3 Neutral Evolution in One- and Two-Locus Systems

Nature selection, neural models

genetic drift: random fluctuations in allele frequencies that necessarily result from sampling finite numbers of gametes in each generation. Sample size, time scale.

3.1 The Wright-Fisher Model

Single finite populations of constant size within which mating is random; other Variant models: separate sexes; family size; generation overlap

Its roots trace to Fisher (1922) and Wright (1931).

Assume:

  • a diploid population with a fixed number (N) of monoecious (hermaphroditic, 雌雄同体) adults
  • random mating (self-fertilization included)
  • discrete generations

Consider a locus with two alleles B and b, without selection advantage:

the probability (\(P_{ij}\)) that i copies of allele B in generation t and j copies of allele B in generation t+1 follows a binomial distribution:

\[P_{ij} = \begin{pmatrix}2N \\j \end{pmatrix}(i/2N)^j[1-(i/2N)]^{2N-j}\]

Letting \(\boldsymbol{P} \in \boldsymbol{R}^{(2N+1)*(2N+1)}\) that

\[\boldsymbol{x}(t+1)=\boldsymbol{x}(t)\boldsymbol{P}\] , where \(\boldsymbol{x}(t) \in \boldsymbol{R}^{1*(2N+1)}\), the probabilities that the allele is present in i=0,1,…,2N copies in generation t

\[x(t) = x(0)P^t\]

Markov chain

Absorbing states: lost or fixed

transition-matrix; diffusion approximation

Moran model

3.2 Loss of heterozygosity by random genetic drift

3.3 Probabilities and times to fixation or loss

3.4 The age of a neutral allele

3.5 Allele-frequency divergence among populations

3.6 Buri’s experiment

3.7 Higher-order allele-frequency moments

3.8 Linkage disequilibrium

3.9 Mutation-drift equilibrium

3.10 The detailed structure of neutral variation

3.10.1 The infinite-alleles model and the associated allele-frequency spectrum

3.10.2 The infinite-sites model and the associated site-frequency spectrum

3.11 The genealogical structure of a population

3.12 Mutation-migration-drift equilibrium

3.12.1 Quantifying population structure: Fst

3.12.2 Mutation-migration-drift equilirium values of Fst