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 |
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.
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 X2 test for linkage and found the X2 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) |
nr |
nr |
n(1-r) |
n |
| 2 | 2 | 2 | 2 |
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) |
pT | nr | ||
| 2 | 2 | ||||
| Pt | nr |
pt | n(1-r) | ||
| 2 | 2 | ||||
| Total | n(1-r+r) |
n(r+1-r) | |||
| 2 | 2 | ||||
= |
n |
= |
n |
||
2 |
2 |
||||