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Molecular and Genomic Evolution Getting at the Gene Pool Figure 26.1 What is Molecular Evolution? • a change in nucleotide &/or amino acid sequences over time – molecular phylogenetics reconstructs lineages based on molecular differences Determining and Comparing Sequences • PCR (Polymerase Chain Reaction) – rapid amplification of DNA from many samples • automated sequencing methods – rapid determination of sequences from PCR Sequence Alignment Figure 26.2 Determining and Comparing Sequences • homologous DNA sequences are compared by alignment – sequences of closely related groups have fewer differences Determining and Comparing Sequences • sequence changes accumulate at different rates in different parts of the genome – regions that encode functional products change relatively slowly • synonymous changes are most common – non-coding regions may change rapidly Figure 26.3 Determining and Comparing Sequences • good historical evidence combined with good molecular evidence give the rate of change of a sequence • some polypeptides have relatively constant amino acid substitution rates over time Determining and Comparing Sequences • a sequence with a constant rate of change can be used as a “molecular clock” – cytochrome c is in the electron transport chain in the mitochondrion of all eukaryotes Figure 26.4 amino acid substitution rate of cytochrome c Figure 26.5 The Origin of New Protein Functions • protein function can change – lysozyme, an antibacterial enzyme, is found in most animals. • tears, saliva, milk, egg whites – some mammals use lysozyme in foregut fermentation, a type of digestion • ruminants • langurs Langurs Figure 26.6 The Origin of New Protein Functions • protein function can change – foregut fermentation arose in langurs and ruminants separately • each descended from non-fermenting recent ancestors – langur and ruminant lysozymes share changes that protect them from digestion Table 26.1 The Origin of New Protein Functions • protein function can change – langur and ruminant lysozymes share changes that protect them from digestion – the hoazin, a foregut fermenting bird, makes a lysozyme with similar changes Figure 26.6 The Origin of New Protein Functions • protein function can change – hoazin, langurs, ruminants all share distinctive amino acid substitutions in the same enzyme – therefore… • they shared a recent common ancestor? • homoplasy, such as convergent evolution, is identified by comparison with patterns of homology. The Origin of New Genes • Gene duplication yields new genes – duplicate genes may change together • rRNA gene tandem arrays share changes so that members retain the same sequence – duplicate genes often change independently • one copy of the gene is required to produce a normal product • a duplicate copy may change its function by mutation The Origin of New Genes • Gene duplication yields new genes – duplicate genes are homologs • paralogs are homologs in the same genome • orthologs are homologs in different genomes – duplication of genes, chromosomes, or entire genomes can occur The Origin of New Protein Functions • duplicated proteins can change without harm to the organism – myoglobin, -globin and -globin gene families arose following gene duplication • each family experienced later duplications Figure 26.9 Figure 26.7 Figure 26.8 C-value Inflation A Relevant Mystery • The C-value paradox – more-complex organisms have more DNA per genome than less-complex organisms – more-complex organisms have more genes than less-complex organisms – more-complex organisms have much higher proportions of non-coding DNA – the non-coding DNA has no known function Molecular Phylogenetics • different molecules change at different rates over time – rapidly changing molecules • useful for recently diverged groups • slow changing molecules for groups that diverged long ago Molecular Phylogenetics • different molecules change sequences at different rates over time – the gene for the small ribosomal RNA subunit changes very slowly – serves as one of the bases for the three domain classification of life Figure 26.10 Molecular phylogenetics reconstructs the history of gene evolution Molecular Phylogenetics • The Big Bird story – Moas • large flightless birds • extinct for ~1000 years • shared New Zealand with kiwis • presumed to share a more recent common ancestor with kiwis than with other large flightless birds on other continents