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Explains why over half of the offspring produced by sexual reproduction will be male and yet in a sizeable number of these organisms, females greatly outnumber males. Introduces the strange world of male-killers, parthenogenetic reproduction, and ultra-selfish genes, using a large variety of fascinating examples. This book is the first to synthesize our knowledge of sex ratio distorters and their evolutionary effects.
LIST OF ILLUSTRATIONS xi LIST OF TABLES xv PREFACE xvii ACKNOWLEDGMENTS xxi PART I: THE SEXES, SEX DETERMINATION, AND SEX RATIOS 1 CHAPTER 1 The Logic of Sex 3 Summary 3 Introduction: Fantasy Sex Ratios! 3 Reproduction--Asexual, Sexual, and Secondary Asexual 4 The Ultimate Aim of Existence: The Passage of Genes 4 Asexual Reproduction 5 Sexual Reproduction 6 Sex Cells of Different Sizes 7 Males and Females: The Root of Sexual Conflict 9 Sexual Selection: Competing Males and Choosy Females 9 Pre- and Postcopulatory Male Competition 11 Female Choice 13 Direct and Indirect Benefits to Choosy Females 13 Sexy Sons 14 Handicaps and Condition-Dependent Handicaps 16 Sensory Exploitation 16 Mate Choice, Reinforcement, and Speciation 17 Cryptic Female Choice 19 Secondary Asexual Reproduction: Parthenogenesis 23 The Twofold Cost of Sex 24 The Diversity of Parthenogens 25 Horses for Courses 27 Why Is Parthenogenesis Rare? 30 Sex Accelerates Evolution 30 Muller's Ratchet 32 The Maintenance of Sex 34 CHAPTER 2 The Sex Ratio 36 Summary 36 Sex Ratio Equality 36 Are Males Parasites of Females? 36 The Ubiquity of the 1:1 Sex Ratio 37 A Divine Explanation 37 The Sex Chromosomes 38 Fisher's Theory 39 Biased Sex Ratios 41 The Trivers-Willard Hypothesis 41 Advantaged Daughters 43 Local Resource Competition Hypothesis 43 Local Mate Competition 43 The Influence of Relatedness on the Sex Ratio 46 Primary, Secondary, and Tertiary Sex Ratios 47 The Problem of Birth 47 CHAPTER 3 Sex Determination 49 Summary 49 Sexual Mosaics, Intersexes, and Hermaphrodites 49 Sex Chromosomes 51 XX/XY Sex Chromosome Systems 52 Human Sex Determination 52 Dosage Compensation 54 Sex Determination in Drosophila 55 ZZ/ZW Sex Determination 57 The Consequences of Female Heterogametry 61 Males without a Y Chromosome 62 Evolution of the Sex Chromosomes 63 Evolution within the Sex Chromosomes 64 Single-Gene Sex Determination 66 Sex Determination in Aphids: Two Homogametic Sexes! 66 Haplo-Diploidy 67 Degrees of Relatedness in Haplo-Diploids 70 Haplo-Diploid Sex Determination 73 Diploid Male Hymenoptera 73 Environmental Sex Determination 77 The Effect of Temperature in Reptiles 77 The Evolution of Environmental Sex Determination 78 Neighbor Sex Effects 79 Where Large Males Are Advantageous 79 Sex Changers 79 Microbes That Change Host Sex 81 Parental Control of the Sex Ratio? 82 CHAPTER 4 Genetic Law Makers and Law Breakers 84 Summary 84 Fair Genes and Foul 84 Conflict within the Genome 86 Driving Sex Chromosomes 87 Drive of Autosomes 89 The Effect of Sex Chromosome Drive on Host Populations 90 B-Chromosome Drive 92 Conflict between Cytoplasmic and Nuclear Genes 93 Misbehaving Mitochondria: Cytoplasmic Male Sterility 93 Cytoplasmic Incompatibility 95 Taxonomic Diversity of Species Showing CI 98 Mechanisms of Incompatibility 99 Vertical Transmission Efficiency of CI Wolbachia 100 Effects of CI Wolbachia Infection on Host Fitness 100 Field Prevalence Levels 101 The Taxonomic Relationships of CI Wolbachia and Their Hosts 102 Cytoplasmic Incompatibility and the Sex Ratio 103 Cytoplasmic Influence on Fertilization in Haplo-Diploids 104 Cytoplasmic Sex Ratio Distortion in ZZ/ZW Insects 104 Why Are Genetic Law Breakers the Exception Rather Than the Rule? 105 The Realization of the "Pretty Improbable" 108 PART II: ULTRASELFISH SYMBIONTS 111 CHAPTER 5 Feminization 113 Summary 113 From Adam to Eve, or Vice Versa 113 Microbe-Induced Sex Change 115 Horizontal and Vertical Transmission 116 Options for Maternally Inherited Symbionts in Male Hosts 116 Feminizing Microbes 118 The Diversity of Feminizers and Their Hosts 119 Sex Determination Systems of Hosts Susceptible to Feminization 122 How Are Genetic Male Crustacea Feminized? 123 The Dual Feminizing Action of Wolbachia 125 Sex Determination Warfare 125 Conflict between Feminizers: No Honor among Thieves 129 CHAPTER 6 Male-Killers 132 Summary 132 Two Types of Male-Killer 132 The Timing of Male Death 133 Late Male-Killers 134 Horizontal versus Vertical Transmission 135 Early Male-Killing 136 Fitness Compensation 136 Taxa Affected by Early Male-Killers 137 Early Male-Killing in Ladybird Beetles 138 The First Ladybird Male-Killer 138 The Diversity of Male-Killers in Ladybird Beetles 139 The Principal Advantage to Male-Killing in Ladybirds: Resource Reallocation 141 When Suicide Is Painless 143 Male-Killing and Kin Selection 144 Avoidance of Cannibalism 146 The Population Dynamics of Male-Killing in Ladybird Beetles 147 Vertical Transmission Efficiency 148 Horizontal Transmission 151 The Cost of Male-Killing in Ladybird Beetles 152 Male-Killing in Ladybirds: Testing the Predictions 153 Male-Killers and Resource Reallocation in Other Insects 153 Can Inbreeding Avoidance Account for Male-Killing Where Resource Reallocation Appears Unimportant? 155 The Extraordinary Case of the Nymphalid Butterflies 157 Biased Sex Ratios in Acraea Butterflies 157 More Hosts of Male-Killing Wolbachia 158 Exceptionally Female-Biased Populations 158 Wolbachias in Other Acraea Species 160 The Case of Danaus chrysippus: A Male-Killer with No Obvious Advantage 161 The Mechanism of Male-Killing 163 Sperm-Killers and Distortions of the Primary Sex Ratio 164 CHAPTER 7 The Evolutionary Consequences of Sex Ratio Distortion 166 Summary 166 Evolutionary Oddities? 166 Male-Killers and the Evolution of Clutch Size 167 The Sex Ratio and Sexual Selection 168 Male Mate Choice 169 Male Reproductive Investment and the Sex Ratio 170 Sex Role Reversal and Female Leks 171 Can Males Choose? 174 The Attractions of Incest 174 Host Coloration, Polymorphism, and Male-Killing 176 Biological Coincidences? 176 Male-Killers Are More Likely in Species with True Warning Colors 177 Sibling Recognition by Color Pattern 178 Sexually Transmitted Disease and the Sex Ratio 179 Male-Killer Suppressors 180 Male-Killer Rescue Genes and Intragenomic Conflict 182 Male-Killer Suppressors and the Presence of More Than One Male-Killer 183 Male-Killers and Mitochondrial DNA 184 The Evolution of Virulence 185 Clade Selection 186 Parthenogenesis: An Escape Route 187 CHAPTER 8 Parthenogenesis Inducers 189 Summary 189 Forsaking Needless Males 189 Parthenogenesis-Inducing Microbes 190 The Wasp and the Wolbachia 190 The Diversity of Symbiont-Induced Parthenogenesis 191 Horizontal Transfer of Parthenogenesis-Inducing Wolbachia 192 How Is Asexuality Induced? 194 The Population Dynamics of Parthenogenesis Inducers and Their Hosts 195 Effects of Parthenogenesis-Inducing Wolbachia on Host Mating 196 How Are Hosts Invaded? 197 Is Parthenogenesis an Escape Route? 198 CHAPTER 9 Sex Ratio Distorters: Implications and Applications 199 Summary 199 Musical Chairs 199 The Evolution of Sex 200 Immediate Benefit Hypotheses 202 Variation and Selection Hypotheses 203 Hedging One's Bets on "Entangled" Banks 203 The Red Queen 204 Deleterious Mutations and the Evolution of Sex 205 The Enigma of Sex 206 The Importance of Ultraselfish Genes 207 Is Wolbachia Special? 209 Implications of Sex Ratio Distorters for Evolutionary Geneticists 210 Can Sex Ratio Distorters Be Useful to Humanity? 211 Helping Beneficial Insects 211 Wolbachia and Pest Control 212 Six Impossible Things 213 GLOSSARY 215 REFERENCES 227 INDEX 241
Michael E. N. Majerus is Professor of Genetics at Cambridge University. He is the author of "Ladybirds, Evolution, Melanism: Evolution in Action and Moths".
Majerus's account is rich in examples from newly published studies and offers an excellent introduction to a rapidly developing discipline. -- John Bonner NewScientist