PART 1 FUNDAMENTALS OF EVOLUTIONARY BIOLOGY
1 Introduction ................................................. 3
1.1 What is disease? ........................................ 5
1.2 What evolution is: fundamental principles ............... 8
1.3 Time ................................................... 13
1.4 Constraints ............................................ 15
1.5 We are not alone ....................................... 15
1.6 Culture and gene-culture coevolution ................... 16
1.7 How evolutionary arguments fit alongside other
biological perspectives ................................ 16
1.8 Evolution and medicine ................................. 17
Key points .................................................. 18
2 Evolutionary theory ......................................... 19
2.1 Introduction ........................................... 19
2.2 What does evolutionary theory explain? ................. 21
2.3 How does evolution work? ............................... 24
2.4 Areas of debate and the limitations of the
adaptationist argument ................................. 39
2.5 Conclusion ............................................. 46
Key points .................................................. 48
3 The molecular basis of variation and inheritance ............ 49
3.1 Introduction ........................................... 49
3.2 Genes and disease ...................................... 49
3.3 The molecular basis of human genetic variation ......... 50
3.4 Factors affecting genetic variation .................... 59
3.5 Single-gene or Mendelian disorders ..................... 65
3.6 No single genes for common diseases .................... 68
3.7 Epigenetic mechanisms as a cause of variation .......... 71
3.8 Non-genetic inheritance ................................ 73
3.9 Conclusion ............................................. 77
Key points .................................................. 77
4 Evolution and development ................................... 79
4.1 Introduction ........................................... 79
4.2 Development: pre-ordained or plastic? .................. 81
4.3 Is development important to evolution? ................. 82
4.4 Developmental plasticity ............................... 84
4.5 Responses to environmental cues during development ..... 86
4.6 Epigenetic processes and development ................... 90
4.7 Learning and instinct .................................. 91
4.8 The evolution of novelty ............................... 93
4.9 Conclusion ............................................. 94
Key points .................................................. 96
5 The human life history ...................................... 97
5.1 Introduction ........................................... 97
5.2 General overview of life-history theory ................ 98
5.3 Body size and shape ................................... 107
5.4 Growth in humans ...................................... 113
5.5 Evolutionary analysis of the distinct features of
human growth .......................................... 124
5.6 Conclusion ............................................ 127
Key points ................................................. 129
6 Human evolution and the origins of human diversity ......... 131
6.1 Introduction .......................................... 131
6.2 The hominoid clade .................................... 131
6.3 Hominin evolution ..................................... 132
6.4 Genomic changes that make us human .................... 147
6.5 Human adaptation to local selection pressures ......... 148
6.6 Are humans still evolving? ............................ 153
6.7 Social and medical implications of human diversity .... 155
6.8 Conclusion ............................................ 157
Key points ................................................. 158
PART 2 EVOLUTION IN HEALTH AND DISEASE
7 An evolutionary framework for understanding human health
and disease ................................................ 161
7.1 Introduction .......................................... 161
7.2 Fundamental principles of evolutionary medicine ....... 162
7.3 Why has evolution left our bodies vulnerable to
disease? .............................................. 165
7.4 An evolutionary classification of ultimate
mechanisms affecting disease risk ..................... 167
7.5 Testing hypotheses in evolutionary medicine ........... 175
Key points ................................................. 175
8 Reproduction ............................................... 177
8.1 Introduction .......................................... 177
8.2 Sexual reproduction ................................... 178
8.3 Why did sex evolve? ................................... 178
8.4 Sex determination ..................................... 181
8.5 Reproductive strategies ............................... 182
8.6 Mate choice ........................................... 183
8.7 Sexual differences in the human ....................... 186
8.8 Sex differences in morbidity and mortality ............ 188
8.9 Human reproductive life cycle ......................... 190
8.10 Conclusion ............................................ 203
Key points ................................................. 204
9 Nutritional and metabolic adaptation ....................... 205
9.1 Introduction .......................................... 205
9.2 Strategies for energy storage ......................... 206
9.3 Human diet: an evolutionary history ................... 210
9.4 How can change in the environment increase disease
risk? ................................................. 218
9.5 Conclusion ............................................ 235
Key points ................................................. 236
10 Coevolution, infection, and immunity ....................... 237
10.1 Introduction .......................................... 237
10.2 Coevolution ........................................... 237
10.3 Humans and their associated species ................... 238
10.4 The challenge of infectious disease ................... 240
10.5 Pathogen emergence .................................... 241
10.6 Pathogen virulence and transmission ................... 242
10.7 Host defenses ......................................... 244
10.8 Public health measures ................................ 252
10.9 Vaccination ........................................... 253
10.10 Antibiotics .......................................... 255
10.11 Conclusion ........................................... 258
Key points ................................................. 258
11 Psychology and behavior .................................... 261
11.1 Introduction .......................................... 261
11.2 Biological determinants of culture and behavior ....... 261
11.3 Evolution of the human brain and behavior ............. 263
11.4 Evolution of social behavior .......................... 264
11.5 Evolutionary perspectives on psychology ............... 278
11.6 Evolutionary psychiatry ............................... 279
11.7 Conclusion ............................................ 284
Key points ................................................. 286
12 Cancer ..................................................... 287
12.1 Introduction .......................................... 287
12.2 Epidemiology of cancer ................................ 288
12.3 Ecology of cancer ..................................... 289
12.4 The biology of cancer ................................. 290
12.5 Cancer in the light of evolutionary mechanisms ........ 293
12.6 Implications of an evolutionary approach for the
prevention and treatment of cancer .................... 299
12.7 Conclusion ............................................ 302
Key points ................................................. 302
13 Evolutionary principles applied to medical practice and
public health .............................................. 303
13.1 Introduction: Understanding health and disease from
an evolutionary perspective ........................... 303
13.2 Testing evolutionary hypotheses in medicine ........... 305
13.3 Clinical examples ..................................... 305
13.4 An evolutionarily mismatched or novel environment ..... 305
13.5 Life-history-associated factors ....................... 311
13.6 Excessive and uncontrolled defense mechanisms ......... 313
13.7 Consequences of coevolution with microbes ............. 315
13.8 Results of evolutionary constraints ................... 317
13.9 An apparently harmful allele is maintained by
balancing selection ................................... 319
13.10 The consequences of sexual selection ................. 321
13.11 The outcomes of cladal and demographic history ....... 322
13.12 Value and limits of an evolutionary medicine
perspective .......................................... 324
Key points ................................................. 326
14 Evolution, medicine, and society ........................... 327
14.1 Introduction .......................................... 327
14.2 Origins of Darwin's theory ............................ 327
14.3 From Darwin to "Social Darwinism" ..................... 331
14.4 Eugenics .............................................. 332
14.5 The "Modern Synthesis," human evolution, and
medicine .............................................. 335
14.6 Evolution, society, and religion ...................... 337
14.7 Evolutionary thought and the human condition .......... 339
Key points ................................................. 339
References .................................................... 341
Index ......................................................... 365
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