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Molecular. Evolution and. Phylogenetics. Masatoshi Nei. Sudhir Kumar ... 7 Phylogenetic Inference: Máximum Parsimony Me

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Molecular Evolution and Phylogenetics

Masatoshi Nei Sudhir Kumar

Contents

Numerical Examples xiii 1 Molecular Basisof Evolution 3 1.1. Evolutionary Tree of Life 3 1.2. Mechanism of Evolution 4 1.3. Structure and Function of Genes 5 1.4. Mutational Changes of DNA Sequences 9 1.5. CodonUsage 11 2 Evolutionary Changeof Arrimo Acid Sequences 17 2.1. Amino Acid Differences and Proportion of Different Amino Acids 17 2.2. Poisson Correction (PC) and Gamma Distances 19 2.3. Bootstrap Variances and Covariances 25 2.4. Amino Acid Substitution Matrix 27 2.5. Mutation Rate and Substitution Rate 29 3 Evolutionary Change of DNA Sequences 33 3.1. Nucleotide Differences Between Sequences 33 3.2. Estimation of the Number of Nucleotide Substitutions 35 3.3. Gamma Distances 43 3.4. Numerical Estimation of Evolutionary Distances 45 3.5. Alignment of Nucleotide Sequences 46 3.6. Handling of Sequence Gaps in the Estimation of Evolutionary Distances 49 4 Synonymous and Nonsynonymous Nucleotide Substitutions 51 4.1. Evolutionary Pathway Methods 52 4.2. Methods Based on Kimura's 2-Parameter Model 62 4.3. Nucleotide Substitutions at Different Codon Positions 67 4.4. Likelihood Methods with Codon Substitution Models 69

CONTENTS

5 Phylogenetic Trees 73 5.1. Types of Phylogenetic Trees 73 5.2. Topological Differences 81 5.3. Tree-Building Methods 83 6 Phylogenetic Inference: Distance Methods

6.1. 6.2. 6.3. 6.4. 6.5.

87

UPGMA 87 Least Squares (LS) Methods 92 Mínimum Evolution (ME) Method 99 Neighbor Joining (NJ) Method 103 Distance Measures to Be Used for Phylogenetic Reconstruction 111

7 Phylogenetic Inference: Máximum Parsimony Methods 115 7.1. Finding Máximum Parsimony (MP) Trees 116 7.2. Strategies of Searching for MP Trees 122 7.3. Consensus Trees 130 7.4. Estimation of Branch Lengths 131 7.5. Weighted Parsimony 133 7.6. MP Methods for Protein Data 138 7.7. Shared Derived Characters 140 8 Phylogenetic Inference: Máximum Likelihood Methods

8.1. 8.2. 8.3. 8.4. 8.5.

9 Accuracies and Statistical Tests of Phylogenetic Trees

9.1. 9.2. 9.3. 9.4. 9.5.

147

Computational Procedure of ML Methods 147 Models ofNucleotideSubstitution 152 Protein Likelihood Methods 159 Theoretical Foundation of ML Methods 162 Parameter Estimation for a Given Topology 163 165

Optimization Principie and Topological Errors 165 Interior Branch Tests 168 Bootstrap Tests 171 Tests of Topological Differences 175 Advantages and Disadvantages of Different Tree-Building Methods 178

10 Molecular Clocks and Línearized Trees 10.1. Molecular Clock Hypothesis 10.2. Relative Rate Tests 191 10.3. Phylogenetic Tests 196 10.4. Linearized Trees 203

187 187

11 Ancestral Nucleotide and AminoAcid Sequences 11.1. Inference of Ancestral Sequences: Parsimony Approach 207 11.2. Inference of Ancestral Sequences: Bayesian Approach 208

207

CONTENTS

11.3. Synonymous and Nonsynonymous Substitutions in Ancestral Branches 216 11.4. Convergent and Parallel Evolution 221 12 Genetic Polymorphism and Evolution

231

12.1. Evolutionary Significance of Genetic Polymorphism 231 12.2. Analysis of Alíele Frequency Data 233 12.3. Genetic Variation in Subdivided Populations 12.4. Genetic Variation for Many Loci 244 12.5. DNA Polymorphism 250 12.6. Statistical Tests for Detecting Selection 258 13 Population Trees from Genetic Markers

236

265

13.1. Genetic Distance for Alíele Frequency Data 265 13.2. Analysis of DNA Sequences by Restriction Enzymes 275 13.3. Analysis of RAPO Data 285 14 Perspectives

291

14.1. Statistical Methods 291 14.2. Cenóme Projects 292 14.3. Molecular Biology and Evolution 294 Appendices

A. B. C. D.

Mathematical Symbols and Notations 297 Geological Timescale 299 Geological Events in the Cenozoic and Mesozoic Eras 301 Organismal Evolution Based on the Fossil Record 303

References Index

329

305

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