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Titel:Charakterisierung der Tryptophan-abhängigen Pigmentsynthese am Modellorganismus Ustilago maydis und Identifizierung von Wirtsspezifitätsfaktoren in den phytopathogenen Brandpilzen Ustilago maydis und Sporisorium reilianum
Autor:Zuther, Katja
Weitere Beteiligte: Bölker, Michael (Prof. Dr.)
Veröffentlicht:2009
URI:https://archiv.ub.uni-marburg.de/diss/z2010/0484
URN: urn:nbn:de:hebis:04-z2010-04844
DOI: https://doi.org/10.17192/z2010.0484
DDC: Biowissenschaften, Biologie
Titel(trans.):Characterisation of the Tryptophan dependent pigment synthesis in the model organism Ustilago maydis and Identification of host specificity factors in the phytopathogenic smut fungi Ustilago maydis and Sporisorium reilianum
Publikationsdatum:2010-09-22
Lizenz:https://rightsstatements.org/vocab/InC-NC/1.0/

Dokument

Schlagwörter:
Phytoalexin, Pigments, Malassezia furfur, Hirse, Wirtsspezifität, Ustilago zeae, Pigmente, Sporisorium reilianum, Tryptophanderivate, Mais, Phytoalexine, Tryptophanstoffwechsel, Head smut disease, Tryptophan, Pityriasis versicolor, Host specificity, Sprorisorium reilianum

Zusammenfassung:
Im ersten Teil der vorliegenden Arbeit wurde am Modellorganismus Ustilago maydis der Tryptophan-abhängige Biosyntheseweg untersucht, der zur Generierung von Indolpigmenten mit biologischen Effekten führt, die erstmals für den Pityriasis versicolor-assoziierten Hautpilz Malassezia furfur nachgewiesen worden waren. Mittels eines revers-genetischen Ansatzes konnte gezeigt werden, dass die Tryptophan-Aminotransferase Tam1 das einzige für die Pigmentbiosynthese verantwortliche Enzym repräsentiert. Die aufgereinigte heterolog exprimierte Tam1 aus U. maydis katalysierte die Umsetzung von Tryptophan und Phenylpyruvat zu Indolpyruvat und Phenylalanin. Ein im Vorfeld der Arbeit durchgeführter vorwärts-genetischer Ansatz hatte zur Identifikation von Mutanten geführt, die unfähig waren, in Anwesenheit von Tryptophan Pigmente zu bilden. Diese Mutanten waren in dem Sulfitreduktase-Gen sir1 betroffen. In vitro-Experimente mit aufgereinigter Tam1 zeigten, dass auch 2-Oxo-4-Methylthiobutanoat, die 2-Oxosäure von Methionin, als Substrat für Tam1 dienen kann, was die Tryptophan-Deaminierung an den Schwefelmetabolismus koppelt. Die Experimente zeigten, dass Indolpyruvat spontan, entweder alleine oder in Kombination mit Tryptophan, in wässriger Umgebung zu einer Vielzahl von farbigen Substanzen reagierte, unter denen sich auch mindestens vier der für M. furfur beschriebenen Substanzen mit biologischen Effekten befanden. Experimente an Keratinozyten-Zellkultur zeigten, dass Indolpyruvatabhängige biologische Effekte wie vermindertes Proliferationsvermögen über den humanen Arylhydrocarbonrezeptor vermittelt werden könnten. Im zweiten Teil der Arbeit wurden mittels Genomvergleich Gene aus Sporisorium reilianum identifiziert, die an der Ausprägung der Wirtsspezifität beteiligt sein könnten. S. reilianum existiert in zwei Varietäten (SRM und SRH), wovon SRM bevorzugt auf Mais und SRH ausschließlich auf Hirse zur Sporenbildung kommt. Um Faktoren aus S. reilianum zu identifizieren, die die Virulenz auf Mais unterstützen und gleichzeitig die Virulenz auf anderen Wirtspflanzen schwächen, wurden die in den Genomen der Maispathogene SRM und U. maydis vorkommenden Gene nach Abwesenheit in den Genomen verwandter Pathogenen anderer Wirtspflanzen untersucht. Dabei wurde ein Gen aus S. reilianum identifiziert, sr13864, das vermutlich für ein sekretiertes Protein kodiert. Mittels RT-PCR wurde gezeigt, dass sr13864 während der biotrophen Phase exprimiert wird. U. maydis-Stämme, in denen das homologe Gen von sr13864, um00823, deletiert wurde, zeigten reduzierte Virulenz auf Mais. Während Expression von um00823 in der um00823-Deletionsmutante den Virulenzdefekt vollständig komplementieren konnte, konnte heterologe Expression von sr13864 in der um00823-Deletionsmutante die reduzierte Virulenz nur teilweise aufheben. Expression von sr13864 in SRH führte zu einer erhöhten Abwehrreaktion auf Hirse in Form von roten Läsionen entlang der Leitbündel, die wahrscheinlich auf die Bildung von 3-Desoyanthocyanidin-Phytoalexin zurückzuführen sind. Damit scheint sr13864 ein Wirtsspezifitätsfaktor mit gleichzeitiger Virulenzfunktion für die Infektion von Mais und Avirulenzfunktion für die Infektion von Hirse zu sein.

Summary:
In the first part of the thesis the model organism Ustilaog maydis was used to study the tryptophan dependent biosynthesis that leads to the generation of indole pigments with biological effects. These indole pigments were originally found in the fungus Malassezia furfur associated with the human skin disease pityriasis versicolor. Using a reverse genetic approach it could be shown that the tryptophan aminotransferase Tam1 represents the only enzyme responsible for pigment biosynthesis. Heterologously expressed and purified Tam1 from U. maydis catalyzed the conversion from tryptophan and phenyl pyruvate into indole pyruvate and phenyl alanine, respectively. Using a forward genetic approach mutants were identified that were incapable of producing pigments in the presence of tryptophan. These mutants were affected in the sulphite reductase gene sir1. In vitro experiments with purified Tam1 showed that also 2-oxo-methylthiobutanoate, the oxoacid of methionine can serve as a substrate for Tam1, thereby coupling tryptophan deamination to sulfur metabolism. The experiments show that indole pyruvate in an aqueous environment spontaneously reacts (either alone or in combination with tryptophan) to form numerous colourful compounds. At least four compounds could be detected that had been described for M. furfur and seem to be associated with biological effects. Experiments with human keratinocytes showed that indole pyruvate dependent biological effects such as impaired proliferation could be directed via the human aryl hydrocarbon receptor. In the second part of the thesis candidate genes for host specificity from Sporisorium reilianum were identified using genome comparison. S. reilianum exists in two varieties (SRM and SRS) of which SRM produces spores in the flower of maize and SRS produces spores in the flower of sorghum. To identify factors in S. reilianum that support virulence on maize and at the same time reduce virulence on other host plants, genes present in the genomes of the maize pathogens SRM and U. maydis were tested for absence in the genomes of related pathogens that have different hosts. Thereby one gene from S. reilianum was identified, sr13864, that presumably codes for a secreted protein. Using RT-PCR it was shown that sr13864 is expressed during the biotrophic phase. U. maydis strains in which the homolog of sr13864, um00823, was deleted, showed reduced virulence on maize. While expression of um00823 in the um00823 deletion mutant could fully complement the virulence defect, heterologous eexpression of sr13864 in the um00823 deletion mutant could only partially restore the virulence. Expression of sr13864 in SRS led to an enhanced defence reaction on sorghum along the vacular bundles as could be deduced from the production of red spots. These red spots most likely represent 3-Deoxyanthocyanidine phytoalexins. Thus, sr13864 seems to be a host specificity factor with a dual function as virulence factor for maize and avirulence factor for sorgum.

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