfalen. This work was supported in part by the Deutsche Forschungsgemeinschaft.
When highly resistant wheat (Triticum aestivum L.) varieties are infected by an avirulent race of the stem rust fungus (Puccinia graminis Pers. f. sp. tritici Erics. and E. Henn.), penetrated host cells undergo rapid necrotization. This hypersensitive cell death is correlated with cellular lignification which efficiently restricts further fungal growth. Three competitive inhibitors of phenylalanine ammonia-lyase, the first enzyme of the general phenylpropanoid pathway and, thus, of lignin biosynthesis, namely a-aminooxyacetate, a-aminooxy-B-phenylpropionic acid, and (1-amino-2-phenylethyl)phosphonic acid, and two highly specific irreversible suicide inhibitors of the lignification-specific enzyme cinnamyl-alcohol dehydrogenase, namely N(O-aminophenyl)sulfinamoyl-tertiobutyl acetate and N(O-hydroxyphenyl)sulfinamoyltertiobutyl acetate, were applied to genetically resistant wheat plants prior to inoculation with stem rust. Treatment with any of these inhibitors decreased the frequency of lignified necrotic host cells and concomitantly led to increased fungal growth. The cinnamyl-alcohol dehydrogenase inhibitors were generally more effective than the phenylalanine ammonia-lyase inhibitors, occasionally allowing some sporulation to occur on the resistant wheat leaves. These results clearly point to a causal relationship between the formation of lignin precursors and the resistance of wheat to stem rust.The lignin content ofhigher plants has long been recognized as an important factor in the resistance response against potential pathogens. Lignin is extremely resistant to microbial degradation and thus constitutes one of the most effective barriers against pathogenic invasion (25). In addition to the role of lignin as a preformed resistance factor, induced lignification has been proposed as an active resistance mechanism of plants to fungi (15). However, it is still a matter of debate whether active lignification processes are causally involved in resistance (25).In the family Gramineae, active lignification appears to be of special importance in induced resistance mechanisms (28), possibly related to the near absence of phytoalexins in this family. In spite of intensive searches for infection-induced During the time of this study, B. M. M. received a Graduiertenstipendium des Landes Nordrhein Westfalen. This work was supported in part by the Deutsche Forschungsgemeinschaft.
Abstract. In cowpea (Vigna s&ensis L.) leaves the pathogenic rust fungus Uromyces vignae and the non-pathogens U. appendiculatus and U. viciae-fabae developed similarly to give rise to more than 80% haustorial mother cells. Whereas U. vignae was able to sporulate, the nonpathogens were stopped either after formation of some haustoria (U. appendiculatus) or immediately after formation of haustorial mother cells (U. viciae-fabae).Approximately 30% of the cells in contact with haustorial mother cells of the two non-pathogens showed autofluorescence and deposition of phloroglucinol/HC1-positive material. The early defence reactions of V. sinensis include induction of phenylalanine ammonia-lyase (PAL, EC 4.3.1.5) and extracellular peroxidase (POD, EC 1.11.1.7) activity as early as 10h and 24h after inoculation, respectively. Probing Western blots with heterologous monospecific anti-PAL serum showed that pathogenesis-induced increases in enzyme activity are the result of de novo synthesis. Native polyacrylamide gel electrophoresis revealed the specific induction of two extracellular acidic POD forms in cowpea inoculated with the non-pathogens. Both PAL and apoplasmic POD activities were highest in U. viciae-fabae-, intermediate in U. appendiculatus-and low in U. vignae-inoculated or talcum-treated control plants. The timing of increasing PAL and extracellular POD activities in relation to deposition of lignin or lignin-like material in mesophyll cell walls indicates the involvement of lignification in the early defence of V. sinensis against biotrophic fungal parasites. Analysis of the substrate specificity of the inducible POD forms, separated and partially purified by chromatofocusing, showed that apoplasmic PODs preferentially oxidize the naturally occurring substrate coniferyl alcohol. These results support the assumption that POD-mediated lignification is involved in the expression of non-host resistance of cowpea to rust fungi. * To whom correspondence should be addressed Abbreviations: IWF = intercellular washing fluid; PAL = phenylalanine ammonia-lyase; p.i. = post inoculation ; POD = peroxidase; SDS-PAGE = sodium dodecyl sulfate-polyacrylamide gel electrophoresis
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