Genes in Conflict
The Biology of Selfish Genetic Elements
Austin Burt
Robert Trivers
Preface
1. SELFISH GENETIC ELEMENTS
Genetic Cooperation and Conflict
Three Ways to Achieve "Drive"
Within-Individual Kinship Conflicts
Rates of Spread
Effects on the Host Population
The Study of Selfish Genetic Elements
Design of This Book
2. AUTOSOMAL KILLERS
The t Haplotype
Discovery
Structure of the t Haplotype
History and Distribution
Genetics of Drive
Importance of Mating System and Gamete Competition
Fate of Resistant Alleles
Selection for Inversions
Recessive Lethals in t Complexes
Enhancers and Suppressors
t and the Major Histocompatability Complex
Heterozygous (+/t) Fitness Effects: Sex Antagonistic?
Accounting for t Frequencies in Nature
Other Gamete Killers
Segregation Distorter in Drosophila
Spore Killers in Fungi
Incidence of Gamete Killers
Maternal-Effect Killers
Medea in Flour Beetles
HSR, scat+, and OmDDK in Mice
The Evolution of Maternal-Effect Killers
Gestational Drive?
Gametophyte Factors in Plants
3. SELFISH SEX CHROMOSOMES
Sex Chromosome Drive in the Diptera
Killer X Chromosomes
Killer Y Chromosomes
Taxonomic Distribution of Killer Sex Chromosomes
Evolutionary Cycles of Sex Determination
Feminizing X (and Y) Chromosomes in Rodents
The Varying Lemming
The Wood Lemming
Other Murids
Other Conflicts: Sex Ratios and Mate Choice
4. GENOMIC IMPRINTING
Imprinting and Parental Investment in Mammals
Igf2 and Igf2r: Oppositely Imprinted, Oppositely Acting Growth Factors in Mice
Growth Effects of Imprinted Genes in Mice and Humans
Evolution of the Imprinting Apparatus
The Mechanisms of Imprinting Involve Methylation and Are Complex
Conflict Between Different Components of the Imprinting Machinery
History of Conflict Reflected in the Imprinting Apparatus
Evolutionary Turnover of the Imprinting Apparatus
Intralocus Interactions, Polar Overdominance, and Paramutation
Transmission Ratio Distortion at Imprinted Loci
Biparental Imprinting and Other Possibilities
Other Traits: Social Interactions after the Period of Parental Investment
Maternal Behavior in Mice
Inbreeding and Dispersal
Kin Recognition
Functional Interpretation of Tissue Effects in Chimeric Mice
Deceit and Selves-Deception
Imprinting and the Sex Chromosomes
Genomic Imprinting in Other Taxa
Flowering Plants
Other Taxa Predicted to Have Imprinting
5. SELFISH MITOCHONDRIAL DNA
Mitochondrial Genomics: A Primer
Mitochondrial Selection within the Individual
"Petite" Mutations in Yeast
Within-Individual Selection and the Evolution of Uniparental Inheritance
Within-Individual Selection under Uniparental Inheritance
DUI: Mother-to-Daughter and Father-to-Son mtDNA Inheritance in Mussels
Cytoplasmic Male Sterility
Uniparental Inheritance Implies Unisexual Selection
Disproportionate Role of mtDNA in Plant Male Sterility
Mechanisms of Mitochondrial Action and Nuclear Reaction
CMS and Restorers in Natural Populations
CMS, Masculinization, and the Evolution of Separate Sexes
Pollen Limitation, Frequency Dependence, and Local Extinction
Resource Reallocation Versus Inbreeding Avoidance
Importance of Mutational Variation
CMS and Paternal Transmission
Other Traces of Mito-Nuclear Conflict
Mitochondria and Apoptosis
Mitochondria and Germ Cell Determination
Mitochondria and RNA Editing
6. GENE CONVERSION AND HOMING
Biased Gene Conversion
Molecular Mechanisms
Effective Selection Coefficients Due to BGC in Fungi
BGC and Genome Evolution
BGC and Evolution of the Meiotic Machinery
Homing and Retrohoming
How HEGs Home
HEGs Usually Associated with Self-Splicing Introns or Inteins
HEGs and Host Mating System
Evolutionary Cycle of Horizontal Transmission, Degeneration, and Loss
HEG Domestication and Mating-Type Switching in Yeast
Group II Introns
Artificial HEGs As Tools for Population Genetic Engineering
The Basic Construct
Increasing the Load
Preventing Natural Resistance and Horizontal Transmission
Population Genetic Engineering
Other Uses
7. TRANSPOSABLE ELEMENTS
Molecular Structure and Mechanisms
DNA Transposons
LINEs and SINEs
LTR Retroelements
Population Biology and Natural Selection
Transposition Rates Low But Greater Than Excision Rates
Natural Selection on the Host Slows the Spread of Transposable Elements
Rapid Spread of P Elements in D. melanogaster
Net Reproductive Rate a Function of Transposition Rate and Effect on Host Fitness
Reducing Harm to the Host
Transposition Rate and Copy Number "Regulation"
Selection for Self-Recognition
Defective and Repressor Elements
Extinction of Active Elements in Host Species
Horizontal Transmission and Long-Term Persistence
Transposable Elements in Inbred and Outcrossed Populations
Beneficial Inserts
Rates of Fixation
Transposable Elements and Host Evolution
Transposable Elements and Chromosomal Rearrangements
Transposable Elements and Genome Size
Co-Option of Transposable Element Functions and Host Defenses
Transposable Elements As Parasites, Not Host Adaptations or Mutualists
Origins
Ancient, Chimeric, and Polyphyletic Origins
8. FEMALE DRIVE
Selfish Centromeres and Female Meiosis
Abnormal Chromosome 10 of Maize
Other Knobs in Maize
Deleterious Effects of Knobs in Maize
Knobs, Supernumerary Segments, and Neocentromeres in Other Species
Meiosis-Specific Centromeres and Holocentric Chromosomes
Selfish Centromeres and Meiosis I
The Importance of Centromere Number: Robertsonian Translocations in Mammals
Sperm-Dependent Female Drive?
Female Drive and Karyotype Evolution
Polar Bodies Rejoining the Germline
9. B CHROMOSOMES
Drive
Types of Drive
Genetics of A and B Factors Affecting B Drive
Transmission Rates inWell-Studied Species
Absence of Drive
Degree of Outcrossing and Drive
Effects on the Phenotype
Effects on Genome Size, Cell Size, and Cell Cycle
Effects on the External Phenotype
Disappearance from Somatic Tissue
B Number and the Odd-Even Effect
Negative Effects of Bs More Pronounced under Harsher Conditions
Is the Sex of Drive Associated with the Sex of Phenotypic Effect?
B Effects on Recombination Among the As
Pairing of A Chromosomes in Hybrids
Neutral and Beneficial Bs
Beneficial B Chromosomes
B Chromosomes in Eyprepocnemis plorans: A Case of Continuous Neutralization?
Structure and Content
Size
Polymorphism
Heterochromatin
Genes
Tandem Repeats
The Origin of Bs
A Factors Associated with B Presence
Genome Size
Chromosome Number
Ploidy
Shape of A Chromosomes
Bs and the Sex Ratio
Paternal Sex Ratio (PSR) in Nasonia
X-B Associations in Orthoptera
Has the Drosophila Y Evolved from a B?
Other Effects of Bs on the Sex Ratio
Male Sterility in Plantago
10. GENOMIC EXCLUSION
Paternal Genome Loss in Males, or Parahaplodiploidy
PGL in Mites
PGL in Scale Insects
PGL in the Coffee Borer Beetle
PGL in Springtails?
Evolution of PGL
PGL and Haplodiploidy
Sciarid Chromosome System
Notable Features of the Sciarid System
An Evolutionary Hypothesis
Mechanisms
PGL in Gall Midges
Hybridogenesis, or Hemiclonal Reproduction
The Topminnow Poeciliopsis
The Water Frog Rana esculenta
The Stick Insect Bacillus rossius-grandii
Evolution of Hybridogenesis
Androgenesis, or Maternal Genome Loss
The Conifer Cupressus dupreziana
The Clam Corbicula
The Stick Insect Bacillus rossius-grandii
11. SELFISH CELL LINEAGES
Mosaics
Somatic Cell Lineage Selection: Cancer and the Adaptive Immune System
Cell Lineage Selection in the Germline
Evolution of the Germline
Selfish Genes and Germline-Limited DNA
Chimeras
Taxonomic Survey of Chimerism
Somatic Chimerism and Polar Bodies
12. SUMMARY AND FUTURE DIRECTIONS
Logic of Selfish Genetic Elements
Molecular Genetics
Selfish Genes and Sex
Fate of a Selfish Gene within a Species
Movement between Species
Distribution among Species
Role in Host Evolution
The HiddenWorld of Selfish Genetic Elements
References
Glossary
Index



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