The following is FORWARDED (from the Canine-Genetics@yahoogroups.com list)
IN ENTIRETY WITH PERMISSION from the original author.

Belated Tullibardine Holiday Greetings to All Chinookdom from snowy
Rossburn, Manitoba, CANADA! Where (34) of (36) Tullibardine owned beloved
Chinooks were legally and successfully imported finally this past Nov. 1st,
2007! (The two elder females, Perry Greene Natanis & Bates Argenta, remain
in Maine with Mum! :-)

Best Regards,
Susan E (Murray) Bragg (aka Mrs. J Jeffrey Bragg)
Chinook Trading Post
Tullibardine Farm Chinooks

Custodian of Records
International Seppala Association
http://www.seppalas.org

"Blanket permission to cross-post and/or forward my 28
Dec. 2007 CanGen submission "21st Century Dog
Breeding" is hereby granted to list members provided
credit to the author is given and the post is
reproduced in its entirety.

J. Jeffrey Bragg
The Seppala Siberian Sleddog Project
http://www.seppalasleddogs.com
http://www.seppalakennels.com

--- In Canine-Genetics@yahoogroups.com, Jeffrey Bragg
<ditkoofseppala@...> wrote:

Two and a half years ago I asked on this list whether
we could not somehow manage to come up with what I
then called a "synthesis of praxis" drawn from the
theoretical principles of population genetics -- a set
of guidelines for dog breeders that would show the way
to a more innocuous mode of dog-breeding than that
which is now practised by the vast majority, a set of
rules, guidelines or principles that would allow us to
breed in accordance with the principle "primum non
nocere" -- "first, do no harm!" Although a few people
acknowledged the desirability of such a document,
nobody came up with a draught text and we never really
managed to get a thorough discussion going of which
principles ought to be included. That thread, headed
"Dynamic balance + heterozygote superiority vs.
screening and selection"
(http://tech.groups.yahoo.com/group/Canine-Genetics/message/17526)
terminated with a brilliant and thought-provoking post
from Jim Seltzer (one of the brilliant minds on this
list whom I hold in great awe and respect), saying
that I ought to consider the concept of the "adaptive
topography" sometimes used by population geneticists
to provide a three-dimensional model of the dynamic
interrelationship of fitness, gene frequencies and
evolutionary adaptation. I thought about that for
quite awhile but never could discover how it could be
made into a tool for practical dog-breeding. In the
end, though, I went ahead and draughted my own list of
principles for 21st-century dog breeding, which I
never published because I never was able to put it
into a final form that I thought adequate.

Recent discussions on this list have given me an eerie
feeling of "this is where I came in," notably those
about "removing genes," the impossibility of using
screening and selection to solve the "genetic defects"
dilemma, the disconnect between geneticists and
breeders -- and the "new direction" thread generated
by the publication of Koharik Arman's paper. I found
Koharik's discussion very familiar-sounding. In
terminology and concept it strongly recalled to mind
the dialogue that I attempted without success to
initiate within the Canadian Kennel Club in the
mid-1990s with the self-publication of a brief
entitled "Purebred Dog Breeds into the Twenty-First
Century -- Achieving Genetic Health for Our Dogs,"
(http://documents.seppalasleddogs.com/html-documents/pbdb21c.htm),
cited as reference number 3 in Koharik's paper. At
this point in this list's discussion it might be
useful for those following it to review the latter
brief published almost twelve years ago, and then to
read Koharik's paper immediately afterward. Perhaps
the reader may then feel, as I cannot help feeling,
that although we may have raised a certain level of
awareness relative to the purposes of this list,
perhaps we have not really gone anywhere since the
days of Dr. Armstrong's efforts to raise these matters
on the web.

Meanwhile the urgent demands of the Seppala Siberian
Sleddog Project have required that I "wing it" as best
I could, creating for purposes of the Project a
coherent body of breeding practices. I have tried to
keep a couple of other developing breeds (such as the
Shiloh Shepherd Dog and the Chinook) under
observation, feeling that they shared similar
challenges to those faced by the SSSD Project; I have
not had the time, though, to read every post to this
list or seriously to investigate what those in other
breeds may have been doing. So, at best, I have had to
work mostly from my own personal knowledge of the
principles of population genetics, within the
parameters of our own evolving SSSD breed, with
relatively little light shed on our problems by the
practices of other breeders.

Not every breed may be in a position for its breeders
to do some of the things we have done in the SSSD
Project -- breeders of Chinooks, for example, cannot
avail themselves of landrace stock from the "country
of origin" of their breed, both because the breed is
synthetic in origin and because its original component
canine strains are not completely known. With the
strong caution, then, that not everything recommended
here may be possible or appropriate for all other
breeds, for every situation, or for any particular
breed other than the Seppala sleddog, I offer for the
list's consideration the following guidelines drawn
from my own imperfect knowledge and limited
experience. Please realise that I am not saying that
you (as an individual dog breeder) must necessarily do
any or all of these things. Still less would I wish to
see any such guidelines imposed by government as laws
or regulations upon the dog breeding community. But I
suggest that if you are concerned about inbreeding,
genetic diversity, and inherited illnesses, you might
wish to consider implementing some of the following
principles whose observance we have found useful in
the Seppala Siberian Sleddog Project.


BALANCE OF SIRES AND DAMS - Make a great effort to
maintain a reasonably equal numerical balance of sires
and dams; don't consistently use fewer individual
sires than dams. The so-called "popular sires"
syndrome has received much discussion and attention.
What may not be as well realised is that this selfsame
syndrome is repeated in miniature in most kennels,
where one or two of the "best" males cover all the
bitches, sire all the litters. (How often has one
heard it said that "the best males should sire all the
litters!") In order to avoid needless reduction of the
effective breeding population, just as many individual
males as bitches should contribute to the population;
this holds true whether we speak of the breed
population as a whole, or of the population within a
single kennel.

INCESTUOUS MATINGS - Do no incest breeding whatsoever
(even if you would rather call it "linebreeding" or
inbreeding). Matings of related individuals closer
than cousins ought not to be contemplated unless it
should become absolutely necessary to prevent loss of
a bloodline. This means: (a) no brother/sister
matings, (b) no father/daughter or mother/son matings,
(c) no half-brother/half-sister matings (i.e., sire
and dam share one parent in common), (d) no
grandsire/granddaughter or grandson/granddam matings.
This does not mean that first-cousin matings (sire and
dam have different parents but the same grandparents)
are okay or recommended; it is simply a case of having
to draw a line somewhere, at a given degree of
consanguinity, in order to say "anything closer than
this is quite beyond the pale and cannot even be
considered." (Otherwise an excuse will be found even
for full-sib matings.) If the available population
diversity within your own breed allows you to draw the
line further out, so much the better.

COEFFICIENT OF INBREEDING - To avoid frank incest
matings within the first three generations of pedigree
is not enough in and of itself. The Coefficient Of
Inbreeding (COI) must also be monitored, preferably
over at least eight to ten generations of the known
pedigree, keeping it as low as possible. To do this
over more than two or three generations requires the
use of computer software such as CompuPed, Breeder's
Assistant, FSpeed, etc. (To calculate a four to six
generation COI only gives a false sense of security;
usually such a COI fails to tell the whole story, and
the ten-generation COI will be found to be
dramatically higher.) In a purebred dog breed COI can
hardly be too low; almost always it is far too high!

It is impossible to recommend an arbitrary figure for
percentage COI, because the situation of each breed is
likely to be different. Probably anything greater than
5% constitutes a distinct threat to genetic health,
yet setting the bar at 5% may be virtually impossible
in many breeds. There are breeds in which breeders may
have to make great efforts to get it as low as 20%; in
at least a few breeds 20% would be alarmingly and
needlessly high. But breeders should KNOW what the
average 10-generation COI level is for their breed, at
least, and seek to keep their own breeding well
beneath that average level! Otherwise the COI just
goes on indefinitely, increasing steadily year by
year. It is easy to point to specific individuals in
numerous breeds with COIs of 70% or more, but it would
be a real challenge to produces examples of less than
5% COI in many breeds.

One ought to take care that the COI trend in one's own
breeding is never upward, but always either downward
or at worst neutral. This is done by averaging the
individual COIs of sire and dam (add the sire's COI
and the dam's COI and divide by two) and then
comparing this average with the COI for the trial
mating or litter that would result from mating those
two individuals. If the litter COI is higher than the
average of the parents, then you are obviously
increasing the overall level of inbreeding by
performing that mating, and the greater the disparity
between the two figures, the more the mating should be
deprecated. You may also wish to look at the same data
from a different perspective by calculating (with the
same pedigree software) the Coefficient of
Relationship (RC) when examining trial matings, the
more easily to ascertain which of two or more
alternative matings has the least-related parents.

NUMBER OF UNIQUE ANCESTORS - You must use a breed
database in conjunction with a pedigree and COI
application such as Breeder's Assistant or FSpeed to
explore COI and trial matings. When you do so, you
should also use it to study the number of unique
ancestors in the known pedigree, the number of
ancestors in common between sire and dam, and the
number of ancestors unique to each parent. These
figures are useful in assessing the potential
diversity of a projected mating and will tell you more
than the simple COI (which, after all, is only a
percentile probability figure predicting the
likelihood that alleles at the same gene locus
contributed by the sire and dam will be identical by
descent).

PEDIGREE ANALYSIS - Also try to carry out in-depth
pedigree analysis for every mating, listing the major
ancestors on which inbreeding occurs in that mating,
noting the number of occurrences and the generation
number of each occurrence. This analysis should be
carried back for at least six ancestral generations,
ideally for eight. This practice will alert the
breeder to undesirable "pile-ups" on key animals and
therefore to potential genetic problems (if such are
known to be associated with such individuals) in the
planned mating. An alternative or supplementary
approach is to use the "percentage of blood" function
of pedigree software such as Breeder's Assistant.

ASSORTATIVE MATING - Instead of inbreeding, use
assortative mating (mating unrelated parents who are
phenotypically similar for the desired traits) to
emphasise or fix greatly desired traits. Assortative
mating is much less dangerous than inbreeding and will
accomplish much the same ends. It should be obvious
that to breed "like to like" for given desired traits
will tend to yield more of what is desired, but if the
parents are not closely related, there is a greatly
reduced chance that other unconsidered traits will be
unknowingly reinforced by such matings.

GENERATION TIME - Genetic losses occur almost
infallibly with each generation in purebred dogs,
whether those losses happen through random drift, from
too few progeny contributing to the next generation,
from the inbreeding/selection cycle, or whatever. For
that reason, the fewer the intervening generations
that occur between foundation stock and current stock,
the less genetic diversity is lost. Breeders should
therefore maintain a high average generation time (age
of the sire at mating plus the age of the dam at
mating, divided by two) for each litter produced: four
years should be considered an appropriate minimum
floor level, five or six is better. It is helpful to
calculate a running average generation time for your
kennel throughout its history, by keeping a grand
average of the average generation times of all litters
produced.

REPEAT BREEDINGS - Do not always use the same sire
for a particular bitch (or vice-versa). Many kennels
make a routine practice of repeating favourite
breedings over and over again. Take care to maintain
diversity in your matings. Endless repetitions of the
same matings greatly reduce the available breeding
combinations both within the individual kennel and for
the breed at large.

SIBLING CONTRIBUTION - Try to ensure that at least two
of every litter (unless it should happen to be one of
those litters that had best be forgotten) contribute
to the next generation; half the litter should be the
ideal, though perhaps a difficult one to maintain. In
every instance in which only one progeny from a given
mating contributes to the next generation,
automatically and infallibly 50% of the available
genetic diversity in that line is permanently lost! If
two progeny contribute the theoretical average loss is
reduced to 25%, still less if more littermates
contribute. This single point is a major source of
losses of genetic diversity, yet it often goes totally
unconsidered by the breeder.

FITNESS INDICATORS - Monitor key indicators of
survival fitness in your canine stock. These are
fertility (percentage of successful matings),
fecundity (average litter size compared to the norm
for your breed), birth weights, nestling viability,
survival to adulthood, and longevity; be sure that
your breeding programme does not trend toward the
reduction of any of these.

FOUNDER BALANCING - It may be valuable to try to
balance the relative contributions of founders (where
possible and appropriate), particularly subsequent to
founder events or genetic bottlenecks. "Founder" is a
relative term. If a breed has a long pedigree history
with original breed foundation stock at thirty or more
generations remove from current stock, it may well be
impossible to balance the contributions of the
original breed founders; their relative contributions
may already be set in stone for all practical
purposes. But founder events tend to occur repeatedly
within the history of a breed. Bottlenecks occur with
dismal regularity. At least the breeder can pay
attention to the most recent founder set that is
clearly identifiable, attempt to prevent the loss of
individual founder lineages that are seriously
under-represented, and seek to balance the relative
contributions. Clearly this is no simple matter and to
suggest that it be applied consistently may be a
counsel of perfection. At least it is one more
possible tool in the breeder's armoury against
diversity losses.

OUTCROSS MATINGS - The great majority of dog breeds
have been bred within a completely closed studbook for
sixty to a hundred years, with little or no fresh
genetic input throughout the entire period from breed
foundation to the present. Similarly, many individual
bloodlines have been treated in exactly the same way,
bred in relative genetic isolation from other
bloodlines. Each breeder ought perhaps to consider the
desirability of locating and using a true outcross
within his or her own breed (unrelated to one's own
stock for at least ten to fifteen generations) at
least once and to integrate the resulting progeny into
one's bloodline.

If there is any possibility to import unrelated stock
from a breed's country of origin, one ought seriously
to consider doing just that. This is mainly possible
in the case of landrace breeds, in which an
autochthonous regional population remains in the
country of origin, independent of exported stock that
may have become a registered breed in other countries.
Examples of such situations would be the population of
desert-bred coursing sighthounds in the Near East,
relative to the Saluki breed in Europe and North
America, or the relict populations of autochthonous
arctic spitz-type sled dogs relative to the modern
Siberian Husky, Alaskan Malamute, Samoyed, et al.

It would be difficult to overestimate the genetic
value of a single import animal, unrelated to the
"registered" breed population for scores of
generations but stemming from exactly the same
fountainhead. This I would term the "Holy Grail" of
the diversity breeder -- the ideal controlled-outcross
situation in which an immediate significant increase
in healthy genetic diversity may be obtained at little
to no cost in terms of breed type and purpose. (That
the Canadian Kennel Club rejected this option for the
Siberian Husky in 1994 demonstrates, I believe, the
true extent to which the umbrella all-breed registries
represent an obstacle to genetic health and true breed
improvement.)

In cases of small, highly-inbred populations for which
there is no landrace resource, it may become necessary
to consider an outcross or outcrosses to similar
breeds. If so, this ought to be faced squarely and
proactively by the breed club concerned and breeding
subsequent to the breed outcross ought to be a
collective endeavour, shared for purposes of more
thorough integration and to reduce the work-load on
any one breeder -- because, no question about it, the
integration of a breed outcross is a major task that
can hardly be undertaken alone by the average breeder.
(The Backcross Project in the Dalmatian breed was an
excellent example of a breed outcross well-purposed
and superbly integrated; but the reaction of the breed
club was deplorable.)

POPULATION GROWTH - In the case of small, developing
breed populations, it should be regarded as important
to monitor and control the growth of the population
(in number) such that there is steady expansion of the
population within the limits of breeders' kennel
capacity and the demand for progeny. Growth by fits
and starts, with overexpansion followed by sudden
cutbacks or population collapse, is very bad for
genetic health. It is difficult, at best, wholly to
avoid population bottlenecking. But its existence and
ever-present possibility should be recognised and to
whatever extent may be possible, breed clubs and the
individual breeders should do whatever they can to
ensure smooth, steady population expansion and to
minimise cutbacks and consequent genetic
bottlenecking.

BALANCED TRAITS - One ought always to evaluate
breeding stock for balanced characteristics: health,
vitality, temperament, working ability, intelligence,
structure, type. Breeders should aim to maintain the
balanced characteristics of a total dog, not just to
produce winners at dog shows, field trials, races,
etc. An all-round, balanced dog will be a much better
hope for the future than a highly-selected, over-bred
dog that is thought to be "best" due to possessing
exaggerated traits in one or two areas, whether it be
a "perfect head," a showy gait, a faster racing speed,
or whatever. First, each individual needs to be a good
dog, and that should come ahead of breed
considerations.

UNFIT BREEDING STOCK - It ought not even to need
saying -- but in these days in which extensive, heroic
and expensive veterinary measures are routinely used
to save otherwise doomed animals, it does need saying:
the breeder ought never to breed from dogs that would
not be alive but for such interventions (excepting, of
course, survivors of physical injuries). It should be
obvious that if we prevent the operation of natural
selection, many of the animals that we use for
breeding purposes are likely to pass on various
transmissible genetic weaknesses.

REPRODUCTIVE TECHNOLOGY - Breeders should also
consider whether it is in their breed's interest
routinely to use elaborate reproductive technology to
produce litters. These days various and sundry
technical means are available which circumvent natural
mating and whelping. Some breeds, indeed, cannot
either mate or whelp a litter without veterinary
intervention -- already! If we use artificial
insemination and hormone assay to effect mating
combinations that cannot be brought about by natural
mating, along with routine C-section to deliver
litters, we may rapidly find ourselves in the position
of having created strains which cannot reproduce
naturally without technological support. We should
also consider whether it is really a good thing to
freeze the semen of outstanding males and thus extend
their breeding life decades into the future; this
practice seems to be universally approved, while no
one appears to have examined what effect such
extension of the influence of individual stud dogs
might have on breed genomes.

ARTIFICIAL SELECTION - This may really be the most
important principle of all, and the most difficult for
the vast majority to accept. Breeders should avoid all
extremes of artificial selection! When one comes to
consider the problem of lost genetic diversity,
inbreeding by itself is only half the story. The hard
truth is that selection itself is just as great a
culprit, if not worse. Inbreeding and selection
combine together in a cyclical fashion in the dog
world to cause the systematic depletion
("depauperisation" to the geneticist) of purebred
genomes. The desire for a cookie-cutter "consistency
of type" causes healthy genetic diversity to be
discarded intentionally at an alarming rate. (An
example of this desire is the person who declared at a
Chinook specialty show that he saw at least five
different types represented there, and that "they had
better get themselves a geneticist or they will never
have a standard type." The Chinook is a working breed
with a dangerously low population and a perilously
narrow genetic base; the kind of diversity that
engendered that comment is hardly to be deprecated in
such circumstances.)

We hear endless discussion about inbreeding and its
evils, and rightly so; but we hear very little about
the dangers of sustained extremes of artificial
selection, which are if anything yet more dangerous
than inbreeding. Together these two factors become an
engine of destruction for genetic diversity. The
constant obsession with having the "best" dog and with
"breeding only the best to the best," whether in
dog-show terms, in dogsled racing, or whatever,
creates a situation in which the best is definitely
the enemy of the good. The endless repetition of the
inbreeding/selection cycle in the quest for a dog that
is better than last year's best, has systematically
stripped away most of the healthy genetic diversity
from today's purebred dogs. Stringent, sustained
selection for cosmetic ideals (shape, number and
intensity of the Dalmatian's spots; shape and
chiselling of the poodle's muzzle; subtleties of
colour and markings in an endless series of breeds) or
narrow ideals of performance or athleticism (top
sprinting speed in racing greyhounds or racing
sleddogs) have for many decades taken absolute
precedence over breeding to provide the kind of
"genetic outfit" that will allow the dog to be healthy
and hardy.

Now that canine diversity has been stripped to the
point that homozygous recessive "defect" genes are
everywhere apparent, the dog fancy proposes to remedy
the situation by embarking upon a new level of
elevated selection, armed with DNA marker testing to
enable the wholesale "elimination" of "defective"
genes. This new wave of super-selection on top of the
already extant depauperisation may well become the
killer wave that will sink the entire ship of purebred
dogdom, AKC, CKC, and The Kennel Club with it. DNA
testing has become a growth industry. This all may be
more about corporate profits and grant money, than
about genetic health. It is up to breeders to have the
common sense to realise that what is being proposed is
a losing game, that already depauperate purebred breed
genomes will not support further massive artificial
selection and the consequent wholesale elimination of
yet more genetic diversity. The "defect" genes cannot
be excised with a scalpel; many other genes that
happen to reside on the same chromosomes will go right
along with the defects, with totally unforeseeable
consequences.

In conclusion, let me say that, although this set of
guidelines cannot be made into hard and fast rules or
(worse yet) regulations -- because the situations of
each individual dog breed and even each breeder are
different -- yet I believe we all need faithfully to
attempt to apply the principles discussed above, in
order that our dogs may have long, healthy lives upon
the earth. We should strive to be faithful stewards of
the genetic heritage of our canine friends. In that
way we may hope that our bloodlines will endure longer
in the dog world, and in the end we may even be
remembered as pioneer 21st Century dog breeders who
strove heroically to correct the errors of the past in
the light of better knowledge of population genetics.

J. Jeffrey Bragg
Seppala Kennels http://www.seppalakennels.com
The Seppala Siberian Sleddog Project
http://www.seppalasleddogs.com

--- End forwarded message ---