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What are Transposons? • “Selfish DNA” • Interspersed repeats • “move” in the genome Bacterial Insertion Sequences • Transpositions inactivate essential genes, killing the host and the IS element it carries • Some transposed sequences enter nonessential regions of the genome allowing regions of genes to survive Transposase • Required for transposition of IS element to new site • Target site direct repeat sequence are immediately adjacent to both ends of the inserted elements Transposase Functions 3’ of Is element to 5’ ends of cut donor DNA McClintock’s Discovery • Spontaneous mutations in maize affect production of enzymes required for pigment 2 Movable Elements 1. Activator elements Encodes mRNA for transposase protein. -recognizes the terminal repeats and catalyzes transposition to a new DNA site 2. Dissociation elements -deletion mutant of Activator that has lost the necessary sequences to move on its own Results of Transposons • Can result in an increase in the number of a transposon if it occurs during S phase of cell cycle • after a DNA transposon leaves a gene, the resulting gap will probably not be repaired correctly • Transposition in germ cells to their new sites is passed on to succeeding generations Phytophthora infestans • Oomycete – aka “water mold” – fungus-like eukaryotic organism “pseudofungus” • Cause of potato blight, which brought about the Irish Potato Famine Genome organization • Transposon-poor regions o Genes needed for survival • Transposon-rich regions o Genes used for pathogenesis • 74% of genome is transposon-rich • Most transposons in genome are inactive Plant Immune System • Pathogen-associated Molecular Patterns (PAMPs) causes response of PAMP-triggered Immunity (PTI) • Pathogens produce effectors to suppress PTI • Plants produce R proteins to suppress effectors • High selective pressure on effector genes leads to rapid evolution RNA Silencing • P. infestans uses sRNA to restrict the activity of transposons Effector Silencing • Proximity to transposons leads to silencing of effector genes • Caused by formation of heterochromatin at and around transposons • Heterochromatin spreads ~300 to 600 bases from transposon • 563 RXLR effectors • 35 within 300bp of transposons • 106 within 600bp • 283 within 2kb