OF PEAS AND PUPS
PART IX
INTRODUCTION:
Selection, we concluded
in Part VIII, is the oldest and most used of the breeding tools. If any leg of
a three-legged stool is most important...selection is the most important of the
breeding tools too. It is also the "handle" for the others tools
since it is necessary for their effective use. Selection is used to choose one
individual at a time...a sire or a dam...The "selection-handled"
tools are used to choose pairs, mates, a sire and a dam, and therefore might
better be called plans than tools.
...The genes already
present within the breed are the materials of the genetic builder. The tools or
plans for the manipulation of these genes are the breeding systems...they
permit the shuffling, re-arranging and aligning of the contents of the germ plasm...As with all forms of building, there are specific
tools for each job. No breeder will succeed with inbreeding where, he should
have outcrossed and the results will be no less
absurd than driving a nail with a saw. He who mates like to like, using no other
tool, is no less foolish than the builder who cuts his boards with a hammer.
Carrying the analogy a step further...a builder who starts with poor material,
will build a poor house, regardless of his ability or the completeness of his
tools. So to build a better Shorthair, we must have good genetic material to
start with and we must use each genetic tool for its proper job.
A BREEDING PLAN
WITHOUT BASIS:
...Before discussing
worthwhile mating plans, we should, solely for the sake of completeness include
that form of mating which accounts for the greatest number of canine offspring.
It is, of course no real "plan" at all. It is random matings...the
uniting of sire and dam without regard to Appearance, Performance, Ancestors, Descendance...convenience being
the "tool". Like drawing their names out of a hat.
Occasionally, someone will point to a good pup from such a chance mating and
use such evidence as proof-positive that all breeding is a matter of dumb
luck...The fact of the matter is that a chance mating which does produce an
exceptional specimen is Dumb Luck, and for each such, there are hundreds, which
tend to act as a drag on the breed...
...We may assume, that the good "meat-dog" which oftentimes
results from random-mated Shorthairs, is largely due to the degree of homozygosity, built in by intense
inbreeding in the foundation stock...after all, the breed is less than 100
years old. This original inbreeding , coupled with the
fact that the total population of the breed has remained small, force toward homozygosity even in random-mated Shorthairs, which is not
present in older and more populous breeds...Or, it is also possible that the
random mating just happened to nick, or it might have been that chance provided
a genetic inbreeding even though no pedigree inbreeding was apparent. Whether heterosis
or inbreeding produced the favorable results will
depend upon what comes from future matings of this
exceptional chance-mated sire.
BREEDING PLANS
BASED ON THE APPEARANCE and/or PERFORMANCE OF THE MATES
Mating Likes to
Likes:
...Its name pretty well
describes the plan. In the strict sense of the term, it means mating the worst
to the worst, as well as mating the best to the best. It is the latter, which
is usually implied for no one mates the worst to the worst, except through
Ignorance, Inexperience or Change...To make full use of the tool, we must use
its selection handle. Art is basic to wise selection. For the Shorthair,
selection may be based on the conformation of the dog or on his hunting ability...ideally,
it is based on both.
...If we mated animals
with like pedigrees, we would probably be inbreeding but, here, we are
concerned with likeness of Appearance and Performance of mates themselves...as
a breeding system. What happens when like animals are mated...both being above
average? We assume the use of but two genetic tools...phenotype selection and mating likes to
likes. There will be considerable similarity between parents and
pups because each pup is made up of two halves of a similar appearing pie. The
crust (phenotype-appearance),
makes the pies, look alike. But how is the breeder to know that the
"TM" on one pie stands for "tis
mince" while the "TM" on the other stands for "taint
mince"?...The two parent-pies, are carefully cut down the middle and just
as carefully, a half from each is placed in a clean new pie plate to make
another pie. This puppy pie LOOKS like the parent pie but its half apple and
half mince...Certainly the results from a like to like mating might no be so
genetically divergent as the mince-apple pie...but, then, even mince pies have
some apples in them...The point is, that although sire and dam have some genes
which produce similar appearance, these genes do not necessarily come from the
same allelic series.
...If, instead of two
tools, we used three our gains could be held. If, with selection and mating
like to like, we used some form of inbreeding, the chances would be much
greater that the similar appearance and performances of the parents would be
due to genes in the same allelic series. This is because, as we have said,
related animals are similar
because they have received common
genes from common
ancestors...If we start with apple pie parents,
we end up with apple pie pups. Homozygosity, contributes to breed improvement mating like to
like, alone, increases homozygosis but little and therefore any gains are shortlived...Like to like matings
reach their limit in a generation or two and if inbreeding does not immediately follow to FIX any
gains made, a rapid return to the average can be expected.
...If the goals of all
Shorthair breeders were the same, within reasonable limits, like to like matings would do the breed no real harm. However, where
there may be a difference, as, for example, in the size of the Shorthair, this
plan can produce profound divergence...It can act as a snow-plow,
disrupting uniformity and dividing the breed right down the middle. The field trialer who wants a smaller dog
and the show people who must have larger dogs to please the judges. I do
not maintain that this divergence is developing to any extreme degree (altho' that it is present cannot be denied) nor that mating
like to like is causing it. My point is that like to like matings
produce wider extremes IF goals differ.
MATING UNLIKES:
...This term, too, is
self-explanatory, or, if we wish to get technical, we can call it
"negative assortive mating on the basis of
somatic resemblance." The term I like best is selective compensatory mating. Its most useful
function is the correction of defects,
but here again, to be most effective, a third tool should be used. Lloyd
Brackett in his "Planned Breeding" (Dog World-Chicago '61) stresses
again and again..."Physical
compensation is the foundation rock upon which all enduring worth is built."
His compensatory matings were accomplished by close
mating within his own strain. His success with the "three tool plan"
is testified by some 90 Show Champion German Shepherds...a record approached by
no other breeder of any breed!
...The correction of
physical faults and the production of intermediate types makes
this tool one whose primary use applies to conformational improvement. It has
little use with regard to field traits except indirectly, that is, except that
we know a dog "Must be built right to work right." We know of no
field dogs which have too much nose, too much run, are too staunch on point, or
are too intense in backing.
...Mating unlikes is a matter of degree. Like mating likes, no two
dogs are identical...Even when we try to mate likes, we never succeed fully.
GERD has an exceptional nose. Our desire is to pass it along and strengthen it.
To this end, we should seek the finest nose in a female of his strain...like to
like in the like, in the same
strain. Since we cannot find any female with a nose to match his,
we will be mating unlikes, whether we like it or not.
GERDA has a fine nose but GERDS is better. The pups will be intermediate. How can we
channel Gerd's great nose into some of his get?..We can continue to breed
back to Gerd, each time picking up a few more of
those great nose genes...but we are getting off the subject.
...Mating unlikes, intentionally, is primarily a conformational tool,
as we have said. If, for a simple example, we have a small male, say 22
1/2" at the shoulder, he should be mated, to correct this fault, to a
larger female. For best results, she should be about as far over he is under,
say 23 1/2". The results should be intermediate...particularly if the
mates are of the same family or "bloodlines". This is not to say, of
course, that when 23 1/2" bitches are mated to 22 1/2" sires, all the
male pups will be 24" and all the female pups will be 22" at
maturity. If breeding were an exact science, it would lose much of its interest
and enjoyment...it is still primarily art and no such slide-rule results can be
predicted, but the guiding principle is still there and working.
...If we neglect
selection and use to plan but the mating of unlikes
what would be the overall results to the GSP as a breed?
-1. Heterozygosity would be increased but little and
therefore but little homozygosity would be lost.
-2. The extremes of type would be lost...the
breed would become more uniform. This would take place rapidly, probably little
over a generation. The return to its present state would be just as fast with a
generation of random mating.
-3. In contrast to mating likes, the pups
would little resemble their parents or grandparents but would have a tendency
to resemble each other a great deal more that in a chance mating.
...Do these
fundamentals of mating unlikes have any practical
application? I'll let you answer that.
SUMMARY:
...Mating unlikes has a its most important
use, the correction of faults...primarily show faults, and the production of
intermediate types. The faults are more certain of correction if a third tool
is added. The fixing of some intermediate types is possible through inbreeding
once they have been established by mating unlikes.
The fixing of such intermediate traits as would result, from genotype of Aa, Bb, Cc, etc. is impossible
with any tool or tools now available.
BREEDING PLANS
based on the Relationship of the Mates:
...The Wall Street
Journal may seem like a strange source of genetic information, however in a
recent issue it reported a genetic breakthrough which is pertinent to our
discussion. The story concerned genetic relationship...By a new technique,
scientists at the Carnegie Institute have been able to separate the "beads
for genes"...to divide our double strand necklace into two single stands.
This is something nature does many times each moment, but this is the first
time man has accomplished the feat. A single strand from one species is exposed
to a single strand from another species. When the two strands
have the same beads...identical genes (allelic pairs or series), they will
combine and cling together in the area of their likeness. By this
process, it was determined that the salmon and man are related by 5%. They
share 5 genes in every hundred. No mention was made of the relationship between
man and dog. Are you man or mouse? Mice and men share 20-25% of their genes in
common. The method is not without error and they have as yet been unable to
count the number of genes in any species but can this knowledge be far off?
...On this basis, all
dogs of all breeds are 99 plus % related...and they are...that is, the single
strand (gene string) from a male of any breed will combine and cling to a
single strand from a female of any breed...they can mate. But what concerns us
here is the relationship between the members of the same breed. This
relationship is based not alone on the number of the allelic pairs in common
but more especially the number of Homozygotic allelic pairs in common,
either dominant or recessive. Two animals which not only possess the same genes
but the same genes in the same form, are identical
twins (even if they didn't have the same parents). Such individuals could not
mate of course, they being the same sex...if they could,
this would constitute self-fertilization...not possible in any animals,
although common in plants.
...GENETIC RELATIONSHIP
within a breed is based upon homozygotic genes in
common. Since we cannot know with certainty these genes, we must use terms
which have the most likelihood of indicating that genetic relationship. But we
must always be mindful that this relationship...and the plans based on it...are
but possibilities, not certainties. The probable relationship between
granddaughter and grandsire is 25%. But it may be 50% or it maybe 0% and we
must remember, these possibilities too. If the
relationship were 25%, mating the two would be inbreeding (Fx
.125)...any mating which causes the relationship between parent and offspring
to exceed 50%, the inbreeding would be more intense (Fx .25), the same as mating full
sibs. If the relationship were 0%, which is possible but improbable, no
inbreeding would have taken place...contrary to what the pedigree says. Thus,
factually, a pedigree inbreeding, could be a genetic outbreeding. We are not building paper houses or paper
dogs...the genes are our building blocks. These possibilities are being
constantly stressed only to emphasize the lack of consistency, the importance
of ART in breeding.
...Merely for
explanation, lets make the GSP a single gene-paired
animals, as we did in discussing relationship in Part VI. There would be but
three types...AA, Aa &aa. Under such circumstances, mating AA to aa would be a perfect outbreeding. Mating AA to AA or aa to aa
would be perfect inbreeding. Aa to Aa would be the breed average. A
breeding which produced AA or aa
pups would have been an inbreeding...that which produced Aa
pups, an outbreeding. This brings us to the strange
fact that a random mating...the average...Aa to Aa, would produce two inbred
pups, AA, aa, and two outbred
pups, Aa, Aa, in the same
litter. Inbreeding results when like genes meet in the zygote...outbreeding results when unlike genes meet in zygote. It is
only art, and lots of it, that can unravel such a tangle. It is obvious that no
one definition will satisfy all circumstances. If difficulties arise even in
such an oversimplified hypothetical case, how much more complex the real ting
can be. So in describing breeding plans, let us not be too positive or too
emphatic. In most cases, the pedigree and the genes will agree within
reasonable limits...it is the exceptions which will arise to plague us and of
which we must always be aware.
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...FIGURE
38, is a summary of breeding plans based upon genetic relationship. All
breeding terms are relative. We would be inbreeding by mating a Shorthair and
a Wirehair...compared to mating a dog and a wolf, it our relationship were
based on the whole genus canis. But for our purpose, and in most
instances, the term relates to a breed...here the FSP...We arbitrarily, set
out starting point at a breed
average of 10,000 homozygous and 10,000 heterozygous genes in the
members of the breed. The 20,000 genes of the Shorthair was merely a figure,
little more than pulled out of the air...In the accompanying illustration the
beads for genes represent the eight-gene string of a typical (or average) gamete shown will produce
complete heterozygosity
for these strings...as do the gametes shown under "perfect outcrossing". The difference between the two lies in
the fact that in the random mating, crossing over could produce some homozygosity, whereas in the perfect outcrossing
no amount of Crossing Over will produce one iota of homozygosity.
This is true for the same reason that no amount of crossing over in the
"perfect inbreeding" will bring the slightest heterozygosity.
In the former instance, for these eight genes, there is no genetic
relationship...the dam is homozygous
dominant, the sire, homozygous
recessive. This is also the reason that at breed average, we
indicate both Rx and Fx at zero. With the minor
exceptions previously mentioned, there is no breed relationship (above the
average) and therefore no inbreeding at that point. We might consider that
point as being Genetic Equilibrium for the breed. |
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...A diagram such as
this, attempts to tell a story briefly and graphically. It omits details, is
satisfied with averages and is therefore somewhat incomplete. What actually
concerns our topic ends with the perfect outbreeding...the
balance is included for prospective. There are those who abhor, even the
thought of mating Lab and Shorthair...yet, they never give the matter a second
thought when they come as close to that point as possible and still stay within
the same breed. Outbreeding, is the crossing of
strains and differs from crossbreeding only in degree....Under generic crosses, we list the
mating of sheep and goat. There seems to be enough relationship so that
fertilization takes place and the fetus starts to
develop but it rarely goes full term. It is either resorbed
or aborted. Cross breeding,
species crosses, etc. need not concern us here except to complete
the picture and to demonstrate that there is a point beyond which heterozygosity cannot go and life be
maintained.
OUTBREEDING:
...OUTBREEDING IS THE
MATING of animals less closely related than the average for the breed. We often
hear it described as the mating of unrelated animals...and relatively, that is
correct, but only animals which are closely related can mate, when we consider
all animals. That's why the average for the breed, is important in
defining the term. We might also say that outbreeding, is a mating
which brings fewer like genes to the zygote than a chance or average mating.
For convenience, and with a reasonable degree of certainty, we may say that a
pedigree which shows no common ancestors in the usual five generation pedigree, is probably an outbreeding.
Of course, the population an outbreeding.
Of course, the population and age of the breed, together with the amount of
inbreeding practiced through the years would also effect
the actuality of such a statement. We must remember too, that there are
instances probably not many, where the gifted breeder has sheparded
the genes in his kennel down through the generation by sheer artistic talent
alone...the pedigree does not show the art involved...Motherline
breeding, which is difficult to read in a pedigree, is such a plan.
...CAN OUTBREEDING
PRODUCE good pups?...Certainly! As a matter of fact, outbreeding will often produce better pups, phenotypically, than any other mating plan,
the first generation either is practiced. Can outbreeding
as a long range plan bring improvement? That is, generation
after generation of outbreeding? Certainly
NOT!
...Outbreeding
is to heterozygosity what inbreeding is to homozygosity. There is not outbreeding
without heterozygosity. In the following figure (39),
we see a theoretically perfect outbreeding. Note the
diversity of the parents. They are genetic opposites, unrelated in a breed
sense, yet they are both inbred and both are homozygotic.
Consider this for a moment...both are inbred...both are homozygotic,
yet they are opposites.

...Both pups are, in
themselves as individuals, good...phenotypically
good. Not genotypically good. The only way they can
become genetically valuable, is to be bred back to their sire...inbred. If, for
example, the little hybrid bitches are bred back to their sire, outcrossing would better describe the mating. OUTCROSSING
is an outbreeding step in an overall inbreeding plan.
Following the outcross, the original inbreeding (or linebreeding)
is resumed. Rare is the dog of any breed, whose genetic quality permits many
generations of close inbreeding without the need for an occasional outcross. In
a genetically perfect dog, there would never be the need for outbreeding. Since there are no such dogs, continued close
breeding, through its process of increasing homozygosity,
is going to eventually double up some recessive defects. To prevent them from
becoming fixed, we outcross. The outcross should be just as mild as consistent
with desired results, to avoid upsetting the genetic applecart...to avoid
excessive heterozygosity.
...OUTBREEDING WORKS ON
THE PRINCIPLE that there are more good genes than bad in all dogs. That most
good genes are dominant, most poor genes recessive...Thus, at first, outbreeding may bring improvement because the favorable dominants mask the unfavorable
recessives...BUT, the recessives are still there...not lost, just
hiding...Continued use creates and maintains heterozygosity
which brings uniformity...a uniformity of mediocracy
to your strain. Factually, I should have said, "to your kennel", it
will bring nothing to your strain because you
can have no strain based on outbreeding. Outbreeding destroys strains by scattering our hard-earned homozygosity to the four winds.
...The outbred animal which proves superior to its parents has
nicked or heterosed. It exhibits hybrid vigor, as explained earlier, and like a falling star, there
is a sudden bright flash in the sky...then darkness. Lush, referred to earlier,
sums it up briefly and accurately..."Outbreeding usually leads to individual excellence but low
breeding-worth." That is, good phenotype...poor genotype. It
is the good genotype which brings breed progress.
...The very rare
pedigree outbreeding which not only brings individual
excellence but also great breeding worth, has, in fact, been no outbreeding at all, from the standpoint of the genes. This
mating has produced homozygosity...like genes have
met...this is the only possible form of prepotency.. Homozygosity is the antithesis
of outbreeding.
...Most dog breeding
books tell us the prime use of outbreeding is to
annex to our own strain some virtue which is either lacking or weak. it sounds easy, if you say it fast...IF we do gain from an
outcross (this is the only practical form of outbreeding)
and this is no certainty, we must immediately fix that gain by inbreeding or it
is lost. Before embarking on any planned outcross, we should exhaust every
possibility that this favorable trait is no where
present in any animal related to our own strain. The closer to home we can pick
up this desired virtue, the more certain, we are of fixing it and the less
danger we suffer from disrupting strain type.
...The dangers from the
improper use of outbreeding are no less great than
those resulting from the improper use of inbreeding or any other genetic
tool...and no less art is required to do one well than the other. Probably the
widest use of planned outbreeding is carried on with
livestock. The increased size and weight which comes with hybrid vigor pays off on the stockyard scales. But, we do not
measure the quality of the Shorthair by the pound..."He can't be all
bad...he hates kids," Outbreeding isn't all bad.
Although I've never heard it mentioned, there is one
use which does much for dogs of all breeds...in a negative sort of way...The
beginner shuns inbreeding like the plague because of what he has been told. The reasons may be
wrong but the idea is not. Pushing the novice toward outbreeding
is a service to any breed. How? Because the inexperienced, through ignorance,
convenience or without realizing it, so often mate faults (like to
like)...Mating likes with outbreeding will no more
fix faults than it fixes virtues. If we must have faults...and we must...the
more "unfixed", the better...Outbreeding is
not a "fixin'" tool but a "mixin'" tool. Inbreeding is the fixin'
tool but both tools, like matches, do not belong in the hands of
children...even those 30-40 years old.
SUMMARY:
...Outbreeding
as a long range plan has little to offer the Shorthair. Outcrossing,
which implies the use of additional tools, has two primary uses...First, to
modify any degeneracy which might develop through homozygosity.
Should necessity require it at all, it should be as mild as possible. Second, to gain or strengthen some characteristic lacking in our
own strain. This step should be taken only after we are sure that it is
available from no closer source. If we gain from the outcross, we must
immediately inbreed to hold it. Outcrossing does not
automatically bring the desired results...and the artistic requirements are no
less rigid for this than any other breeding tool.
Part 1 | Part 2 | Part 3 | Part 4 | Part 5 | Part 6 | Part 7 | Part 8 | Part 9 | Part 10 | Part 11 | Part 12 | Part 13 | Part 14 | Corrections to Part 12 & 13
Copyright 2001. Dr. James G. McCue, Jr. All rights
reserved. Postscript: And his legacy lives on in the German
Shorthaired Pointers of today. May they
always be healthy and bred with forethought and planning.