Wednesday, 4 May 2016

evolution - Variance in Fst in the infinite island model


The most famous result in the study of structured populations come from Sewall Wright. He showed that in an island model, where each subpopulation is of size $N$ and the migration rate is $m$, then the pairwise $F_{ST}$ is


$$F_{ST} = \frac{1}{4Nm+1}$$


This equation gives the expected $F_{ST}$. Because populations are finite in size ($N$), genetic drift yield this value to vary.


What is the variance in $F_{ST}$ in the infinite island model?




References


evolution in mendelian population is the original paper who derived this result from Sewall Wright.


Indirect measures of gene flow and migration: FST≠$\frac{1}{4Nm+1}$ is an influential paper in the field.


GENE FLOW IN NATURAL POPULATIONS is a famous review as well.




Answer



From Lewontin and Krakauer 1973, the ratio


$$\frac{F_{ST}(d-1)}{\bar F_{ST}}$$


approximatively follows $\chi^2$ distribution of degree $k=d-1$. Here $d$ is the number of demes (number of islands), $F_{ST}$ is the random variable of the $\chi^2$ distribution and $\bar F_{ST}$ is the average $F_{ST}$ that is $\bar F_{ST} = \frac{\sum F_{ST}}{n}$, where $n$ is the number of $F_{ST}$ values.


The variance of a $\chi^2$ distribution is $2k$, therefore


$$var\left(\frac{F_{ST}(d-1)}{\bar F_{ST}}\right) = 2d-2$$


Taking $\frac{d-1}{\bar F_{ST}}$ out of the ratio, the variance of $F_{ST}$ becomes


$$var(F_{ST})=\left(\frac{d-1}{\bar F_{ST}}\right)^2(2d-2)$$


, which simplifies into


$$var(F_{ST}) = \frac{2(d-1)^3}{\bar F_{ST}^2}$$





The above expression is probably the most interesting result but one could go further and express the variance independently from the mean (by replacing $\bar F_{ST}$ by Slatkin 1991 expectation for $\bar F_{ST}$ in a finite island. It yields to


$$var(F_{ST}) = \frac{2(d-1)^3}{\left(\frac{1}{1+4Nm(\frac{d}{d-1})^2}\right)^2}$$


, which again "simplifies" into


$$var(F_{ST}) = \frac{2 \left(4 d^2 m N+d^2-2 d+1\right)^2}{d-1}$$


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