Elements of evolutionary genetics:
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[2012]
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245 | 1 | 0 | |a Elements of evolutionary genetics |c Brian Charlesworth and Deborah Charlesworth |
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adam_text | Titel: Elements of evolutionary genetics
Autor: Charlesworth, Brian
Jahr: 2012
Contents
Preface xxiii
Introduction xxv
Variability and its measurement 1
Chapter summary 1
Introduction 2
1.1. Classical and quantitative genetic studies of variability 3
l.l.i. Discrete variability 3
l.l.ii. Quantitative variability 4
l.l.ii.a. Describing quantitative variability 4
l.l.ii.b. Genotypes and phenotypes 7
l.l.ii.c. The genetic basis of quantitative variability 8
l.l.ii.d. Modeling quantitative trait inheritance 10
1.1.iii. Concealed variability and mutational variability 11
l.l.iii.a. Introduction 11
l.l.iii.b. The genetics of inbreeding depression 11
1.1.iv. Limitations of the classical and quantitative genetic
approaches 13
1.2. Molecular variability 14
1.2.i. Gel electrophoresis of proteins 14
1.2.i.a. Introduction 14
1.2.i.b. The discovery of high molecular diversity 16
1.2.ii. Detecting variability by restriction mapping 20
1.2.iii. DNA sequencing 21
1.2.iii.a. Introduction 21
1.2.iii.b. DNA sequence polymorphisms; SNPs 22
1.2.iii.c. Other types of DNA sequence variants: repeat
number polymorphisms and copy number
polymorphisms 26
1.2.iv. Measuring DNA sequence variability 27
1.2.iv.a. Introduction 27
1.2.iv.b. Statistics for describing variability 27
1.2.v. Haplotypes 31
1.2.v.a. Definition and use in measuring variability 31
1.2.v.b. The effect of recombination on haplotype
variability 32
1.3. The causes and maintenance of variability 33
CONTENTS
1.3.i. Conservation of variability by the mechanism ^
of inheritance
1.3.i.a. Hardy-Weinberg equilibrium
1.3.i.b. Gene conversion and mutation 35
1.3.ii. Nonrandom mating and identity by descent 35
1.3.ii.a. Identity by descent and coalescence of alleles in
common ancestors 36
1.3.ii.b. Genotype frequencies with inbreeding and
assortative mating 39
1.3.ii.c. Regular systems of mating 40
1.3.iii. Mutation: the source of new variants 42
1.3.iii.a. The nature of mutations 42
1.3-iii-b. Mutation rates 43
1.3.iii.c. Mutation and evolutionary change 43
Problems 45
Basic selection theory and the maintenance of variation 47
Chapter summary 47
Introduction 48
2.1. Basic selection theory: constant fitnesses 49
2.1.i. Haploid populations 49
2.1.ii. Diploid populations 52
2.1.ii.a. Introduction 52
2.1.ii.b. Directional selection 54
2.1.ii.c. Heterozygote advantage and the maintenance of
variation 56
2.1.ii.d. Biological implications of heterozygote
advantage 59
2.1.ii.e. Mean fitness 61
2.1.ii.f. Heterozygote disadvantage 63
2.2. Variable fitnesses 64
2.2.i. Frequency-dependent selection 65
2.2.i.a. Theory 65
2.2.i.b. Examples of frequency dependence 65
2.2.ii. Fitnesses that vary in time 72
2.2.iii. Spatial variation in fitnesses, hard versus soft selection,
and speciation 77
2.2.iv. The biochemical basis of fitness differences 81
Problems ^
Directional selection and adaptation 35
Chapter summary fiI-
CONTENTS ix
Introduction 86
3.1. Directional selection on a pair of allelic variants 88
3.1.i. Introduction 88
3.l.ii. Haploid populations 88
3.1.iii. Autosomal inheritance with random mating 90
3.l.iii.a. Introduction 90
3.1.iii.b. Results for weak selection 90
3.1.iii.c. Behavior at extreme frequencies 93
3.l.iii.d. Continuous time 93
3.1.iv. Biological applications 94
3.1.iv.a. Estimating the strength of selection 94
3.1.iv.b. Experimental evolution 95
3.1.iv.c. Haldane s sieve 96
3.1.iv.d. Developmental constraints on evolution 96
3.1.v. More complex selection models: relaxing some of the
assumptions 97
3.1.v.a. Autosomal inheritance with sex differences
in selection 97
3.1.v.b. Selection on an X-linked gene: Haldane s sieve and
sexual antagonism 98
3-l.v.c. Selection in inbreeding populations 100
3.1.v.d. Kin selection 101
3.2. Survival and fixation probability of a favorable mutation 106
3.2.i. The problem 106
3.2.U. The solution 107
3.2.iii. Biological implications 110
3.3. Selection on quantitative traits 114
3.3.i. Introduction 114
3.3.ii. Quantitative genetic parameters 114
3.3.ii.a. Introduction 114
3.3.ii.b. Components of genetic variance and covariances
between relatives 115
3.3.iii. Response to selection 118
3.3.iii.a. Change over one generation 118
3.3.iii.b. Selection coefficients at individual sites 122
3.3.iii.c. Long-term responses to selection 124
3.3.iv. Multiple traits 126
3.4. The evolutionary genetics of adaptation of phenotypes
and proteins 129
3.4.i. Introduction 129
3.4.ii. Fishers geometric model of adaptation 130
3.4.iii. Adaptive evolution of sequences 133
3.4.iii.a. Introduction 133
CONTENTS
X
3.4.iii.b. The distribution of the fitness effects of
mutations fixed in an adaptive walk
3.4.iv. Testing the theory
3.4.iv.a. The theoretical predictions
3.4.iv.b. Genetics of interpopulation and interspecies
crosses
3.4.iv.c. Experimental evolution
3.4.v. Multiple fitness peaks
Problems
4 Migration, mutation, and selection 143
Chapter summary *43
Introduction 144
4.1. Migration and selection 145
4.1.i. Migration and selection involving two populations 145
4.l.ii. Migration and selection in a continuous habitat 148
4.1.ii.a. Clines 148
4.l.ii.b. Modeling clines 150
4.1.ii.c. Using cline widths to measure selection 155
4.l.ii.d. Hybrid zones 157
4.1 .ii.e. The wave of advance of a favorable mutation 158
4.2. Mutation and selection 159
4.2.i. Introduction 159
4.2.ii. Mutation-selection equilibrium 160
4.2.ii.a. Autosomes with random mating 160
4.2.ii.b. Other cases 161
4.2.ii.c. Mutation and selection at multiple sites 162
4.2.ii.d. Estimating mutation rates from equilibrium
frequencies 163
4.3. Genetic load 164
4.3.i. Introduction 164
4.3.ii. Mutational load 164
4.3.ii.a. Autosomal inheritance with random mating 164
4.3.ii.b. Other cases 165
4.3.ii.c. Multiple sites 165
4.3.iii. Segregational load 168
4.3.iv. The cost of selection
4.4. Inbreeding depression
4.4.i. Introduction
4 4 ii* Mutation and inbreeding depression in a randomly
mating population
4.4.ii.a. The effects of mutation and selection at a
single site
133
135
135
136
137
138
141
CONTENTS
4.4.ii.b. Multiple sites 171
4.4.ii.c. The inbreeding load 173
4.4.iii. Heterozygote advantage and inbreeding depression 174
4.4.iv. Testing alternative explanations of inbreeding
depression 175
4.4.iv.a. Use of BiA ratios 175
4.4.iv.b. Alternative approaches 179
4.4.v. Does dominance evolve? 182
4.5. Genetic variation in fitness components and other quantitative
traits 183
4.5.i. Mutational variation 183
4.5.i.a. Direct effects of mutations on fitness 183
4.5.i.b. Mutation and stabilizing selection 186
4.5.ii. Variability maintained by selection 191
Problems 192
The evolutionary effects of finite population size:
basic theory 195
Chapter summary 195
Introduction 196
5.1. Modeling genetic drift 199
5.1.i. Drift without mutation 199
5.1 .i.a. The Wright-Fisher model 199
5.1.i.b. Approach to genetic uniformity of a
population 202
5.1.i.e. Increase in variance of allele frequencies 204
5.1.ii. Drift and mutation 205
5.1.ii.a. Identity in state of a pair of alleles and the infinite
alleles model 205
5.1.ii.b. The infinite sites model 207
5.1.iii. The coalescent process 207
5.l.iii.a. Basic properties 207
5. l.iii.b. The genealogy of two alleles 207
5.l.iii.c. Gene genealogies in general 209
5.l.iii.d. Mutations, the coalescent process, and sequence
diversity 209
5.l.iii.e. Simulating the coalescent process 211
5.1.iii.f. Microsatellite loci 215
5.2. Effective population size 216
5.2.i. Introduction 216
5.2-ii. Effective population size and the coalescent process 217
5.2.ii.a. Defining effective population size 217
5.2.ii.b. Calculating N with constant population size 218
CONTENTS
5.2.iii. Effective population sizes for some biologically
interesting cases 221
5.2.iii.a. Different modes of inheritance (constant
population size) 222
5.2.iii.b. Comparisons of different modes of
inheritance 223
5.2.iii.c. Varying population size 225
5.2.iii.d. Age- and stage-structured populations and the
Moran model 226
5.2.iii.e. Estimating effective population sizes 227
5.3. Probability distributions of allele frequencies 228
5.3.i. Introduction 228
5.3.ii. The Wright-Fisher model with mutation and
selection 229
5.3.ii.a. The transition from one generation to the next.. .229
5.3.ii.b. The probability distribution of variant
frequencies 230
5.3.ii.c. Diffusion equations 231
5.3.iii. Allele frequency distributions: beyond the Wright-Fisher
model 231
5.3.iii.a. Variance effective population size 231
5.3.iii.b. Solving the forward diffusion equation 232
5.3.iii.c. The equilibrium distribution with mutation
between neutral variants 233
5.3.iii.d. The equilibrium distribution with mutation and
selection 239
5.3.iii.e. Biological implications 240
Appendix to Chapter 5: Diffusion equations 241
5A.1. General formulation 241
5 A.2. Determining the equilibrium distribution of allele
frequencies 242
Problems 243
Molecular evolution and variation 245
Chapter summary 245
Introduction 246
6.1. Measuring rates of DNA sequence evolution 247
6.1.i. Comparing a pair of sequences 247
6. l.ii. Estimating rates of divergence 248
6.l.ii.a. Basic principles 248
6.1.ii.b. Correcting for multiple changes 250
6.1.ii.c. Stochasticity of the substitution process 251
6.1.ii.d. Correcting for reverse and multiple changes i i *252
CONTENTS Xiii
6.1.ii.e. Nonsynonymous and synonymous changes 254
6.1.ii.f. Approximate rate constancy of molecular sequence
evolution 256
6.2. The causes of DNA sequence evolution 257
6.2.i. Introduction 257
6.2.ii. Neutral sequence evolution 257
6.2.ii.a. Expectations for sequence divergence under
neutrality 257
6.2.ii.b. Fixation of neutral mutations and divergence of
sequences from closely related species 258
6.2.ii.c. Testing the neutrality of sequence evolution 260
6.2.iii. Selection in finite populations 261
6.2.iii.a. Rates of substitution for unique mutations 261
6.2.iii.b. Rates of substitution for recurring mutations 264
6.2.iv. Implications for molecular evolution and for studying
selection 264
6.2.v. Detecting positive selection when purifying selection also
acts: codon-based models 266
6.3. The distribution of variant frequencies under mutation,
selection, and drift 268
6.3.i. Introduction 268
6.3.ii. Statistical equilibrium under mutation, selection,
and drift 270
6.3.ii.a. The general model 270
6.3.ii.b. The equilibrium frequency distribution under
neutrality 272
6.3.ii.c. Properties of the statistical equilibrium with
selection: rates of evolution 275
6.3.ii.d. Properties of the statistical equilibrium with
selection: variability 277
6.4. Testing for selection from polymorphism and divergence data .. .279
6.4.i. Introduction 279
6.4.ii. The McDonald-Kreitman (MK) test 280
6.4.ii.a. Introduction 280
6.4.ii.b. Principle of the MK test 281
6.4.ii.c. Potential difficulties with the MK test 284
6.4.ii.d. Estimating the fraction of amino acid fixations
caused by positive selection 285
6.4-iii. Tests based on variant frequencies 287
6.4.iii.a. Introduction 287
6.4.iii.b. Tajima s D 288
6.4.iii.c. Fu s and Li s test statistics 289
6.4.iii.d. Other test statistics 290
CONTENTS
6.4-iii.e- Problems of interpretation of tests using variant
frequencies 290
6.4.iv. Estimating selection intensities 291
6.4.iv.a. Introduction 291
6,4.iv.b. Fitting the distribution of variants to a one-way
mutation model 292
6.4.iv.c. Other methods and implications of the results 294
6.4.iv.d. Using polymorphism and divergence statistics 296
Appendix to Chapter 6: More diffusion equation results 297
6A.1. Fixation probabilities 297
6A.2. The equilibrium distribution of variant frequencies with
one-way mutations 299
Problems 301
Genetic effects of spatial structure 303
-snl
Chapter summary
Introduction 504
7.1. Describing population differentiation 305
7.1.i. Use of genetic markers to describe differences among
populations 305
7.1.i.a. Variances in allele frequencies and FST 305
7.1.i.b. Gene diversities for allelic variants 308
7.1.i.e. Nucleotide site diversity 309
7.1.i.d. Other methods 309
7.l.ii. Empirical patterns 310
7.2. The theory of neutral variation with spatial structure 310
7.2.i. General principles 313
7.2.i.a. Introduction 313
7.2.i.b. Fast time scales and migration models 313
7.2.ii. Mean coalescence times for migration models 315
7.2.ii.a. Introduction 315
7.2.ii.b. The island model 317
7.2.ii.c. Stepping-stone models 320
7.2.ii.d. Mean coalescence times for general migration
models 322
7.2.ii.e. The strong migration limit 328
7.2.iii. Continuous populations 329
7.2.iii.a. General considerations 329
7.2.iii.b. Genetic differentiation in continuous
populations 332
7.2.iv. Colonization and extinction models 333
7.2.iv.a. Introduction
7.2.iv.b. The exchangeable deme model 334
CONTENTS XV
7.2.v. Sampling of alleles from spatially structured
populations 338
7.2.v.a. Introduction 338
7.2.v.b. The large deme number approximation 340
7.2.V.C. Scattering and collecting phases 341
7.2.v.d. Within-population coalescence times 347
7.2.vi. The effects of population history 348
7.2.vi.a. Introduction 348
7.2.vi.b. Distinguishing historical changes from stable
structure 348
7.3. The theory of selection with spatial structure 350
7.3.i. Introduction 350
7.3.ii. Fixation probabilities in structured populations 350
7.3.iii. Variant frequency distributions with selection and spatial
structure 353
7.3.iii.a. The infinite-deme island model 353
7.3.iii.b. Quantitative traits and Qsr values 359
7.3.iii.c. The sampling properties of alleles under the large
deme number approximation 360
7.3.iv. Effects of population structure on mean fitness and selection
among demes 361
7.3.iv.a. The effects of drift on population mean fitness 361
7.3.iv.b. Inter-deme selection 362
Problems 364
8 Multiple sites and loci 366
Chapter summary 366
Introduction 368
8.1. Describing the state of a population 368
8.1.i. Haplotype frequencies 368
8.1.ii. Measures of linkage disequilibrium 370
8.1.ii.a. Pairs of sites 370
8.l.ii.b. Multiple sites 373
8.1.iii. Empirical results on linkage disequilibrium 374
8.1.iii.a. Observations based on classical genetics 374
8.1.iii.b. Results from electrophoretic surveys 376
8.1.iii.c. Results from surveys of DNA sequence
variability 376
8.1.iii.d. Mapping disease genes and quantitative
trait loci 378
8.2. Causes of linkage disequilibrium: neutral sites 378
8.2.i. Introduction 378
8.2.ii. Neutral sites at two loci in an infinite population 379
CONTENTS
379
8.2.ii.a. Random mating *
8.2.ii.b. Inbreeding populations - 382
8.2.iii. The effects of finite population size on linkage
disequilibrium . * . *
8.2.iii.a. Linkage disequilibrium and drift in a panmictic
population 383
8 2 iii b Linkage disequilibrium and the coalescent
384
process
8.2.iii.c. Estimating recombination rates 386
8.2.iii.d. The effects of spatial structure 387
8.2.iii.e. Other factors affecting linkage disequilibrium 389
8.3. Neutral and nearly neutral variants linked to selected sites 390
8.3.i. Introduction 3^0
8.3.ii. The effects of balancing selection 393
8.3.ii.a. The effects of balancing selection on neutral
variability 393
8.3.ii.b. Associative overdominance caused by
heterozygote advantage 396
8.3.ii.c. Testing for the signature of balancing selection:
the HKA test 397
8.3.ii.d. Testing for balancing selection: Dv D^ and
haplotype tests 398
8.3.iii. Background selection 398
8.3.iii.a. Reduced neutral variability caused by linked
deleterious mutations 398
8.3.iii.b. Associative overdominance due to linkage
disequilibrium with deleterious mutations 403
8.3.iii.c. Associative overdominance due to identity
disequilibrium 404
8.3.iv. The joint effects of spatial and genetic structure 406
8.3.v. Hitchhiking by favorable mutations (selective sweeps) .. .407
8.3.v.a. Effects on levels of neutral variability at
linked sites 407
8.3.v.b. Effects on variant frequency spectra 413
8.3.v.c. Selective sweeps with recombination 415
8.3.v.d. Partial sweeps 416
8.4. Selection at multiple sites in the genome 418
8.4.i. Introduction 41g
8.4.ii. Two-site systems 41g
8.4.ii.a. General description
8.4.ii.b. Two-site fitness matrices 421
8.4.U.C. Properties of stable equilibria ...!. !!!!!! !! *423
8.4,ii.d. Weak selection and loose linkage 425
CONTENTS XVii
8.4.iii. Multisite systems 429
8.4.iii.a. Properties of equilibria 429
8.4.iii.b. Dynamics under QLE 430
8.4.iv. The evolutionary dynamics of quantitative traits 430
8.4.iv.a. The infinitesimal model 430
8.4.iv.b. Extending the infinitesimal model 434
8.4.v. Wright s shifting balance model of evolution 434
8.4.v.a. Epistasis and multiple peaks of mean fitness 434
8.4.v.b. The fixation of chromosomal rearrangements 436
8.4.vi. The evolution of hybrid fitness loss 437
Appendix to Chapter 8: Linkage disequilibrium at neutral sites in
finite populations 439
8A.1. The general method 439
8A.2. Application to linkage disequilibrium 440
Problems 442
9 The evolution of breeding systems, sex ratios, and life
histories 444
Chapter summary 444
Introduction 445
9.1. Sexual versus asexual reproduction; inbreeding versus
outcrossing 447
9.1.i. Introduction 447
9.1.ii. Asexual versus sexual reproduction 447
9.1.ii.a. Introduction 447
9.1.ii.b. The cost of sex 447
9.1.ii.c. The maintenance of sexual reproduction 449
9-l.iii. Sexual systems and mating systems 450
9.1.iii.a. Sexual systems 450
9.1.iii.b. Mating systems 450
9.1.iii.c. Self-incompatibility 452
9.l.iii.d. Detecting outcrossing and estimating outcrossing
rates in natural populations 453
9.2. The evolution of inbreeding and outbreeding 455
9.2.i. Introduction 455
9.2.ii. The evolution of selfing rates 456
9.2.ii.a. Introduction 456
9.2.ii.b. The transmission advantage of selfing
(the cost of outcrossing) 458
9.2.ii.c. Why do many organisms outcross? Inbreeding
depression and the evolution of selfing rates 458
9.2.ii.d. More realistic models of inbreeding depression;
purging of deleterious mutations 464
xviii
CONTENTS
9 2.ii.e. Origins and evolution of self-incompatibility
.465
systems
9.2.iii. The breakdown of self-incompatibility systems 466
9.2.iii.a. The breakdown of heterostyly * *466
9.2.iii.b. The breakdown of homomorphic SI 467
9.3. The evolution of separate sexes 467
9.3.i. Introduction ^67
9.3.ii. The evolutionary stability of cosexuality 468
9.3.iii. Pathways to dioecy from cosexuality 468
9.3.iv. When will unisexuals invade cosexual populations? 471
9.3.iv.a. The establishment of cytoplasmic male sterility
factors in populations 471
9.3.iv.b. The establishment of nuclear male sterility
factors 472
9.3.iv.c. Testing the theory 474
9.3.iv.d. Androdioecy 476
9.3.v. The transition from gynodioecy to dioecy 478
9.3.v.a. The effect of the presence of females 478
9.3.v.b. The importance of linkage 480
9.4. Evolutionarily stable strategies, sex ratios, and sex allocation 483
9.4.i. Introduction 483
9.4.ii. Evolutionarily Stable Strategies 484
9.4.iii. The ESS sex ratio in a panmictic population 486
9.4.iii.a. Equal costs of male and female offspring 486
9.4.iii.b. Unequal costs of sons and daughters 486
9.4.iv. Extraordinary sex ratios 488
9.4.iv.a. Local mate competition 488
9.4.iv.b. Haplodiploid eusocial insects 490
9.4.v. Sex allocation in cosexuals 492
9.4.v.a. Theoretical models 492
9.4.v.b. Empirical tests 493
Age-structured populations and the evolution of life
histories 494
9.5.i. Introduction 494
9.5.ii. Demographic models 494
9.5.ii.a. Basic concepts and notation 494
9.5.ii.b. Predicting population growth 496
9.5.ii.c. Population genetics for age-structured
populations 502
9.5.iii. Selection equations 5Q2
9.5.ni.a. The general recursion relations with no sex
differences 5Q2
9.5.
CONTENTS Xix
9.5.iii.b. Equilibrium conditions 504
9.5.iii.c. Stability and invasion conditions 505
9.5.iii.d. Weak selection approximations 506
9.5.iii.e. Relationships between different fitness
measures 507
9.5.iii.f. More general situations 508
9.5.iv. Implications for population genetic studies 508
9.5.v. Principles of life history evolution 510
9.5.v.a. The sensitivity of fitness to changes in demographic
parameters 510
9.5.v.b. Trade-off models 512
9.5.v.c. Other life history models 514
9.5.v.d. The evolution of aging 515
Problems 519
10 Some topics in genome evolution 521
Chapter summary 521
Introduction 522
10.1. Evolution of noncoding sequences and codon usage 523
lO.l.i. Introduction 523
10. l.i.a. Base composition 523
lO.l.i.b. Codon usage bias 524
10. l.ii. Evolutionary forces affecting base composition 525
lO.l.ii.a. Introduction 525
lO.l.ii.b. Mutation and genetic drift 526
lO.l.ii.c. Lower effectiveness of selection with reduced
recombination 527
lO.l.ii.d. Biased gene conversion 527
lO.l.ii.e. Departures from equilibrium base
composition 529
lO.l.iii. Evolutionary forces affecting codon usage 530
lO.l.iii.a. Measuring codon usage bias 530
lO.l.iii.b. Causes of selection for codon usage bias 533
lO.l.iv. Estimating the intensity of selection or biased gene
conversion 535
lO.l.iv.a. The intensity of selection and BGC on
noncoding sites 536
lO.l.iv.b. The intensity of selection on codon usage 537
10.2. The evolutionary causes and consequences of genetic
recombination 539
10.2.i. Introduction 539
10.2.ii. The cost of recombination 540
XX
CONTENTS
10.2.iii. The evolution of close linkage 543
10.2.iii.a. Inversion polymorphisms 543
10.2iii.b. Supergenes 543
10.2.iii.c. Centromeric and telomeric suppression of
crossing over 544
10.2.iii.d. Congealing of the genome versus selection for
recombination 545
10.2.iv. Possible advantages of sex and recombination:
deterministic mechanisms 546
10.2.iv.a. Introduction 546
10.2.iv.b. Fluctuating epistasis 548
10.2.iv.c. Moving optima 548
10.2.iv.d. Mutation and selection 549
10.2.V. Stochastic processes 550
10.2.v.a. General considerations 550
10.2.v.b. Selective sweeps 551
10.2.v.c. Background selection 552
10.2.v.d. Muller s ratchet 552
10.2.v.e. Reverse mutation and Hill-Robertson
interference 556
10.2.vi. Empirical evidence concerning the consequences of
recombination 556
10.2.vi.a. Reduced levels of adaptation in regions with low
recombination 556
10.2.vi.b. Discriminating between the different possible
advantages of recombination 559
10.3. Selfish DNA
10.3.i. Introduction 561
10-3-ii. Gametic selection and segregation distortion 561
10.3.ii.a Gametic selection 561
10.3.U1. Meiotic drive and segregation distortion 562
10.3.iii.a. Introduction 562
10.3.iii.b. Segregation distortion 563
Jn a - j ExamPles of segregation distorter systems 565
10.3.ni.d. The evolution of segregation distorter
systems 568
10.3.iv. Transposable elements . ^ 571
10.3.iv.a. Introduction
lotiv c TP Ker1Ut °n °f TES in Senomes 573
10 3 v d P btaV1°rJin genomes and populations 574
I0.3.iv.d. Population dynamics of TEs 576
CONTENTS XXi
10.3.iv.e. Discriminating among possible regulators of
element abundance 580
10.3.iv.f. Genomes and genomic regions with high TE
contents 584
Problems 585
Appendix 587
Al. Basic mathematics 587
Al.i. Differentiation 587
Al.ii. Representing functions by series 588
Al.ii.a. Taylor s series 588
Al.ii.b. Approximate numerical solutions of equations .. .589
Al.ii.c. Some useful series 589
Al.iii. Integration 590
Al.iv. Vectors and matrices 591
Al.iv.a. Some definitions 591
Al.iv.b. The inverse of a matrix and the solution of linear
equations 593
Al.iv.c. Eigenvalues and eigenvectors 594
Al.iv.d. The spectral expansion of a matrix 596
A2. Basic probability and statistics 597
A2.i. Introduction 597
A2.ii. The rules of probability 597
A2.iii. The moments of a distribution 598
A2.iv. Pairs of variables 599
A2.v. Some important probability distributions 601
A2.v.a. The binomial distribution 601
A2.v.b. The multinomial distribution 602
A2.v.c. The geometric and exponential distributions 602
A2.v.d. The Poisson distribution 603
A2.v.e.The normal distribution 603
A2.v.f. Other distributions 605
A2.vi. Statistical estimation 605
A2.vi.a. Introduction 605
A2.vi.b. Maximum likelihood 606
A2.vi.c. Bayesian methods 608
Answers to problems 609
Chapter 1 609
Chapter 2 611
Chapter 3 614
Chapter 4 618
xxii
CONTENTS
Chapter 5 621
Chapter 6 624
Chapter 7 629
Chapter 8 633
Chapter 9 637
Chapter 10 640
References 642
Author Index 715
Subject Index 723
|
any_adam_object | 1 |
author | Charlesworth, Brian 1945- Charlesworth, Dorothy ca. 20. Jh |
author_GND | (DE-588)1089153910 (DE-588)1309681058 |
author_facet | Charlesworth, Brian 1945- Charlesworth, Dorothy ca. 20. Jh |
author_role | aut aut |
author_sort | Charlesworth, Brian 1945- |
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callnumber-first | Q - Science |
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dewey-ones | 572 - Biochemistry |
dewey-raw | 572.8/38 |
dewey-search | 572.8/38 |
dewey-sort | 3572.8 238 |
dewey-tens | 570 - Biology |
discipline | Biologie |
format | Book |
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id | DE-604.BV040450830 |
illustrated | Illustrated |
indexdate | 2024-07-10T00:24:12Z |
institution | BVB |
isbn | 9780981519425 0981519423 |
language | English |
lccn | 2009042482 |
oai_aleph_id | oai:aleph.bib-bvb.de:BVB01-025298491 |
oclc_num | 731094156 |
open_access_boolean | |
owner | DE-M49 DE-BY-TUM DE-83 DE-19 DE-BY-UBM |
owner_facet | DE-M49 DE-BY-TUM DE-83 DE-19 DE-BY-UBM |
physical | xxvii, 734 Seiten Illustrationen 26 cm |
publishDate | 2012 |
publishDateSearch | 2012 |
publishDateSort | 2012 |
publisher | Roberts and Company Publishers |
record_format | marc |
spelling | Charlesworth, Brian 1945- (DE-588)1089153910 aut Elements of evolutionary genetics Brian Charlesworth and Deborah Charlesworth Greenwood Village ; Colorado Roberts and Company Publishers [2012] © 2012 xxvii, 734 Seiten Illustrationen 26 cm txt rdacontent n rdamedia nc rdacarrier Literaturverz. S. 642 - 713 Evolutionary genetics Genetik (DE-588)4071711-2 gnd rswk-swf Evolution (DE-588)4071050-6 gnd rswk-swf Evolution (DE-588)4071050-6 s Genetik (DE-588)4071711-2 s b DE-604 Charlesworth, Dorothy ca. 20. Jh. (DE-588)1309681058 aut HBZ Datenaustausch application/pdf http://bvbr.bib-bvb.de:8991/F?func=service&doc_library=BVB01&local_base=BVB01&doc_number=025298491&sequence=000001&line_number=0001&func_code=DB_RECORDS&service_type=MEDIA Inhaltsverzeichnis |
spellingShingle | Charlesworth, Brian 1945- Charlesworth, Dorothy ca. 20. Jh Elements of evolutionary genetics Evolutionary genetics Genetik (DE-588)4071711-2 gnd Evolution (DE-588)4071050-6 gnd |
subject_GND | (DE-588)4071711-2 (DE-588)4071050-6 |
title | Elements of evolutionary genetics |
title_auth | Elements of evolutionary genetics |
title_exact_search | Elements of evolutionary genetics |
title_full | Elements of evolutionary genetics Brian Charlesworth and Deborah Charlesworth |
title_fullStr | Elements of evolutionary genetics Brian Charlesworth and Deborah Charlesworth |
title_full_unstemmed | Elements of evolutionary genetics Brian Charlesworth and Deborah Charlesworth |
title_short | Elements of evolutionary genetics |
title_sort | elements of evolutionary genetics |
topic | Evolutionary genetics Genetik (DE-588)4071711-2 gnd Evolution (DE-588)4071050-6 gnd |
topic_facet | Evolutionary genetics Genetik Evolution |
url | http://bvbr.bib-bvb.de:8991/F?func=service&doc_library=BVB01&local_base=BVB01&doc_number=025298491&sequence=000001&line_number=0001&func_code=DB_RECORDS&service_type=MEDIA |
work_keys_str_mv | AT charlesworthbrian elementsofevolutionarygenetics AT charlesworthdorothy elementsofevolutionarygenetics |