Polymorphic limit (Wright-Fisher model)

guillefix 4th November 2016 at 2:43pm

(See Arrival of the frequent for context)

If NLμ1NL\mu \gg 1, the population naturally spreads over different genotypes, a regime called the polymorphic limit. See Polymorphic limit (Wright-Fisher model) tiddler for more.

To model neutral exploration, we let 1+sp=δpq1+s_p = \delta_{pq}, where δpq \delta_{pq} is a Kronecker delta, so that only qq has some fitness, and all other phenotypes have 00 fitness, and so, even if a mutation produces them, no offspring can inherit from them. At every generation, all offspring inherits from Nq\mathcal{N}_q only, and thus the population can only spread by mutations over a single generation jump, and it is most likely to stay mostly within Nq\mathcal{N}_q, if NN is large enough.

We should note that equations, like Eq.3 would be the same, even though we assumed that all the individuals are in Nq\mathcal{N}_q, because, as 1+sp=δpq1+s_p = \delta_{pq}, all the selection weight is in Nq\mathcal{N}_q, which produces the same results. More precisely, in the expression i=1NΦp~(gi,si)N(1+si)j=1N(1+sj)\sum_{i=1}^N \tilde{\Phi_p}(g_i, s_i) \frac{N(1+s_i)}{\sum_{j=1}^N (1+s_j)} only NN' (the number of individuals in Nq\mathcal{N}_q) elements are non-00 in the sum and so in the mean-field approx (where we assume Φp~(gi,si)\tilde{\Phi_p}(g_i, s_i) is constant) the NN' from the sum cancels the j=1N(1+sj)=j=1Nδpq=N\sum_{j=1}^N (1+s_j) = \sum_{j=1}^N \delta_{pq} = N' from the denominator, leaving a NN on the top.

In the mean-field approximation the expected number of individuals with phenotype pp produced per generation is now independent of time, and given by Eq. 3. (we thus simply call mp(t)=mpm_p(t) = m_p), under the corresponding assumptions, because even if not all of the population are in Nq\mathcal{N}_q, the assumption of fitness, we've made gives selective weight only to those in Nq\mathcal{N}_q (see Wright-Fisher model).

As we said above, the number of individuals with genotype pp (p-type) will follow a binomial distribution, with probability mp/Nm_p/N of success (getting p-type offspring), and number of trials NN, and therefore the probability to get at least one such individual is:

P(at least on p-type offpsring)=1P(no p-type offspring)=1(1mp/N)N1empP(\text{at least on p-type offpsring}) = 1 - P(\text{no p-type offspring}) = 1 - (1 - m_p/N)^N \approx 1 - e^{-m_p}

After TT generations, we have run the Bernoulli trial TNTN times, and thus the number of p-type individuals we have gotten, summed over all the TT generation also follows a Binomial distribution, but with NTNT samples, and same probability. Thus

P(at least on p-type offpsring over T generations)1empTP(\text{at least on p-type offpsring over T generations}) \approx 1 - e^{-m_p T}

Thus, the time when {{the probability of having discovered a p-type individual (produced a p-type offspring)} is α\alpha} is found by:

α=1empT\alpha = 1 - e^{-m_p T}

empT=1αe^{-m_p T} = 1- \alpha

mpT=ln(1α)-m_p T = \ln{(1- \alpha)}

T=ln(1α)NLμΦpq T = \frac{-\ln{(1- \alpha)}}{N L\mu \Phi_{pq}}

Eq. 4

Where we used Eq. 3 in Arrival of the frequent.