Double
crossovers do not occur as often as one would expect, particularly if the
distance between markers is fairly short. In theory, the probability of having a
double crossover between any three markers is the probability of the two single
crossovers. Consider the following map, showing percent recombination.
lz 10.7 su 14.8 gl
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The
theoretical probability of a DCO would be (.107)(.148) = .0158 or 1.58%. In the
population of 740 individuals that this map was based on, this would give
(.0158)(740) = 11.69, or rounding up, 12 DCOs.
However,
in an experiment where the actual frequency of double crossovers was calculated,
it was found to be 6/740 = .008 or 0.8%. The
term "coefficient of coincidence" describes the ratio of
observed to expected double crossovers. In this case it would be: c of c = 6/12
= .50 or 50%. This means that only half as many DCOs were observed as would be
expected if crossovers in one interval had no effect on crossovers in the next
interval. Obviously, having a crossover in one interval did interfere with
having a crossover in the next interval.
The
term "interference" describes the following relationship:
1
- coefficient of coincidence. Thus, for these data, the interference is 1 - 0.5
= 0.5. Interference usually increases as the distance between two markers
becomes smaller, until a point where double crossing over does not occur. In
this case, the coefficient of coincidence = 0 and interference = 1. This
distance is usually about 10 % recombination. When the distance between two
markers is greater than 45 % recombination, then the coefficient of coincidence
= 1 and interference is 0. In other words, you would see (i) complete
interference at 10% recombination and below, (ii) no interference at 45 %
recombination and above, and (iii) steadily decreasing values of interference as
you went from 10 to 45 % recombination.
What
are the implications?
In medium density genome map construction, you are usually dealing with average
distances in the 5 to 10 cM range, so interference should be complete and there
should be very few double crossovers between markers. Relative to % recombination and cM, at about 10%
recombination and below, the % recombination value = cM; from 10% and upward,
the cM value will be larger than the % recombination value. The reason is that
in making the % recombination to cM conversion, you try to take into account an
average number of DCOs. As we've shown, DCOs can give you "parental"
combinations of markers and thus cause you to underestimate the number of
crossover events that actually occurred.
For example, consider the following percentages of recombination and the
corresponding Kosambi cM values:
Kosambi
cM = (0.25)ln ((1+ 2(observed recombination)/1-2 (observed recombination)).
|
%
recombination |
Kosambi
cM |
|
10 |
10.1 |
|
20 |
21.1 |
|
30 |
34.6 |