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Investigation of Host Specificity Mechanisms of Sporisorium Reilianum in Maize and Sorghum

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Investigation of Host Specificity Mechanisms of Sporisorium Reilianum in Maize and Sorghum Investigation of host specificity mechanisms of Sporisorium reilianum in maize and sorghum Von der Fakultät für Mathematik, Informatik und Naturwissenschaften der RWTH Aachen University zur Erlangung des akademischen Grades einer Doktorin der Naturwissenschaften genehmigte Dissertation vorgelegt von M. Sc. Alana Poloni aus Caxias do Sul, Brazil Berichter: Prof. Dr. Jan Schirawski Prof. Dr. Alan Slusarenko Tag der mündlichen Prüfung: 19.02.2015 Diese Dissertation ist auf den Internetseiten der Hochschulbibliothek online verfügbar. “…nonconformity is a great asset to a scientist. We must be curious to see if what we see is what we seem to see. We must analyse it, open it up, turn it over, look underneath it, and look behind." James G. Horsfall, 1975 Content Summary ................................................................................................................................................ 1 Zusammenfassung ................................................................................................................................ 2 1. Introduction ........................................................................................................................................ 3 1.1 Host specificity in plant pathogens ................................................................................................ 3 1.2 Host resistance and plant immunity .............................................................................................. 4 1.3 Downstream responses in plant defense ...................................................................................... 7 1.3.1 Reactive oxygen species ........................................................................................................ 7 1.3.2 Reinforcement of the cell wall................................................................................................. 8 1.3.3 Phytohormones ...................................................................................................................... 8 1.3.4 Antimicrobial compounds ....................................................................................................... 9 1.3.5 Pathogenesis related proteins (PRs) ...................................................................................... 9 1.4 The smut fungi ............................................................................................................................. 11 1.4.1 Head smut of maize and sorghum is caused by Sporisorium reilianum .............................. 12 Aims of this study .............................................................................................................................. 14 2. Materials and Methods .................................................................................................................... 15 2.1 Materials ...................................................................................................................................... 15 2.1.1 Plant material ........................................................................................................................ 15 2.1.2 Fungal strains ....................................................................................................................... 15 2.2 Methods ....................................................................................................................................... 17 2.2.1 Plant inoculation ................................................................................................................... 17 2.2.2 Microscopic characterization of plant infection ..................................................................... 18 2.2.3 Genomic DNA extraction from S. reilianum .......................................................................... 18 2.2.4 Genomic DNA extraction from infected maize and sorghum ............................................... 18 2.2.5 RNA extraction from infected maize and sorghum ............................................................... 19 2.2.6 RNA quantification and cDNA synthesis .............................................................................. 20 2.2.7 Polymerase Chain Reaction (PCR) ...................................................................................... 20 2.2.8 Gel electrophoresis ............................................................................................................... 21 2.2.9 Quantitative RT-PCR ............................................................................................................ 21 2.2.10 Staining of plant material for H2O2 , callose, lignin and plant cell death ............................. 22 2.2.11 Sample preparation and transmission electron microscopy (TEM).................................... 23 2.2.12 Sample preparation for scanning electron microscopy (SEM) ........................................... 25 2.2.13 Laser microdissection (LMD) .............................................................................................. 25 2.2.14 Cell separation by mechanical methods ............................................................................. 26 2.2.15 Transcriptome analysis ....................................................................................................... 26 3. Results .............................................................................................................................................. 28 3.1 The fungal side of the interaction: colonization of maize and sorghum by S. reilianum ............. 28 3.1.1 S. reilianum f. sp. reilianum successfully infects sorghum, while S. reilianum f. sp. zeae does not trigger symptoms in sorghum inflorescences ................................................................ 28 3.1.2 SRZ causes head smut in maize cobs and tassels, while SRS only induces phyllody in cobs as the strongest symptom ............................................................................................................. 31 3.1.3 Sorghum stems are colonized only by SRS ......................................................................... 32 3.1.4 Both SRS and SRZ can reach maize stems ........................................................................ 35 3.1.5 Quantification of fungal genomic DNA reveals contrasting proliferation behavior of SRS and SRZ in maize and sorghum ........................................................................................................... 36 3.1.6 Transmission electron microscopy shows differences in SRS and SRZ cell wall thickness in maize ............................................................................................................................................. 38 3.1.7 No differences in apressorium morphology were observed between SRS and SRZ on maize, but differences appear on sorghum ................................................................................... 42 3.1.8 SRS seems to form a more tight physical interaction with sorghum than SRZ ................... 44 3.2 The plant side: defense responses in maize and sorghum ......................................................... 45 3.2.1 H2O2 accumulates at penetrating hyphae of SRZ in sorghum ............................................ 45 3.2.2 Sorghum cells are reinforced against SRS colonization ...................................................... 46 3.2.3 No plant cell death is observed when plants are inoculated with S. reilianum ..................... 50 3.2.4 Phytoalexin deposition is a late response during the interaction of sorghum with SRZ ...... 51 3.2.5 Expression analysis of marker genes reveals differential plant responses for SRS and SRZ in maize and sorghum ................................................................................................................... 52 3.2.6 Maize transcriptome shows mild differences in gene regulation between SRZ and SRS- infected plants ................................................................................................................................ 55 3.2.7 Transcriptome analysis in sorghum shows strong differences in gene regulation between SRZ- and SRS-infected plants ...................................................................................................... 62 3.3 Cell separation in sorghum: establishment of methods to isolate ............................................... 72 mesophyll and vascular bundle cells in sorghum leaves ................................................................. 72 3.3.1 Isolation of vascular bundles and mesophyll cells by laser microdissection (LMD) ............. 72 3.3.2 Vascular bundles and mesophyll cells isolation by mechanical and enzymatic methods ... 74 4. Discussion........................................................................................................................................ 77 4.1 The Sporisorium reilianum-maize pathosystem .......................................................................... 77 4.2 The Sporisorium reilianum-sorghum pathosystem ...................................................................... 82 5. References ....................................................................................................................................... 90 6. Appendix ........................................................................................................................................ 105 6.1 List of Abbreviations .................................................................................................................
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