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Sex Wars:
Genes, Bacteria, and Biased Sex Ratios
Michael E. N. Majerus

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TABLE OF CONTENTS:

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

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File created: 10/23/2013

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