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Species trees and gene trees Fundamentals of molecular phylogenies Genetic distances • Comparison of 50 amino acids (fig 9.6) Constructing trees 1 2 3 4 1 0 2 2 0 3 8 8 0 4 22 22 22 0 5 22 22 22 14 5 0 Molecular clocks • DNA fixes substitutions at a roughly clock-like rate Distantly related Related pairs of species have a large number of substitutions in the cytochome c gene Number of substitutions in cytochrome c gene Different types of DNA segments evolve at different rates Time since last common ancestor (millions of years) Coalescing genes • Gene trees and species trees Species tree Gene tree Coalescing genes • Genetic loci have their own genealogy 10 haploid individuals pass down their copies of alleles at a particular gene locus Alleles coalesce Fig 9.2, 9.3 Coalescing genes • Coalescence of alleles varies Coalescence of alleles in human genes (Templeton, 2006) Gene trees versus species trees: rates of evolution 1. Monomorphic founding population Constant rate of evolution A B C 2. Monomorphic founding population Unequal rates of evolution A A B C B C A B C Gene trees versus species trees A A B C B C A B 2. Polymorphic founding population Constant rate of evolution C Gene trees versus species trees Three biological factors that can produce true genealogical discordance between a gene tree and a species tree “Non-canonical” trees • Horizontal gene transfer • Example: The ACR3 gene • • • an arsenic permease involved in the cell’s response to arsenic toxicity effluxes arsenite from the cell The gene is often found on plasmids Mega - http://www.megasoftware.net/ Silva - http://www.arb-silva.de/ “Non-canonical” trees • What might explain this result? Clymene dolphins Median-joining network of cytochrome b haplotypes of Stenella clymene (red circles), S. coeruleoalba (green circles) and S. longirostris (blue circles). Circle size is proportional to the number of individuals exhibiting the correspondent haplotype. Length of lines separating the haplotypes is proportional to the number of mutational steps. White circles indicate missing, intermediate haplotypes. (Amaral 2014) Non-canonical trees Stenella coeruleoalba Stenella clymene Stenella longirostris Stenella clymene “Non-canonical” trees • Gene duplication Orthologs Paralogs Pelagic species Substrate spawner Most extant marine teleosts spawn highly hydrated pelagic eggs, due to differential proteolysis of vitellogenin (Vtg)-derived yolk proteins. This tree represents sequences of vitellogenin sequences extracted from species that spawn in the pelagic zone and one that spawns against substrate