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Crossbreeding Draft Horses
by Karen Gruner
Interesting things happen when we crossbreed animals. We have all
seen litters of crossbred puppies with wide variations in appearance. We have
also seen litters of crossbred puppies that are remarkably uniform. The same
thing happens when crossbreeding horses, but since we get at best one offspring
per mare per year, the results are often harder to see than when looking at
puppies.
Over the last 70 years considerable change in type has occurred between
the Brabant and American Belgian. We can all see this change, and it is this
difference that has driven myself and others to choose the Brabant over the
American Belgian.
Phenotype vs. Genotype The difference we can see
between these two types of horse is called the phenotype. Underlying what we can
see is a genetic base called the genotype. The genotype determines the
phenotype. So all animals (ourselves included) have a genetic makeup of specific
chromosomes called the genotype and an external appearance called the phenotype.
When crossbreeding, considerable variation in both genotype and phenotype
is possible, even between offspring from the same set of parents. I will try to
explain what the possible genotype of crossbred animals could be, using Brabants
and American Belgians as my example. The genotype of a 75% Brabant horse could
vary between 50% and 100%, making the 75% Brabant designation only an average
measure of the genetic type of the crossbred animal. We see this all the time.
If you have seen a number of Brabant-American Belgian cross horses you know that
all 50% or 75% Brabants do not look alike. How can some 50% horses be
indistinguishable from 100% horses, while some 75% horses look distinctly
crossbred?
Chromosome Pairs Horses have 64 total chromosomes. The
chromosomes come in homologous pairs (one chromosome from the dam and one from
the sire with the same function). So horses have 32 pairs of chromosomes. The
chromosomes are generally numbered by pairs, so the chromosomes are called
chromosome 1 to chromosome 32, each number corresponding to a pair. For the sake
of simplicity, let's consider five chromosome pairs, and let's call them Bi for
Brabant and bi for American Belgian, where denotes a number 1 through 5. Let's
mate our hypothetical 5 chromosome horses, a Brabant stallion to an American
Belgian mare. The genotypes of these horses would be:
Brabant stallion B1B1 B2B2 B3B3 B4B4
B5B5 |
American Belgian mare b1b1 b2b2 b3b3 b4b4 b5b5 |
When the sex cells are formed (meiosis), the pairs split. Each egg the mare
produces gets one chromosome from each pair. Similarly, each sperm cell the
stallion produces gets one chromosome from each pair. The following egg and
sperm cells would be created:
sperm B1 B2 B3 B4 B5 |
egg b1 b2 b3 b4 b5 |
Remember, this discussion is over simplificatied, since the stallion really
has two slightly different B1 chromosomes, one from his dam (B1d) and one from
his sire (B1s), and on down the line for all the chromosomes. So when the sperm
cell is made, it could have B1d, B2s, B3d, B4d, and B5s, or any other
combination. The same goes for the mare. For the sake of this discussion, we
won't take this into account. Another interesting complicating phenomenon occurs
during meiosis called crossover. During crossover, sections of a specific
chromosome, called genes, can exchange between the Bis and Bid chromosomes, so
the resulting chromosome becomes a blend of sire and dam. Again for this
discussion we won't take this into account. When this sperm and egg unite to
form the 50% Brabant offspring, the following genotype results:
F1 B1b1 B2b2 B3b3 B4b4 B5b5
First Generation Every F1 (first generation) Brabant-American
Belgian cross has 50% Brabant chromosomes (Bi) and 50% American Belgian
chromosomes (bi). These horses tend to look like crossbreds, although the
Brabant chromosomes seem to be rather dominant to the American Belgian
chromosomes. As a result 50% F1 crosses have a nice Brabant type (phenotype).
What happens when we cross two F1 50% Brabant horses? Now things can get
very interesting! During meiosis (formation of egg and sperm cells) the
homologous pairs of chromosomes split. Each egg and sperm carries half the
genetic information of the parent. Each egg and sperm has one of each of the
pairs. A minimum of 2x2x2x2x2 = 32 possible different genotypes can occur for
the gametes (eggs or sperm) in our five chromosome pair example. Twelve of the
32 possible egg or sperm types are shown below.
| egg or sperm types |
| 1 |
2 |
3 |
4 |
5 |
6 |
7 |
8 |
9 |
10 |
11 |
12 |
| B1 |
b1 |
b1 |
b1 |
b1 |
b1 |
B1 |
B1 |
B1 |
B1 |
b1 |
b1 |
| B2 |
B2 |
b2 |
b2 |
b2 |
b2 |
b2 |
B2 |
B2 |
B2 |
b2 |
B2 |
| B3 |
B3 |
B3 |
b3 |
b3 |
b3 |
b3 |
b3 |
B3 |
B3 |
B3 |
B3 |
| B4 |
B4 |
B4 |
B4 |
b4 |
b4 |
b4 |
b4 |
b4 |
B4 |
b4 |
b4 |
| B5 |
B5 |
B5 |
B5 |
B5 |
b5 |
b5 |
b5 |
b5 |
b5 |
b5 |
B5 |
Most of the possible gametes are a combination of Brabant and American
Belgian in genotype. Of course pure Brabant gametes (type 1) may be formed, as
well as pure American Belgian gametes (type 6). Any combination in between can
also be formed. So we see right away that if an egg of type 1 is fertilized by a
sperm of type 1, our supposedly 50% second generation (F2) offspring is
genotypically a full Brabant. Conversely, two type 6 gametes could fuse to
create a 100% American Belgian offspring. But most of the possible gametes carry
both Brabant and American Belgian genes, so most of the offspring will be
cross-bred. In fact, on the average, we expect the offspring to have 50% Brabant
chromosomes and 50% American Belgian chromosomes. But it is clear that there can
be a wide variation in appearance between F2 50% cross-breds since there is a
wide variation in possible genotype. I fact, there are theoretically 32 x 32 =
1024 different possible genotypes just for our 5 chromosome example!
Now let's look at what can happen with a 50% Brabant crossed with a 100%
Brabant. Note that all 75% Brabant horses are necessarily F2 (or higher) crosses
on at least one side. For our example we'll let the mare be the full Brabant and
the stallion be the crossbred. The stallion can produce the sperm cells shown
above plus the other 20 possible combinations not shown. The mare can only
produce full Brabant eggs, so each of the offspring is guaranteed to be at least
50% Brabant. If the egg unites with a sperm cell of type 1, the offspring will
be genetically full Brabant. If, however, the egg unites with a sperm cell of
type 6, the offspring will be genetically 50% Brabant. When we look at 75%
Brabant crossbreds we therefore often see a wide variation in appearance
(phenotype), even among full sibling offspring. Again, since most of the sperm
cells a 50% Brabant stallion produces have both American Belgian and Brabant
chromosomes, we expect that on the average the offspring will be 75% Brabant.
So when we quote percentage Brabant for a horse other than an F1 cross, we
are reporting only the average genetic makeup. We do not really know if the
horse is 50%, 75%, or 100% Brabant.
Karen Gruner is editor of "Trace Chains," the newsletter of the
American Brabant Association, from which
this article has been reprinted with permission. |