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Horizontal gene transfer The transfer of genetic information from one genome to another. 1 Conjugation = transfer of DNA from one organism to another by means of a plasmid. 2 Transformation = uptake of free DNA from the environment. 3 Transduction = transfer of DNA from one organism to another by a bacteriophage. 4 5 An organism into which genetic information from a different organism has been incorporated as a stable part of its genome is a transgenic organism. Aeqorea victoria green fluorescent protein (GFP) ANDi 6 Orthology Paralogy Xenology 7 Horizontal gene transfer is suspected when there is a discrepancy between gene phylogeny and species phylogeny, in particular when the tree reflects geographical proximity rather than phylogenetic affinity. 8 Patchy phylogenetic distribution Aedes aegypti 9 nucleus cytoplasm (CuZn-SOD) SOD = superoxide dismutase cyanobacteria + chloroplast (Fe-SOD) bacteria + mitochondria (Mn-SOD)10 Leiognathus daura Photobacterium leiognathi (CuZnSOD) 11 12 Crown Gall Disease 13 Agrobacterium tumefaciens 14 Plant Gene 15 16 P M P elements & hybrid dysgenesis Margaret Kidwell 17 P elements were not found in any D. melanogaster strains collected before 1950, and collections made subsequently showed increasing frequencies of P with decreasing age. North and South America Europe, Africa, Middle East Australia, Far East 18 Two hypotheses: 1. Most D. melanogaster strains in nature carry P elements, but they tend to lose them in the laboratory. 2. P elements were recently introduced into D. melanogaster populations in nature. 19 The recent acquisition hypothesis is supported by: 1. P strains that have been monitored in the lab for ~15 y were never observed to lose P. 2. There is a geographical cline in the temporal appearance of P in nature. 3. There is evidence for horizontal gene transfer. 20 21 Species tree P-element tree 22 Drosophila melanogaster 23 Drosophila willistoni 24 Proctolaelaps regalis 25 Conditions for horizontal Pelement transfer: (1) two Drosophila females from the donor and the recipient species must lay eggs in proximity to one another (2) the recipient egg must be less than 3 hours old (< 512 cells) (3) the germline of the recipient embryo must incorporate a complete copy of P Marilyn A. Houck (4) the receiving embryo must survive the biting injuries 26 type-C virogene 27 28 29 Informational genes: Genes involved in replication, transcription, reverse transcription, and translation. Operational genes: All others. 30 The complexity hypothesis informational operational 31 Genes move within the genome and between genomes. Genetic “Mutatis mutandis.” 32 Promiscuous DNA 33 Einat Hazkani-Covo et al. Mitochondrial-sequence invasions into the nuclear genome 34 Numts* (nuclear mitochondrial DNA sequences) are a type of promiscuous DNA, i.e., nuclear sequences of mitochondrial origin. *pronounced “new mights” 35 The transfer of functional genes from the mitochondria to the nucleus is thought to have has stopped in evolution after the emergence of animals (~1,000 MYA). 36 The reason is thought to be the differences between the nuclear and mitochondrial genetic codes. 37 The transfer of nonfunctional pieces of mitochondrial genetic information continues to this day. Numts have been found so far in 83 eukaryote species. 38 Most species whose genomes have been completely sequenced contain very few numts. Saccharomyces cerevisiae Caenorhabditis elegans Drosophila melanogaster Plasmodium falciparum 17 numts 3 numts 3 numts 3 numts 39 In the human genome we find ~1,000 numts total length = 831 Kb ~0.02% of the nuclear genome 40 Numts: Evolution’s misplaced witnesses 41 42 Junk DNA Domestic Imported 43