Geoscience Reference
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possible states. Each nucleotide can contain either a purine (state
þ
1) or
pyrimidine (state
1) base. The genome of each haploid individual is then
represented by a sequence of L sites. Each individual i in a population
consisting of J individuals is represented as (S i 1 , S i 2 ,
, S i L ), where S i u is the
uth site in the genome of individual i. The genetic similarity between individ-
ual i and individual j can be defined as
...
L X
L
1
q ij
S i u S j u ;
¼
ð
1
Þ
u ¼ 1
with q ij
1, 1] where 1 means the two individuals are genetically identically.
The genetic similarity in Eq. (1) can be written in terms of the fraction of
identical sites (f ij )
2
[
¼
1
L Lf ij
q ij
f ij
2f ij
¼
L 1
1
:
ð
2
Þ
and f ij
is:
1
þ
q ij
f ij
¼
:
ð
3
Þ
2
Each nucleotide in the offspring is inherited at random, thus ignoring linkage
between neighbouring nucleotides, but with a small probability of error
determined by the mutation rate. Say that the individual k inherited the
nucleotide in site u from its parent G(k): what is the probability that k will
have exactly the same nucleotide (i.e.
1) as G(k)? The probability of
no mutation and mutation in site u, respectively, is:
þ
1or
8
<
PS G ðÞ
¼
S u
¼
1
m
u
;
u
ð
4
Þ
:
PS G ðÞ
S u
u
¼
¼ m
:
u
In order to track divergence in the initial population with J individuals, we
have to calculate at each interaction event the similarity values between the
parents of the offspring k (i.e. G 1 (k)andG 2 (k) ) and each individual j in the
population. Which is the expected fraction of nucleotides in the offspring k
shared with each individual j in the population (E[f kj ])? If we assume the
same mutation rate among nucleotides,
i
i
i
m
1 ¼ m
2 ¼ ... m
L ¼ m
, then from (4)
this expected fraction is:
þ
PS G 1 ðÞ
u
1
2
f G 1 ðÞ j PS G 1 ðÞ
u
S u
f G 1 ðÞ j
S u
Ef kj
½
¼
¼
1
¼
ð
5
Þ
þ
PS G 2 ðÞ
u
1
2
f G 2 ðÞ j PS G 2 ðÞ
u
S u
f G 2 ðÞ j
S u
þ
¼
1
¼
and after substituting (4) in (5) gives:
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