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 White
Paper
Significant Genetic Relationships
© 1998, Dr. John B.
Armstrong, Ph.D.
CompuPed Home Page
Breeding article by Dr. J. Armstrong
How would you determine the impact of a famous Champion on his
breed?
A famous champion who has won many shows and earned many titles
may be (or have been), quite popular as a stud and may have sired
more winning progeney than other contemporary males, but that does
not guarantee that he will have more impact five or ten generations
down the line than another dog who was bred only two or three times.
So in order to answer this question we will need to do some genetic
calculations.
Percent Contribution
If sufficient data is available, one way of determining the
significance of an ancestor is to calculate his genetic contribution
to his descendants. The percentage of contribution (aka
percentage of blood) is determined by the way genes
are passed from the parents to the progeny. An individual inherits
one set of chromosomes, and the genes they carry, from its sire and
a second, homologous (equivalent) set from its dam. Thus each parent
always passes on (contributes) 50% of its genes to each of its
offspring and it therefore seems reasonable to expect that 25% of an
individual's genes will come from each grandparent, 12.5% from each
great-grandparent, and so on.
Each generation of ancestors in a pedigree contributes 100% of
the genes for the target animal (proband). So the percentage of
contribution from each ancestor at a given generational level is 100
divided by the number of ancestors in that generation. Since the
number of ancestors doubles in each preceding generation, the
contribution from each ancestor in each preceding generation is
halved.
However, this is not like mixing paint! Percentage of blood for
ancestors beyond the parents are probabilities, not certainties.
When a male passes on one set of his chromosomes, they will include
a random selection of the genes he inherited from both of his
parents, but there is no guarantee that the genes he passes on will
contain an equal amount from each of his parents and in fact it
rarely is. There is even a small chance (very small) that he will
pass on those genes from only one of his parents. On the other hand,
the more times that an ancestor appears in a pedigree behind
different offspring, the more likely it will be that the percentage
of genes passed down from each of its parents will be closer to 50%.
By the time we get back 10 generations, the contribution from
each of the 1024 ancestors would, in theory, amount to slightly less
than 0.1%. However, in the pedigree of the average purebred dog,
there are seldom more than 100-200 different (unique) ancestors and
some may appear 50 times or more. These are the significant
ancestors that make the major genetic contributions.
If you have a pedigree, you can calculate % contribution of any
repeats simply by multiplying the number of times each ancestor
appears in any generation by the appropriate percentage for that
generation and then add together all of the calculated percentage of
contributions from each generation. The table listed below shows the
percentage of blood inherited from each ancestor at the given
generation levels. Generation "1" is the parents.
| Genetic Contribution of Ancestors |
| Generation |
1 |
2 |
3 |
4 |
5 |
6 |
7 |
8 |
9 |
10 |
| % Contribution |
50.0 |
25.0 |
12.5 |
6.25 |
3.125 |
1.563 |
0.781 |
0.391 |
0.195 |
0.098 |
Databases exist for many breeds that will contain the data
enabling you to extend a pedigree to 10 generations or more. Manual
computation, though tedious, is still possible, but hardly
convenient. However, the CompuPed pedigree program will quickly
calculate % contribution for selected ancestors or all ancestors for
a specified number of generations, providing you with information on
which dogs have been most influential.
Is there a quick way of
determining how genetically similar two dogs are?
Suppose a breeder has two bitches (A and B) she wants to mate to
different males. After careful research the breeder identifies three
potentially suitable males (C, D and E), all of which look equally
good. The breeder hopes to get a male puppy from one litter and a
female from the other, and would like to eventually breed these
puppies to each other. The objective could be to pick the
combination that will minimize the potential inbreeding.
Alternatively, the breeder may be looking for two dogs that are not
close relatives yet have similar heritage.
One approach would be to produce hypothetical litters for all
combinations: AC, AD, AE, BC, BD and BE, and then look at the
possibilities for the second generation. There will be six if shared
grandparents are not permited, and 36 if there are no restrictions.
These potential litters could then be evaluated for inbreeding or
the % contribution of significant ancestors. This will certainly
provide the data, but is unnecessarily tedious.
The Coefficient of
Relationship
The relationship coefficient (RC) provides a way of objectively
assessing the similarity of two pedigrees by giving a number that is
a direct measure of shared ancestry. In most human populations, two
individuals picked at random would likely have a RC of 0, a brother
and sister 50% and identical twins 100%. Other relationships would
fall somewhere between 0 and 50%.
The number generated may be viewed as analogous to the %
composition, except that you are comparing two dogs instead of
looking at one. A brother and sister will have an RC value of 50% as
long as they have no ancestors that are repeated. Once ancestors
start to repeat, the individuals no longer have an inbreeding
coefficient of zero. Two sibs from a highly inbred line may have an
RC of 80% or more, and two dogs that are not sibs may have an RC
above 50%.
The formula for the RC is:
RAB = 2fAB ÷ [(1 +
FA)(1 + FB)]½
where fAB is the inbreeding coefficient of
a hypothetical litter between A and B, and FA and
FB are the inbreeding coefficients for the two
individuals, A and B.
A simpler approach to the breeder's problem would be to compute
the RCs for C vs D and E, and D vs E. This is not an easy pencil and
paper calculation. However, presented with just such a problem, it
took me about 2 minutes to obtain the three RCs with the latest
version of CompuPed. My results were RCCD = 10.4%,
RCCE = 13.4%, RCDE = 17.2%.
D and E share the most common ancestry, while C and D share the
least, and respectively so would the progeny from their two
prospective litters. To minimize inbreeding and maximize diversity,
all else being equal, C and D would be the best choice. (These
values actually all fall below the average for the breed, which is
about 23%.)
The Kinship
Coefficient
The fAB term in the RC equation is sometimes
called the "kinship coefficient" and may also be used as a measure
of the relationship between two individuals. It's computation is the
same as that of an inbreeding coefficient for a hypothetical litter
between the two dogs. (It doesn't matter if they are the same sex.)
The mean kinship (mki)
for individual i is is the average of the kinship
coefficients (fij) between i and all the
other breedable individuals in the population:
A conservation biologist would consider the individual with the
lowest mean kinship to be the most genetically valuable in
terms of maintaining diversity in the population, and would try to
favor that individual in a breeding program.
Acknowledgement. . .
We warmly thank Dr. Armstrong for giving us permission to post
this informative information on our web site. We would also like to
encourage you to vist Dr. Armstrong's very comprehensive Genetic Diversity web site at
the University of Ottawa for more information on many related
subjects.
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