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Moran process : ウィキペディア英語版
Moran process
A Moran process or Moran model is a simple stochastic process used in biology to describe finite populations. The process is named after Patrick Moran, who first proposed the model in 1958. It can be used to model variety-increasing processes such as mutation as well as variety-reducing effects such as genetic drift and natural selection. The process can describe the probabilistic dynamics in a finite population of constant size ''N'' in which two alleles A and B are competing for dominance. The two alleles are considered to be true replicators (i.e. entities that make copies of themselves).
In each time step a random individual (which is of either type A or B) is chosen for reproduction and a random individual is chosen for death; thus ensuring that the population size remains constant. To model selection, one type has to have a higher fitness and is thus more likely to be chosen for reproduction.
The same individual can be chosen for death and for reproduction in the same step.
==Neutral drift==
Neutral drift is the idea that a neutral mutation can spread throughout a population, so that eventually the original allele is lost. A neutral mutation does not bring any fitness advantage or disadvantage to its bearer. The simple case of the Moran process can describe this phenomenon.
If the number of A individuals is given by ''i'' then the Moran process is defined on the state space . Since the number of A individuals can change at most by one at each time step, a transition exists only between state ''i'' and state and . Thus the transition matrix of the stochastic process is tri-diagonal in shape and the transition probabilities are
:\begin
P_&=1\\
P_ &= \frac \frac\\
P_ &= 1- P_ - P_\\
P_ &= \frac \frac\\
P_&=1.
\end
The entry P_ denotes the probability to go from state ''i'' to state ''j''. To understand the formulas for the transition probabilities one has to look at the definition of the process which states that always one individual will be chosen for reproduction and one is chosen for death. Once the A individuals have died out, they will never be reintroduced into the population since the process does not model mutations (A cannot be reintroduced into the population once it has died out and ''vice versa'') and thus P_=1. For the same reason the population of A individuals will always stay ''N'' once they have reached that number and taken over the population and thus P_=1. The states 0 and ''N'' are called ''absorbing'' while the states are called ''transient''. The intermediate transition probabilities can be explained by considering the first term to be the probability to choose the individual whose abundance will increase by one and the second term the probability to choose the other type for death. Obviously, if the same type is chosen for reproduction and for death, then the abundance of one type does not change.
Eventually the population will reach one of the absorbing states and then stay there forever. In the transient states, random fluctuations will occur but eventually the population of A will either go extinct or reach fixation. This is one of the most important differences to deterministic processes which cannot model random events. The expected value and the variance of the number of A individuals at timepoint ''t'' can be computed when an initial state is given:
: \begin
E(= i ) &= i \\
Var(X(t)|X(0) = i) &= \tfrac \left(1-\tfrac \right ) \frac \right )^t}}
\end
: \begin
E (| X(t-1) = i ) &= (i-1)P_ + iP_ + (i+1)P_\\
&= 2ip(1-p) + i(p^2 + (1-p)^2) \\
&= i.
\end
Writing Y = X(t) and Z = X(t-1), and applying the law of total expectation, E() = E. Applying the argument repeatedly gives E() = E(), or E(= i ) = i.
For the variance the calculation runs as follows. Writing V_t = Var(X(t)|X(0) = i), we have
: \begin
V_1 &= E \left(= i \right ) - E()^2 \\
&= (i-1)^2p(1-p) + i^2 \left (p^2+(1-p)^2 \right ) + (i+1)^2p(1-p) - i^2 \\
&= 2p(1-p)
\end
For all , (X(t)|X(t-1) = i) and (X(1)|X(0) = i) are identically distributed, so their variances are equal. Writing as before Y = X(t) and Z = X(t-1), and applying the law of total variance,
: \begin
Var(Y) &= E() + Var(E()) \\
&= E \left ((\frac \right) \left(1-\frac \right ) \right ) + Var(Z)\\
&= \left(\frac \right ) \left (1-\frac \right) + \left(1-\frac\right)Var(Z).
\end
If X(0) = i, we obtain
:V_t = V_1 + \left (1-\frac \right)V_.
Rewriting this equation as
: V_t - \frac} = \left (1-\frac \right )\left(V_-\frac}\right) = \left (1-\frac \right)^ \left(V_1-\frac}\right)
yields
: V_t = V_1 \frac \right)^t}}
as desired.
----
}}
The probability of A to reach fixation is called ''fixation probability''. For the simple Moran process this probability is
Since all individuals have the same fitness, they also have the same chance of becoming the ancestor of the whole population; this probability is and thus the sum of all ''i'' probabilities (for all A individuals) is just The mean time to absorption starting in state ''i'' is given by
: k_i = N \left(\sum_^ \frac + \sum_^ \frac \right )

The variable y_i^ = k_^j- k_^j is used and the equation becomes
: \begin
y_^ &= y_i^ -\frac \\ \\
\sum_^m y_i^ &= (k_^j- k_^j) + (k_^j- k_^j) + \cdots + (k_^j- k_^j) + (k_^j- k_^j) \\
&= k_^j - k_^j \\
\sum_^m y_i^ &= k_^j \\ \\
y_1^ &= (k_^j- k_^j) = k_^j \\
y_2^ &= y_1^ -\frac = k_1^ -\frac \\
y_3^ &= k_1^ -\frac -\frac \\
& \vdots \\
y_i^ &= k_1^ -\sum_^ \frac = \begin k_1^j & j \geq i\\ k_1^j - \frac & j \leq i \end \\ \\
k_i^j &= \sum_^i y_m^ = \begin i \cdot k_1^j & j \geq i\\ i \cdot k_1^j - \frac & j \leq i \end
\end
Knowing that k_N^j = 0 and
:q_j = P_=\frac \frac
we can calculate k_1^j:
: \begin
k_N^j = \sum_^m y_i^ = N \cdot k_1^j &- \frac = 0 \\
k_1^j &= \frac
\end
Therefore
: k_i^j = \begin \frac \cdot k_j^j & j \geq i\\ \frac \cdot k_j^j & j \leq i\end
with k_j^j = N. Now , the total time until fixation starting from state ''i'', can be calculated
: \begin
k_i = \sum_^k_i^j &= \sum_^k_i^j + \sum_^k_i^j \\
&= \sum_^N \frac + \sum_^N \frac
\end
----
}}
For large ''N'' the approximation
: \lim_ k_i \approx -N^2 \left((1-x_i) \ln(1-x_i) + x_i \ln(x_i) \right )
holds.

抄文引用元・出典: フリー百科事典『 ウィキペディア(Wikipedia)
ウィキペディアで「Moran process」の詳細全文を読む



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