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Evolutionary Biology Concepts. “What is behind is not important!”- or is it? Molecular Evolution Phylogenetic Inference. Evolution. Change in living organisms via reproduction. "Change over time"-Kentucky School Boards. Levels of Evolution. Species Population gene frequencies
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Evolutionary Biology Concepts “What is behind is not important!”- or is it? Molecular Evolution Phylogenetic Inference Chuck Staben
Evolution Change in living organisms via reproduction "Change over time"-Kentucky School Boards Chuck Staben
Levels of Evolution • Species • Population • gene frequencies • Organismal • genomic • Molecular Chuck Staben
Branching Descent Evolutionary Tree Family Tree Chuck Staben
Phylogeny Branching diagram showing the ancestral relations among species. “Tree of Life” History of evolutionary change FRAMEWORK for INFERENCE Chuck Staben
Phylogenetic Inference Mom's hair color? Which form of reproduction? Chuck Staben
Mom-Inferred vs Real • “Blonde” Phenotype • Recombination • Male/hair color • Multigenic hair color? • Potential Asexual Propagation • Date of vasectomy? courtesy, Ms. J. Rae Staben Chuck Staben
Inferring the Framework • How do we describe phylogenies? • How do we infer phylogenies? Chuck Staben
Classical Phylogeny Molecular Phylogeny Inheritance RNA Protein Function DNA Chuck Staben
A B C D E F Node Internode Phylogenetic Trees Sister Taxa Terminal Taxa Ancestor Root Chuck Staben
A B C D E F Paraphyletic Group Clade Monophyletic Group More Trees Chuck Staben
A B C D E F Trees-3 Polyphyletic Group Chuck Staben
Add Root Rooted vs Unrooted Trees Chuck Staben
A B C D E F Extinction Chuck Staben
Speciation • Poorly understood • “…the mystery of mysteries…”-Darwin • Reproductive isolation/divergence Chuck Staben
Population Genetic Forces Hardy-Weinberg Paradigm p+q=1 p2 + 2pq + q2 =1 • Natural Selection (fitness) • Drift (homozygosity by chance) • much greater in small populations • Mutation/Recombination (variation) • Migration • homogenizes gene pools Chuck Staben
DNA, protein sequence change Rate=1 change/6 aa sites per 108 yrs Rate=0.16 x 10-9 yrs (normal ~ 1 per 10-9 yrs per site) Chuck Staben
Multiple Changes/No Change ..CCU AUA GGG.. ..CCC AUA GGG.. ..CCC AUG GGG.. ..CCC AUG GGC.. ..CCU AUG GGC.. ..CCU AUA GGC.. 5 mutations 1 DNA change 0 amino acid changes (net) Underestimate Evolution Chuck Staben
Mechanisms of DNA Sequence Change Neutral Drift vs Natural Selection • For a 1000 base gene, 41000 sequences! Selection (Jukes) Neutral (Kimura) Chuck Staben
Rate varies Gene-to-Gene Chuck Staben
Rate varies Site-to-Site Coding>Silent??? Chuck Staben
Constraints on “Silent” Changes • Codon Biases-translation rates • Transcription elongation rates • polymerase ‘pause’ sites • “Silent” regulatory elements • select for or against presence/absence • Overall genome structure Chuck Staben
Neutralism in Eukaryotes vs Prokaryotes-“Slightly deleterious mutations” Models Most non-coding sites are neutral? Coding/noncoding can be flexible? Reconsider evolutionary mechanisms? Chuck Staben
Evolutionary Genetic Forces Hardy-Weinberg Paradigm p+q=1 p2 + 2pq + q2 =1 • Natural Selection (fitness) • Drift (homozygosity by chance) • much greater in small populations • Mutation/Recombination (variation) • Migration • homogenizes gene pools Genome Recombination? Chuck Staben
DNA, Protein Similarity • Similarity by common descent • phylogenetic • Similarity by convergence • functional importance • Similarity by chance • random variation not limitless • particular problem in wide divergence Chuck Staben
Homology-similar by common descent Chuck Staben
Inferring Trees and Ancestors CCCAGG CCCAAG-> CCCAAG CCCAAA-> CCTAAA CCTAAA-> CCTAAC MANY traps, problems Chuck Staben
Paralogs Homology, Orthology, Paralogy Orthologs Chuck Staben
Paralogy Trap Chuck Staben
Improper Inference Man is a mouse, not a rat! Chuck Staben
Convergence Globin Common Ancestor Convergence Leghemoglobin Chuck Staben
Our Goals • Infer Phylogeny • Optimality criteria • Algorithm • Phylogenetic inference • (interesting ones) Chuck Staben
Watch Out “The danger of generating incorrect results is inherently greater in computational phylogenetics than in many other fields of science.” “…the limiting factor in phylogenetic analysis is not so much in the facility of software applicaition as in the conceptual understanding of what the software is doing with the data.” Chuck Staben