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Understanding Inbred Pedigrees
By Suzon Murray
You just bought
a purebred, papered animal. You’re proud of that animal, those papers,
that pedigree. You’re thinking about breeding. Now you’ve got to do your
homework. Now it’s time to take that pedigree apart and understand what
you’ve got in terms of genetics, in terms of blood. It’s time to think
about the genetics you’re going to be producing when you cross that
pedigree with another. Questions begin to form in your mind. Should your
program include inbreeding or avoid such practices? But exactly what is
inbreeding and why is it important? And why should you pay attention to
it in your own breeding program?
Most
man-developed purebred breeds are the product of " inbreeding."
Inbreeding by definition is the mating of two animals more closely
related than the general population. In creating a breed, breeders cross
individuals with desirable characteristics to other closely related
individuals with those same characteristics to increase the regularity
with which those traits are passed on. Eventually a distinct line of
animals comes into being with set characteristics that are passed on
without fail. This is what is called prepotency, the ability to stamp
one or many traits on offspring regularly, even offspring that is the
result of crossing to animals outside the breed. Prepotency most often
(though not always) comes from a high number of genes that are
homozygous. Homozygosity is what inbreeding is all about.
OK, so what is
homozygosity anyway? Each parent of an individual contributes one set of
chromosomes to the genetic makeup of that individual. These pairs are
the map that defines all the characteristics of that individual. Each
chromosome is made up of individual genes. The genes on one set of
chromosomes match up with genes at the same location on the other set.
When the two genes are identical they are said to be " homozygous" , and
when they are different, they are " heterozygous." In purebred animals,
the starting assumption is that 50% of all an individual’s genes are
homozygous.
Genes are
either " dominant" or " recessive." In heterozygous pairs, if there is a
dominant and a recessive gene, the dominant gene is the one that is
expressed. However, if a gene pair is homozygous and dominant, you are
given a 100% chance of passing that trait on, no matter what. If you
have a recessive quality and you raise your percentage of homozygous
genes in a breed population, you raise your chances of pairing up those
recessive genes, allowing their qualities to be expressed. The rub,
unfortunately, comes in regard to these recessive genes. As you raise
the number of homozygous pairs, you raise your chances of matching
recessive genes whose traits are less than desirable (infertility,
deformities, immune-deficiencies, et cetera).
There are
schools of thought that state any amount of inbreeding is going to
decrease the " vigor " of an individual. Other schools argue that until
total homozygosity has reached 65% there is little danger. Breeders need
to come to terms with how much homozygosity they are willing to create
in order to produce the stock desired.
This is where
analyzing a pedigree comes in. There are many ways to do this.
The first to be
discussed makes a simple distinction between " inbred " and " line
bred." In this system inbreeding is defined as the crossing of very
closely related individuals (such as half-brother/sister), with line
breeding to mean the crossing of less closely related individuals (for
example, having the same great-grandparent on both sides).The
distinction is made between the two with a simple formula of "
generation numbers."
Each generation
is given a numerical value. The parents are given the number 1, the
grandparents 2, great-grandparents 3 and so on.Then for any ancestors
appearing in both the paternal and maternal sides of the pedigree, the
values are added and a total reached. If this number is below 6, the
individual is said to be inbred, if the number is between 6 and 9 then
the individual is classified line bred, beyond 9 it is considered to be
effectively out crossed.
For example:
|
|
F1 |
F2 |
F3 |
F4 |
F5 |
|
|
Beaux
Arts |
Orgueilleux |
Diviseur |
Quicko |
Gringalet |
|
|
Italienne |
|
|
Vigoureuese |
Bourbacky |
|
|
Mirza |
|
|
Irma |
Polo |
Bacchus |
|
|
Hermione |
|
|
Etincelle |
Trompeur |
|
|
Amande |
|
|
Tyronienne |
Gaulois |
Questeur |
Ultra |
|
|
Rola |
|
|
Tulipe |
Neptune |
|
|
Jonquille |
|
|
Etincelle |
Trompeur |
Bacchus |
|
|
Parfait |
|
|
Amande |
Millefore |
|
|
Verveine |
|
Fedon |
|
|
|
|
|
|
|
Hirondelle |
Arcol du
Bourg |
Diviseur |
Quicko |
Gringalet |
|
|
Italienne |
|
|
Vigoureuse |
Bourbacky |
|
|
Mirza |
|
|
Irma |
Polo |
Bacchus |
|
|
Hermione |
|
|
Etincelle |
Trompeur |
|
|
Amande |
|
|
Utopie du
Bourg |
Diviseur |
Quicko |
Gringalet |
|
|
Italienne |
|
|
Vigoureuse |
Bourbacky |
|
|
Mirza |
|
|
Oba |
Diviseur |
Quicko |
|
|
Vigoureuse |
|
|
Victoire |
Printemps |
|
|
Jonquille |
This pedigree
is fictitious, but it is an excellent example of both an inbred and a
line-bred pedigree. In looking at the numbers we see that Fedon has
Diviseur in the third generation in both the paternal and maternal
side. 3 + 3=6; therefore this individual can be considered mildly "
inbred." If Diviseur had appeared as a grand parent on both sides,
then the combined values would equal four and the degree of inbreeding
would be more acute. Then we see that Fedon has Etincelle in the third
generation on the paternal side and in the fourth generation on the
maternal side. 3 + 4=7, so Fedon is considered linebred to Etincelle.
However, we also see he has Jonquille in the fifth generation on both
sides. 5+5=10, which is greater than nine; so this pairing is
considered to be too far away to have any bearing on the pedigree.
Remember,
inbreeding indicates a higher degree of homozygosity. This method is
good to quickly determine the general degree of inbreeding in a
pedigree, but it does not give us an accurate idea of the degree of
increased homozygosity that could be present in this individual. For
that, you will need the next method.
In his work,
Systems of Breeding, Sewell Wright developed a formula to determine
the degree of homozygosity likely to be present in an inbred
individual. The number, which is calculated by this formula, is called
the coefficient of inbreeding. It is an estimate of the percentage of
homozygous pairs above and beyond the assumed 50% for a purebred
animal.
Fx=S(1/2)a+b
(1+Fa)
Wright’s
Formula states that the coefficient of inbreeding is equal to the sum
of one-half raised to a power equal to the number of generations from
the sire to the common ancestor and back to the dam times one plus the
coefficient of inbreeding, if any, of the common ancestor. In other
words, if we start with the base 50%, Wrights Formula tells us what
percentage we must add to that to have an accurate idea of the
increased homozygosity of the individual. If an animal has a
coefficient of 25% it would mean that 25% of the REMAINING 50% (or
12.5% of the total) are likely to be homozygous. So the total gene
pairs that are homozygous would be 50% + 12.5% or 62.5%.
However, if
you’re sane, you forgot your algebra shortly after you learned it.
Fear not, there’s a slightly longer means of calculating this formula
by hand that requires only addition and the use of the table below.
|
Calculating the Co-effient of inbreeding |
|
|
F1 |
F2 |
F3 |
F4 |
F5 |
F6 |
F7 |
|
F1 |
50% |
25% |
12.5% |
6.25% |
3.13% |
1.56% |
.78 |
|
F2 |
25% |
12.5% |
6.25% |
3.13% |
1.56% |
.78% |
.39% |
|
F3 |
12.5% |
6.25% |
3.13% |
1.56% |
.78% |
.39% |
.19% |
|
F4 |
6.25% |
3.13% |
1.56% |
.78% |
.39% |
.19% |
.095% |
|
F5 |
3.13% |
1.56% |
.78% |
.39% |
.19% |
.095% |
.05% |
|
F6 |
1.56% |
.78% |
.39% |
.19% |
.095% |
.05% |
.025% |
Each pedigree
is divided into generations just as in the previous method. The
parents are expressed as generation F1, the grandparents as F2, the
great-grandparents as F3 et cetera. To use the above chart, simply
identify the common ancestor(s) in a pedigree. For example: In the
pedigree used as an example previously we find Diviseur on the
paternal side in the F3 generation. We also find him on the maternal
side twice in the F3 generation and once in F4. For each combination
(from paternal to maternal) we must calculate the percentages and add
them together (in other words, coefficients are collective). F3 +
F3=3.13%
F3 + F3= 3.13%
F3 + F4=1.56%
So Diviseur
has contributed 7.81% of the remaining 50% to the increased
homozygosity. But we cannot stop there. We also find Irma in the F3
generation of both the sire and dam. So to our total we must add
another 3.13% to make 10.94% of the remaining 50%.But wait, we need to
look a little farther. We see that Etincelle is the mother of Irma. We
have already calculated Irma so there is no need to calculated
Etincelle, EXCEPT that Etincelle ALSO appears in a different context
as the mother of Tyrolienne. This makes her a separate factor and we
must now calculate for her as well. On the Paternal side, she appears
in the F3 generation as the mother of Tyrolienne. On the maternal side
she appears in the F4 generation as the mother of Irma.
F3 + F4 =
1.56%
Our final
calculation looks like this: Diviseur 7.81% + Irma 3.13% + Etincelle
1.56%, for a grand total of 12.50% of the remaining 50% or 6.25% of
the total. 50% + 6.25% =56.25% homozygous.Get it? Work through several
on your own and it will begin to be clear. The secret is to keep a
sharp eye for ALL the combinations. Remember you are only looking for
matches between the two sides of the pedigree; matches on the same
side are irrelevant. Work backwards from the individual whose pedigree
it is. Once you find a common ancestor, there is no need to also
calculate for all of his ancestors (which are also on the pedigree as
many times as he is) UNLESS they appear in a different relationship.
Finally we
come to Percentage of Blood. This calculation assumes that any
individual is the product of the sum of its genetic makeup.n this
method, all generations in the pedigree together are equal to 100%. So
the parents would each contribute 50% of the total genetic material,
the grandparents 25% and so on. In an inbred pedigree each incident of
a given individual must be added together to give the total of his
contribution to the whole. This method differs from Wright’s Formula
in that the coefficient of inbreeding expresses the percentage of
homozygosity (the percentage of both genes in a pair being alike)
while percentage of blood only calculates the probability of one of
the genes of any given pair is present in the individual, passed down
from the ancestor in question.
Using the
values in the following table will allow you to quickly calculate the
percentage of blood for any given ancestor. When the ancestor only
appears once then his contribution will be the basic value for the
generation in which he appears. However, if he appears more than once,
then you must add together all the values for each and every time he
appears in the pedigree.
|
Percentage
of Blood |
|
F1 |
F2 |
F3 |
F4 |
F5 |
F6 |
F7 |
|
50% |
25% |
12.5% |
6.25% |
3.13% |
1.56% |
.78% |
Let’s look
at our sample pedigree again. Beaux Arts and Hirondelle, as parents,
each contribute 50%. Each grandparent (Orgueilleux, Tyrolienne,
Arcol du Bourg and Utopie du Bourg) contributes 25%.Now things start
to need a bit of addition.
Diviseur
appears 3 times in the F3 generation and once in F4 which makes his
total contribution 12.5% x 3 + 6.25% = 43.75% (in other word, for
any gene pair, there is a 43.75% chance that one of the pair
descended directly from Diviseur). Irma appears twice in the F3
generation, making her total contribution 25%.Etincelle appears once
in F3 and twice in F4 for a total of 25%. Quicko appears 3 times in
the F4 generation for a contribution of 18.75%. You should be
getting the picture now. Finish the calculations for each individual
on your own to become an expert.
Now that
you’ve looked at all that, you’re back to square one. Now you must
personally decide how much homozygosity you want to deal with in
your animals.Remember, homozygosity is not necessarily bad. That’s
how purebreds get to be uniform in the first place. If an individual
has perfect genes, it can, in theory, be 100% homozygous without
consequence...however, no individual is perfect. Somewhere lurking
in there are genes none of us want to know about. But to decide
whether inbreeding is beneficial or detrimental to your goals, look
at your stock, or the stock your stock came from.Is it well
conformed? Does it have any growth problems? Is its over-all health
good? Does it seem to have a degree of intelligence? Is it fertile?
Have a good life span? Is the size staying consistent or is it
getting smaller with each generation? Are you producing a few
outstanding individuals, but also a lot of culls? Do buyers shy away
from your stock because it’s inbred?
By now you
may have an opinion on inbreeding. Perhaps after looking at all the
individuals that make up one animal you’ve either decide inbreeding
is too risky or produces stock you don’t want and can’t sell.
Perhaps you’ll find it works well for one or two generations and
then needs to be out crossed. Maybe you’re of the opinion that a
mild form of inbreeding (line breeding) is the way to go. It could
be that you’re dead set against it, or 100% for it. The point is
every breeding operation is as individual as it’s animals and what
works with the stock at one farm might not work at another. But I
hope I’ve given you a little food for thought and that it will help
you find what’s best for you and your animals.
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