The evolutionary models discussed in the previous chapters assume an infinite population that can be divided in to infinitessimal parts. Finite populations can also be studied using a model called a Moran Process (first described in 1958).

Consider a population of two types of fixed size $N$. This can be represented as a vector of the form: $(i, N-i)$ where $i\geq 0$ represents the number of individuals of the first type.

The term **neutral** drift refers to the fact that the two types reproduce at the same rate.

The Moran process is as follows:

- At a given time step: select a random individual for reproduction and a random individual for elimination
- The eliminated individual is replaced by a new individual of the same type as the individual chosen for reproduction.
- Proceed to the next time step.
- The process terminates when there is only one type of individual in the population.

Here is some simple Python code that simulates such a Process assuming an initial population of $(3, 3)$:

In [1]:

```
import numpy as np
import matplotlib.pyplot as plt
%matplotlib inline
def neutral_moran(N, i=1, seed=0):
"""
Return the population counts for the Moran process with neutral drift.
"""
population = [0 for _ in range(i)] + [1 for _ in range(N - i)]
counts = [(population.count(0), population.count(1))]
np.random.seed(seed)
while len(set(population)) == 2:
reproduce_index = np.random.randint(N)
eliminate_index = np.random.randint(N)
population[eliminate_index] = population[reproduce_index]
counts.append((population.count(0), population.count(1)))
return counts
N = 6
plt.plot(neutral_moran(N=N, i=3, seed=6));
```

In [2]:

```
def neutral_fixation(N, i=None, repetitions=10):
"""
Repeat the neutral Moran process and calculate the fixation probability
"""
fixation_count = 0
for seed in range(repetitions):
final_counts = neutral_moran(N=N, i=i, seed=seed)
if final_counts[-1][0] > 0:
fixation_count += 1
return fixation_count / repetitions
```

In [3]:

```
probabilities = [neutral_fixation(N, i=i, repetitions=500) for i in range(1, N)]
plt.scatter(range(1, N), probabilities)
plt.xlabel("$i$")
plt.ylabel("$x_i$");
```

We see that as the initial population starts with more of a given type, the chance that that type "takes over" (becomes fixed) grows.

This Moran Process is a specific case of a Markov Process:

- A given state of the system can be described by a single integer $0\leq i\leq N$;
The state to state transition probabilities are given by:

$$ \begin{aligned} p_{i, i-1}&=\frac{i(N - i)}{N^2}\\ p_{i, i+1}&=\frac{i(N - i)}{N^2}\\ p_{i, i}&=1 - p_{i, i-1} - p_{i, i+1} \end{aligned} $$

We also have two absorbing states (when the Moran process ends):

$$p_{00}=1\qquad p_{0i}=0\text{ for all }i>0$$

$$ p_{NN}=1\qquad p_{Ni}=0\text{ for all } N>i $$

these transitions can be represented as a matrix. Here for example is the matrix for $N=6$:

In [4]:

```
N = 6
p = np.zeros((N + 1, N + 1))
p[0, 0] = 1
p[N, N] = 1
for i in range(1, N):
for j in [i - 1, i + 1]:
p[i, j] = i * (N - i) / (N ** 2)
p[i, i] = 1 - sum(p[i, :])
p.round(2)
```

Out[4]:

The above corresponds to a particular type of Markov process called a Birth-Death process

A birth death process is a Markov process with the following properties:

- $p_{i,i+1}+p_{i,i-1}\leq 1$
- $p_{ii}=1-p_{i,i+1}-p_{i,i-1}$
- $p_{00}=1$ and $p_{NN}=1$

Thus we have two absorbing states: $\{0, N\}$. Let us denote by $x_i$ the probability of being in $state$ $i$ and eventually reaching state $N$.

We have the following linear system:

\begin{align} x_0&=0\\ x_i&=p_{i,i-1}x_{i-1}+p_{ii}x_i+p_{i,i+1}x_{i+1}\text{ for all }0< i< N-1\\ x_N&=1\\ \end{align}Given a birth death process as defined above, the fixation probability $x_i$ is given by:

$$x_i=\frac{1+\sum_{j=1}^{i-1}\prod_{k=1}^j\gamma_k}{1+\sum_{j=1}^{N-1}\prod_{k=1}^j\gamma_k}$$where:

$$ \gamma_k = \frac{p_{k,k-1}}{p_{k,k+1}} $$We have:

$$ \begin{aligned} p_{i,i+1}x_{i+1} & = -p_{i,i-1}x_{i-1} + x_i(1 - p_{ii}) \\ p_{i,i+1}x_{i+1} & = p_{i,i-1}(x_{i} - x_{i-1}) + x_ip_{i,i+1} \\ x_{i+1} - x_i & = \frac{p_{i, i-1}}{p_{i, i+1}}(x_i-x_{i-1})=\gamma_i(x_i-x_{i-1}) \end{aligned} $$We observe that:

$$ \begin{aligned} x_2 - x_1 &= \gamma_1(x_1-x_{0})=\gamma_1x_1\\ x_3 - x_2 &= \gamma_2(x_2-x_1)=\gamma_2\gamma_1x_1\\ x_4 - x_3 &= \gamma_3(x_3-x_2)=\gamma_3\gamma_2\gamma_1x_1\\ &\; \vdots & \\ x_{i+1} - x_i &= \gamma_i(x_i-x_{i-1})=\prod_{k=1}^i\gamma_kx_1\\ &\; \vdots & \\ x_{N} - x_{N-1} &= \gamma_{N-1}(x_{N-1}-x_{N-2})=\prod_{k=1}^{N-1}\gamma_kx_1\\ \end{aligned} $$thus we have:

$$x_i=\sum_{j=0}^{i-1}x_{j+1}-x_j=\left(1+\sum_{j=1}^{i-1}\prod_{k=1}^j\gamma_k\right)x_1$$we complete the proof by solving the following equation to obtain $x_1$:

$$x_N=1=\left(1+\sum_{j=1}^{N-1}\prod_{k=1}^j\gamma_k\right)x_1$$In the case of neutral drift (considered above) we have:

$$p_{i,i-1}=p_{i,i+1}$$thus:

$$ \gamma_i=1 $$so:

$$ x_i=\frac{1+\sum_{j=1}^{i-1}\prod_{k=1}^j\gamma_k}{1+\sum_{j=1}^{N-1}\prod_{k=1}^j\gamma_k}=\frac{1+i-1}{1+N-1}=\frac{i}{N} $$In [5]:

```
probabilities = [neutral_fixation(N, i=i, repetitions=500) for i in range(1, N)]
plt.scatter(range(1, N), probabilities, label="Simulated")
plt.plot(range(1, N), [i / N for i in range(1, N)], label="Theoretic: $i/N$", linestyle="dashed")
plt.xlabel("$i$")
plt.ylabel("$x_i$")
plt.legend();
```

The fixation probability in a Moran process is the probability that a give type starting with $i=1$ individuals takes over an entire population. We denote the fixation probabilities of the first/second type as $\rho_1$ and $\rho_2$ respectively and we have:

$$ \rho_1=x_1 $$$$ \rho_2=1-x_{N-1} $$We will now consider a Moran process on a game:

Consider a matrix $A\in\mathbb{R}^{m\times n}$ representing a game with two strategies.

$$ A= \begin{pmatrix} a & b\\ c & d \end{pmatrix} $$The Moran process is as follows:

- At a given time step: all individuals play all other individuals.
- Obtain their fitness as given by the game.
- Randomly select an individual proportional to their fitness as an individual to be reproduced
- Uniformly select an individual to be replaced
- Proceed to the next time step.
- The process terminates when there is only one type of individual in the population.

Assuming $i$ individuals of the first type, the fitness of both types is given respectively by:

$$f_{1i}=\frac{a(i-1)+b(N-i)}{N-1}$$$$f_{2i}=\frac{c(i)+d(N-i-1)}{N-1}$$The transition probabilities are then given by:

$$p_{i,i+1}=\frac{if_{1i}}{if_{1i} + (N-i)f_{2i}}\frac{N-i}{N}$$$$p_{i,i-1}=\frac{(N-i)f_{2i}}{if_{1i} + (N-i)f_{2i}}\frac{i}{N}$$which gives:

$$\gamma_i=\frac{f_{2i}}{f_{1i}}$$thus:

$$ x_i=\frac{1+\sum_{j=1}^{i-1}\prod_{k=1}^j\gamma_k}{1+\sum_{j=1}^{N-1}\prod_{k=1}^j\gamma_k} $$Here is some code to carry out this calculation:

In [6]:

```
def theoretic_fixation(N, game, i=1):
"""
Calculate x_i as given by the above formula
"""
f_ones = np.array([(game[0, 0] * (i - 1) + game[0, 1] * (N - i)) / (N - 1) for i in range(1, N)])
f_twos = np.array([(game[1, 0] * i + game[1, 1] * (N - i - 1)) / (N - 1) for i in range(1, N)])
gammas = f_twos / f_ones
return (1 + np.sum(np.cumprod(gammas[:i-1]))) / (1 + np.sum(np.cumprod(gammas)))
```

Here is an example of calculating $x_1$ for the following game for $N=4$:

$$ A = \begin{pmatrix} 4 & 1\\ 1 & 4 \end{pmatrix} $$In [7]:

```
A = np.array([[4, 1],
[1, 4]])
theoretic_fixation(N=4, i=1, game=A)
```

Out[7]:

Applying the theorem gives:

$$ \begin{aligned} f_{1i}&=\frac{4(i - 1) + 4 - i}{3} = \frac{4i-4+4-i}{3}=i\\ f_{2i}&=\frac{i + 4(3 - i)}{3} = \frac{12-3i}{3}=4-i \end{aligned} $$$$ \gamma_i = \frac{f_{2i}}{f_{1i}}=\frac{4-i}{i}=\frac{4}{i}-1 $$Thus:

$$ \begin{aligned} x_1 & =\frac{1 + \sum_{j=1}^{0}\prod_{k=1}^{j}\gamma_k}{1 + \sum_{j=1}^{4 - 1}\prod_{k=1}^{j}\gamma_k}\\ & =\frac{1}{1 + \sum_{j=1}^{3}\prod_{k=1}^{j}\gamma_k}\\ & =\frac{1}{1 + \gamma_1 + \gamma_1\times \gamma_2 + \gamma_1 \times \gamma_2 \times \gamma_3}\\ & =\frac{1}{1+3+3\times 1 + 3 \times 1\times \frac{1}{3}} = \frac{1}{1 + 3 + 3 + 1}=\frac{1}{8}\\ \end{aligned} $$Here is some code to simulate a Moran process.

In [8]:

```
def moran(N, game, i=1, seed=0):
"""
Return the population counts for
the Moran process on a 2 by 2 game
"""
population = [0 for _ in range(i)] + [1 for _ in range(N - i)]
counts = [(population.count(0), population.count(1))]
np.random.seed(seed)
while len(set(population)) == 2:
scores = []
for i, player in enumerate(population):
total = 0
for j, opponent in enumerate(population):
if i != j:
total += game[player, opponent]
scores.append(total)
total_score = sum(scores)
probabilities = [score / total_score for score in scores]
reproduce_index = np.random.choice(range(N), p=probabilities)
eliminate_index = np.random.randint(N)
population[eliminate_index] = population[reproduce_index]
counts.append((population.count(0), population.count(1)))
return counts
def fixation(N, game, i=None, repetitions=10):
"""
Repeat the Moran process and calculate the fixation probability
"""
fixation_count = 0
for seed in range(repetitions):
final_counts = moran(N=N, i=i, game=game, seed=seed)
if final_counts[-1][0] > 0:
fixation_count += 1
return fixation_count / repetitions
```

In [9]:

```
N = 8
plt.plot(moran(N=N, i=1, seed=44, game=A));
```

Here is how the fixation probabilities vary for different initial populations:

In [10]:

```
probabilities = [fixation(N, i=i, game=A, repetitions=500) for i in range(1, N)]
plt.scatter(range(1, N), probabilities, label="Simulated")
plt.plot(range(1, N), [i / N for i in range(1, N)], label="Neutral: $i/N$", linestyle="dashed")
plt.plot(range(1, N), [theoretic_fixation(N=N, i=i, game=A) for i in range(1, N)], label="Theoretic")
plt.xlabel("$i$")
plt.ylabel("$x_i$")
plt.legend();
```

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