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Evolutionary Genomics and Proteomics
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Evolutionary Genomics and Proteomics
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Evolutionary
Genomics and
Proteomics
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\,' Mark Pagel
, Andrew Pomiankowski
Editors
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Sinauer Associates, Inc. Publishers
Sunderland, Massachusetts 01375
ADout the \,;over:
The cover illustration shows part of an undirected phylogenetic tree for a short
protein sequence. The proteins are represented as strings of colored spheres
(amino acids), and proteins connected by a bar differ from each other by a single
amino acid. The image alludes both to the evolutionary relationships revealed
by genomics and proteomics and to the structure of biological networks. At the
upper right, a ribosome translates a freshly transcribed mRNA into a protein.
Art and cover layout by Joanne Delphia.
Evolutionary Genomics and Proteomics
Copyright 2008 by Sinauer Associates, Inc.
All rights reserved. This book may not be reproduced in whole or in part with-
out permission from the publisher.
For information, address:
Sinauer Associates
23 Plumtree Road
Sunderland, MA 01375 U.S.A.
FAX:413-549-1118
Email: publish@sinauer.com
Internet: www.sinauer.cOIn
Library of Congress Cataloging-in-Publication Data
Evolutionary genomics and proteomics/edited by Mark Pagel,
Andrew Pomiankowski.
p.;cm.
Includes bibliographicall'eferences and index.
ISBN-13: 978-0-87893-654-0 (paperbound)
ISBN-13: 978-0-87893-655-7 (hardcover)
1. Evolutionary genetics. 2. Genomics. 3.. Proteomics.
[DNLM: 1. Evolution, Molecular-Revie~. 2.
Genome-genetics-Review. 3. Proteome-genetics-Review. QU 475
E956 2008] 1.Pagel, Mark D. II. Pomiankowski, Andrew. QH390.E98
2008 572.8'38-dc22
2007035125
Printed in China
5 4 3 2 1
Table of Contents
Preface XIII
Contributors xv
CHAPTER
1 The Organismal Prospect
Mark Pagel and Andrew P011l;ankowski
Evolutionary Genetics in the Time of Genomics
The Organism in the Genome 3
Complexity and Regulation 4
The Origins of Novelty 6
Robustness and Evolvability 8
Genome Immunity 9
Identifiying Selective Change 9
What If Evolutionary Genomic and Proteomic Scientists
Succeed? 10
CHAPTER 2 Evolutionary Systems Biology II
Eugene V. Koon;n and Yur; T. Wolf
Systems Biology: The New Big Science of "Omes" 11
Systems Biology in the Light of Evolution: The Quest for the Link
between Genomic and Phenotypic Evolution 13
Correlations between Variables Characterizing Genome Evolution
and Functioning 16
The Multidimensional Space of Gmics 20
Glimpses of Biology behind the Correlations 22
Some Generalizations: Evolutionary Systems Biology-
a Nascent Field in Turmoil 24
Acknowledgments 25
CHAPTER 3 The Origin of New Genes 27
ClllIallzhllFan, J. J. Emerson, and Many"an Long
Introduction 27
Recent Discoveries of New Genes 27
Mechanisms to Generate New Genes 30
Evolutionary Forces for New Gene Retention 32
The Location and Movement of New Genes 33
Inferring the Functionality of New Genes 33
General Methods to Detect New Genes 35
Early filldings 35
Comparative molecular cytogenetic allalyses 35
Computatiollal genomic analysis 35
Comparative Genomic Hybridization to Detect New Genes 36
Challellges ill using array-based CGH ill new gene studies 39
Outlooks and Perspectives 42
CHAPTER 4 Lateral Gene Transfer 45
w. Ford Doolittle, Camilla L. Nesbg, Eric Bapteste, and Olga Zhaxybayeva
Introduction 45
Defining terms 45
A Very Brief History of Lateral Genomics 46
Mechanisms of Lateral Gene Transfer 46
Transduction 47
Conjugation 48
Transformation 50
Detecting LGT 51
Compositional methods 51
Unusual phyletic similarity patterns 51
Phylogenetic incongruence 52
PatcilY (anomalous) gene distribution among phyla and species 52
Strain-specific genes (patchy distribution within species) 53
Physical methods
57
Ever-Present Alternatives 58
Quantifying LGT 61
GenOllle-centric approaclles 61
Gene-centric methods 61
An LGT-Resistant Core 63
The complexity hypothesis 64
The tiny universal core 65
Do core genes have a common history? 66
LGT and Microevolution 67
HR and the biological species concept (jor bacteria) 67
Paralogs and xenologs 68
Adaptive LGT 69
Neutral LGT 69
LGT and Macroevolution 70
Eukaryotes, the Other Iceberg 72
LGT from inside: endosymbiotic gem transfer 72
LGT from outside: prokaryotes to eukaryotes 76
Eukaryotes to eukanjotes 77
Eukaryotes to prokaryotes 78
Reuniting Process and Pattern 78
CHAPTER 5 Evolution of Genomic Expression 81
Bemardo Lei/lOS,Christian R. Landry, PierreFontanillas, SlIsan C. p, Reml,
Rob KlIlathinal, KyleM. Brown, and Daniel L. Hartl
Introduction 81
The Complex Regulation of Genomic Expression 81
Classical Transcriptional Regulation in
Cis and TrailS 83
Promoters, enhallcers, repressors, transcription factors, and regulatory proteins 83
Epigenetic Regulation and Chromatin Modifications 86
DNA methylation 87
Histone modifications 88
Chromosome territories and nue/ear architecture 89
Post-Transcriptional Regulation 91
Measuring Attributes of Genomic Expression with Experimental
and Computational Tools 93
Experimental approaches 93
Coplputational approaches 99
The Evolution of Genomic Expression: What Do We Know? 103
Genomic expression and morphological evolution 104
Molecular evolution of regulatory sequences 105
Stabilizing selectiOlJ,positive selection, and neutrality
of gene expression levels 107
GelJomic attributes and rates of gelJe expression evolution 109
Inheritance of gene expression levels: Regulatory variation in
cis and trans 111
GelJotype-by-environment interactions, sex-biased genes, and epistasis 115
Gene Regulatory Networks, Subnetworks, and Modules 117
Conclusion 118
CHAPTER
6 The Evolution of Proteome Complexity
and Diversity 119
Laszlo Pntthy
Introduction 119
Evolution of Protein Complexity 120
Evolution of Proteome Complexity 123
Multidomain Proteome Complexity 126
Proteome Interaction Networks 127
Multidomain Proteins and Organismic Complexity 133
Factors Favoring the Formation of Multidomain Proteins 136
Conclusion 137
Acknowledgments 139
CHAPTER
7 Genomic Redundancy and Dispensability 141
Laurence D. Hurst and CsabaPal
Introd uction 141
Are "Dispensable" Genes Really Dispensable? 143
Nonessential genes are typically under strong purijtjing selection 146
By What Mechanisms Are Nontrivially Dispensable Genes Dispensable? 148
Network architecture and distributed robustness 151
How do duplicates provide backup? 153
Backup by paralogs or by distributed robustness: whicll is more important? 156
Why Did Dispensability Evolve? 157
General conditions for the evolution of redundancy 158
Specific models for dispensability support the "side-consequence" llypot/zesis 160
Discussion 164
CHAPTER 8 Genome Defense 167
Christopher B. Schaefer,Mary Grace Gall, and Timothy H. Bestor
Introduction 167
Transposable elemellts 167
Transposons compromise genomic stability 169
Transposable elements are a greater threat to sexual hosts 170
Genome defense systems must continuously evolve 171
1rll1l~J1U~Uflt'J llllU wt: C.ljlHft:JlUIt U) 6t:.twrm:"
Mechanisms of Genome Defense 173
RNA interference 173
Cosuppression 176
Qllelling 176
Meiotic silencing by unpaired DNA 177
DNA eliminatioll 177
Trallscriptional gene silencing 178
Systemic silellcing 180
Cytosine Methylation 181
Restrictionmodijication in bacteria 181
5-Methylcytosine and heritable silencing in eukaryotes 181
DNA methylation in vertebrates 183
Methylation ill insects 185
Methylation ill plants 185
Methylation in
N. crassa 186
Editing as a Form of Defensive Mutagenesis 187
Restriction of retroviruses by G-to-A hypermutatioll 187
A-to-I editing of SINE elements 189
Conclusion 189
CHAPTER
9 Sex-Biased Genomic Expression 193
Brian Oliver
Introduction 193
One Species, Two Optimal Genomes 194
Y Chromosomes 195
X Chromosomes 198
Comparing Mammalian, Drosophila, and C. elegans X Chromosomes 201
Moving Forward 204
CHAPTER 10 Sex Chromosome Origins and Evolution 207
D. Charlesworth
Introduction: The Diversity of Sex Chromosomes 207
The Origins of Sex-Determining Loci and Recombination Suppression
around the Sex-Determining Locus Region 213
Evolution of an initial sex-determining region 213
Recombination suppression 214
Evolution of separate sexes and proto-sex cllromosomes 214
Evolution in Young and Proto-Sex Chromosomes: Sexually Antagonistic Genes 217
Determining the Age of Sex Chromosomes 218
Phylogenetic evidence 218
Divergence between X- and 'Y-linked sequences 218
Degeneration and Loss of Adaptation of Genes on the Nonrecombining Sex
Chromosome 219
Degeneration due to deleterious mutations 221
Interference with the spread offavorable Y-linked mutations 223
Selective sweeps 223
Evidence for degeneration processes 223
Do Y Chromosomes Always Degenerate, and What Determines
Which Genes Are Retained? 225
Transposable element accumulation 227
Low gene density of MSY regions 227
How Is Recombination Suppressed? 228
Chromosome inversions 228
Progressive recombination suppression 228
Further Chromosome Rearrangements 231
Neo-Sex Chromosome Formation 231
Molecular evolutionary evidence for genetic degeneration in
neo-sex chromosomes 232
Neo-sex chromosomes in species other than Drosophila 234
Evolution and Adaptation of Ancient Sex Chromosomes: Evolution of Dosage
Compensation and of Changed Gene Content 235
Dosage compensation 235
Discussion: Adaptation of Sex Chromosomes 238
CHAPTER
11 Molecular Signatures of Adaptive Evolution 241
Alan Filipski, Sonja Prohaska,Sudhir Kumar
Introduction 241
Codon Usage Bias and the Estimates of Selection Ratio
(00) 243
Hypermutability of CpG Dinucleotides and the Estimates of Selection Ratio 247
Using Pseudogenes to Estimate Nonsynonymous Mutation Rates 248
Rate of Intron Divergence as a Proxy for Nonsynonymous Mutation Rates 250
Direct Estimates of Mutation Rates in the Laboratory 252
Conclusion 253
Acknowledgments 254
CHAPTER
12 Gene Networks and Natural Selection 255
Andreas Wagner
Introduction 255
Is There a Network Biology? 256
Natural Selection and Network Parts 257
Natural Selection and Small-Scale, Local Network Features 261
Natural Selection and Global Network Structure 263
Natural selection and the degree distribution 264
Global features of transcriptional regulation networks 266
Alternative pathways in transcriptional regulation networks 266
Conclusion 269
CHAPTER
13 Human Evolutionary Genomics 271
Ines Hellmann and Rasmus Nielsen
Introd uction 271
Mutations-the Sources of Genome Variability 272
Point mutations 272
Insertions, deletions, and inversions 274
Recombination 276
Transposable Elements 278
Evolution through Gene Duplication 280
Analyses of Protein-Coding Regions 281
Comparative data 283
Detecting selective sweeps from SNP data 286
Human-Specific Genetic Adaptations 288
Selection on Noncoding Sequences 290
Regulatory sequences 292
Expression Data 293
Amount of Selection Affecting the Human Genome
and the Genetic Load of Humans 295
Conclusion 297
References 299
Index 337
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