| Preface | xi |
| Acknowledgments | xv |
Chapter 1 | The Language of Selection | 3 |
| Plan of the Book | 3 |
| Darwinian Dynamics | 5 |
| Major Evolutionary Transitions | 7 |
| Cooperation and Conflict | 8 |
| Fisherian Fitness | 9 |
| Deconstructing Fitness | 11 |
| Selection as Fitness Covariance | 13 |
| Mathematical Models | 16 |
| Adequacy Criterion for Understanding Fitness | 18 |
| Definitions of Basic Concepts | 18 |
Chapter 2 | Origin of Fitness | 19 |
| Complementarity | 19 |
| Spontaneous Creation | 20 |
| Self-Replication and the Origin of Fitness | 20 |
| Replicator Dynamics | 21 |
| Design Analysis of Molecular Replicator | 22 |
| Life History Evolution | 24 |
| Survival of the Fittest | 25 |
| Survival of Anybody | 26 |
| Overview of the Origin of Fitness | 27 |
Chapter 3 | The First Individuals | 29 |
| Origin of Gene Networks | 29 |
| Cooperation and Conflict | 32 |
| Survival of the First | 33 |
| Evolutionary Transitions Are Inherently Nonlinear | 35 |
| Origin of Hypercycles | 35 |
| Quasispecies | 37 |
| Population Structure | 39 |
| Kin Selection in Evolutionary Transitions | 40 |
| Conflict Mediation through Individuality | 42 |
| Further Evolution of the Cell | 43 |
| Heritable Capacities of Single Cells | 45 |
| Reconsidering Adaptedness and Fitness | 46 |
| Early Transitions in Evolution | 47 |
Chapter 4 | Evolution of Interactions | 49 |
| Gene Frequency Change | 49 |
| Population Growth | 50 |
| Frequency-Dependent Selection | 51 |
| Constant Selection | 51 |
| Adaptive Topography | 52 |
| Frequency Dependence Decouples Fitness in a Selection Hierarchy | 53 |
| Selection as Covariance | 55 |
| Fisher's Fundamental Theorem | 57 |
| Evolution in Hierarchically Structured Populations | 62 |
| Evolution of Multicellular Organisms | 64 |
| Kin Selection | 66 |
| Game Theory | 70 |
| Modification of Genetic Constraints | 71 |
| Population Dynamics and Natural Selection | 72 |
| Fitness Minima | 73 |
| Prisoner's Dilemma | 75 |
| Spatial Structure and the Evolution of Cooperation | 77 |
| The Problem of Frequency Dependence | 80 |
Chapter 5 | Multilevel Selection of the Organism | 81 |
| A Scenario | 81 |
| A Model for the Emergence of Organisms | 81 |
| Recurrence Equations | 84 |
| Within-Organism Mutation Selection Model | 86 |
| Mutation Rate | 87 |
| Covariance Methods | 89 |
| The Risk of Development | 90 |
| Increase of Cooperation | 91 |
| Level of Cooperation among Cells within Organisms | 93 |
| Fitness of Organisms | 95 |
| Effect of Sex and Diploidy on the Emerging Organism | 101 |
| Strengths and Weaknesses of the Model | 102 |
Chapter 6 | Rediscovering Individuality | 107 |
| Evolutionary Individuals | 107 |
| Two-Locus Modifier Model | 109 |
| Model Parameters | 112 |
| Equilibria of the System | 112 |
| Evolution of the Germ Line | 113 |
| Evolution of the Mutation Rate | 118 |
| Evolution of Self-Policing | 119 |
| Evolution of Adult Size | 121 |
| Effect of Transition on the Level of Cooperation | 122 |
| Increase of Fitness Covariance at Organism Level | 122 |
| Heritability of Fitness and the Evolution of Individuality | 124 |
| Sex and Individuality | 127 |
| Origin of Multicellular Life | 129 |
| Transitions in Individuality | 131 |
Chapter 7 | Fitness Explanations | 133 |
| Overview of Fitness and Natural Selection | 133 |
| Trading Fitness through Cooperation | 135 |
| Kinship and Population Structure | 136 |
| Conflict Mediation | 137 |
| Reconsidering Fitness | 137 |
| The "Tautology Problem" | 140 |
| Surrogates for Natural Selection | 142 |
| Evolution of Selfing | 145 |
| Cost of Sex | 146 |
| Immortality, Death, and the Life Cycle | 150 |
| Kin Selection of Altruism | 152 |
| Heterozygote Superiority | 154 |
| Sickle Cell Anemia | 156 |
| Darwin's Dilemmas | 158 |
Chapter 8 | A Philosophy of Fitness | 161 |
| Dynamics of Design | 161 |
| What Makes Biology Different? | 163 |
| Success and Design | 164 |
| Long-Term versus Short-Term Measures of Fitness | 166 |
| Darwinian Dynamics | 167 |
| Natural Selection as a Biological Law | 169 |
| Paradigms for Natural Selection | 173 |
| Fitness in Darwinian Dynamics | 174 |
| The Insufficiency of Individual Fitness | 175 |
| Heritability and Natural Selection | 179 |
| Schema for Natural Selection | 181 |
| The Propensity Interpretation of Fitness | 183 |
| Brandon's Approach | 186 |
| Heritable Capacities as Components of Design | 189 |
| Overall Adaptedness of Organisms | 191 |
| Masking of Adaptiveness | 194 |
| Are Adaptation Concepts Necessary? | 196 |
| F-Fitness and Evolutionary Explanations | 197 |
| Explaining Fitness | 200 |
Appendix A | Supporting Analyses | 203 |
| Statistics of Fitness and Selection | 203 |
| Equilibria for Modifier Model (G = 0) | 217 |
| Cost of Sex in Diploids | 218 |
Appendix B | Fitness Phrases | 222 |
Appendix C | Notation | 226 |
| Notes | 233 |
| References | 237 |
| Index | 253 |