Part I
Source: Mather, K. 1963. The Measurement of Linkage in Heredity. John
Wiley & Sons. New York. pgs 44-55.
Estimation of 'r', the recombination faction of two
linked loci should have the most precise estimate possible.
Precision is measured by the smallest standard of error
of r Maximum likelihood estimation of r is the most
precise estimate. We use maximum likelihood to find
the estimate of r that best fits the observed data.
We maximize the term:
Likelihood = L = n! (m1)a1
(m2)a2....(mi)ai
Likelihood = L = a1!a2!...ai!
"n" is the total number of observations and the a
's
are the observed numbers of each class. The m
's
are the expected proportions determined by r. The logarithm of the above
expression is at a maximum when the expression is at a maximum.
log(L) = C + a2 log m1 + a2 log m2 + ai log mi
C is the log of n!
which is a constant.
C is the log of a1! a2!...ai!
The derivative of a constant is zero. We take the derivative of r with
respect to r and set the derivative equal to zero to find the maximum
value of r.
Let us assume that we have already conducted the X
test for linkage and found the X
value to be significant. Now we want to estimate r. The cross was:
pt x pt
PT pt
This is a testcross with one parent a double heterozygote for the P
and T genes in coupling phase linkage.
| classes |
PpTt |
PPtt |
ppTT |
pptt |
Total |
| Observed |
191 |
37 |
36 |
203 |
467 |
| Expected |
n(1-r)
2 |
nr
2 |
nr
2 |
n(1-r)
2 |
n |
In the absence of linkage, we would expect a 1:1:1:1
ratio.
| |
1/4
PT |
1/4Pt |
1/4pT |
1/4pt |
| 1pt |
1/4PpTt |
1/4Pptt |
1/4ppTt |
1/4pptt |
However, the observed results show an excess of the non-recombinant
PT and pt classes and a deficiency of the recombinant classes PT and
pT. How do we get the expected ratios?
| class |
expected |
class |
expected |
| PT |
n(1-r)
2 |
pT |
nr
2 |
| Pt |
nr
2 |
pt |
n(1-r)
2 |
| Total |
n(1-r+r)
2 |
|
n(r+1-r)
2 |
| |
=
n
= 2 |
|
=
n
= 2 |