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Population Genetics II: Drift, Effective Population Size, and Coalescent Thinking

Manual: General · Subject: Evolutionary Biology

Examine stochastic evolution, sampling variance, and the inferential framework of genealogies and coalescence.

Random Genetic Drift

Stochastic change

Drift is random allele-frequency change due to finite sampling of gametes or individuals. It is strongest in small populations, can fix neutral alleles by chance, and can overpower weak selection when ∣s∣≲1/Ne|s| \lesssim 1/N_e, where NeN_e is effective population size.

Effective population size

Effective population size is the size of an idealized population that would experience the same degree of drift as the real population. It is often smaller than census size because of unequal sex ratios, fluctuating sizes, variance in reproductive success, and selection at linked sites.

⚠️

Interpretive caution

NeN_e is not simply the number of individuals alive. It is a parameter summarizing the genetic consequences of demographic and reproductive structure.

Coalescent theory

Coalescent models trace sampled alleles backward in time to common ancestors. They provide a powerful way to infer demographic history, recombination, selection, and population structure from sequence data. Under neutrality, the expected time to coalescence scales with NeN_e.

Selection versus drift

Selection

  • Directional and environment-dependent.
  • Acts on fitness differences and can be predictable.

Drift

  • Stochastic and stronger in small populations.
  • Can fix or lose alleles regardless of effect.

Which population is most affected by drift, all else equal?

Give one reason effective population size can be much smaller than census size.

Using coalescent logic in inference

  1. 1

    Sample aligned sequences from the population of interest.

  2. 2

    Estimate genealogical relationships and summary statistics such as diversity and linkage patterns.

  3. 3

    Fit neutral or demographic models to the data.

  4. 4

    Test whether the observed patterns require selection, population structure, or growth/bottlenecks.

  5. 5

    Compare competing models using likelihood or Bayesian approaches.

Why is coalescent theory especially useful in genomics?

What does a selective sweep do to local genetic variation?