PATTERNING IN DOGS, WITH PARTICULAR REFERENCE TO THE GSP
Although there is general agreement amongst geneticists
as to the modes of inheritance of most aspects of coat colour and pattern
in dogs, confusion can arise due to different lettering systems being employed
by geneticists to describe the same genes. The lettering system used by Dr
Clarence C. Little (1957) in his book, "The Inheritance of Coat Colour
in Dogs" is widely used, and will be followed in this chapter, apart
from the description of the "A" series alleles, which is that used
by Willis (1977). All dogs have the same number of chromosomes and the same
number of genes controlling the colour and pattern of their coats. The particular
alleles of those genes, present in a particular breed, however, are in many
cases quite limited, and the appearance of a colour or pattern not normally
found in a breed is usually indicative of cross-breeding. For example, a "GSP"
with orange or lemon patches is most likely to be a cross-bred animal. Why
this is so will be made clear in this chapter.
Some of the genes mentioned are of little significance to the GSP, so they are included without detailed discussion. Of greatest interest to most breeders are the "B" series alleles, which determine whether the animal will be black or liver and the "S" series which determine whether it will be solid coloured or non-solid. There are of course, variations in the amount and distribution of the non-coloured areas and also in the amount of "ticking" or "roaning" which is present in the white or non-coloured areas. These variations are dealt with separately, with illustrations. Because of the confusion which exists amongst breeders and fanciers alike, I have dealt most fully with the "B" and "S" loci. Existing GSP books have included confusing information such as "ticking is..a dominant gene", given in a context which could lead one to believe that the genes for ticked (non-solid) coat patterning were dominant to those for solid colour-patterning. That, of course, is the reverse of the true situation. In fact, the true statement "ticking is dominant", refers to the amount of colouring found in the non-solid areas of the coat, i.e. clear white (non-ticked) in contrast to ticking or roaning in the non-solid areas. Even more misleading are statements which describe dogs as being "liver and whites with solids behind them" as if the solids might reappear in subsequent matings, or "black puppies cropping up legitimately in liver to liver litters" or even "black to liver may produce tricolours". All such statements reveal a misunderstanding of the inheritance of coat colour and pattern in the GSP. To clarify this issue, I have included herein all the possible matings of blacks to livers and solids to non-solids, together with the only legitimately possible outcomes. To simplify matters, all non-solid coloured animals, regardless of the distribution or patterning of the coloured hairs found within the non-solid areas of their coats (ticked, roaned or clear white), are grouped together, with "s" to represent them. Because GSP fanciers tend to refer to most non-solid animals as "ticked", I have added "ticked", when referring to non-solid. The looseness of this terminology must be kept in mind when the genes controlling the amount and distribution of coloured hairs in white areas are discussed. Using this simplified system, I will then be dealing with only four possible phenotypes, solid liver, solid black, liver/non-solid ("ticked") and black/non-solid ("ticked"). The nine possible genotypes, are as follows: liver/non-solid ("ticked"), heterozygous-solid/liver, homozygous-solid/liver, heterozygous-black/non-solid ("ticked"), homozygous-black/nor-solid ("ticked"), heterozygous-solid/homozygous-black, heterozygous-solid/heterozygous-black, homozygous-solid/heterozygous-black, and homozygous-solid/homozygous-black. The distribution of the coloured patches, and the coloured hairs in the white areas of the coat are not of great significance to most breeders and I have no wish to fill this book with tables of the possible outcomes ofmatings between "Irish spotted" or "extreme-white piebald" animals, or ticked to roaned or clear whites. I have included however, illustrations of these various pattern types and their modes of inheritance. Readers may work out possible outcomes for themselves, using the given colour/pattern tables as a guide. There is some disagreement amongst authors on the subject of ticking vs roaning, so I have included a hypothetical explanation of the subject based on my own observations and breeding experience. In Dr Little's book, the section on GSPs attributes no genes for colour paling to the breed. Nor does he acknowledge the presence of black, no doubt due to the fact that it is outlawed in the breed in his country (the USA). My own experience suggests that a colour paling factor (or factors) does exist. Dr Little attributes the varying depth of colour in Chesapeake Bay Retrievers to the recessive allele "c^". Since this breed, like the GSP, is genetically liver in colour ("b") it seems possible that the same allele ("c''11") is responsible for the light liver colour sometimes seen in GSPs.
Burns (1952), using material gleaned from Danish studbooks, describes a litter of eleven GSP pups, from two "normal" coloured parents, in which nine were "brown" (normal) coloured and two were "cinnamon-coloured". Burns ascribes this phenomenon to a modifying gene "Z", named by Steiger (1936). Willis (1977) mentions (without naming them) five alleles for colour paling in the German Shepherd so it well may be that another gene (or genes) other than the "C" locus gene is responsible for the depth of colour in liver GSPs. However, I thought it worthwhile to point out that a problem exists (most fanciers prefer the darker livers) and the "C" locus gene alleles provide a possible explanation. I have therefore included illustrations of GSPs with variations in depth of colour, caused by "C" locus alleles. Interestingly enough, there does not seem to be a "pale" version of black, although black dogs certainly can produce light liver offspring.
GENE LOCI FOR COAT COLOUR AND
1. Gene locus "A" (agouti): The alleles at the "A" locus are as follows:-
A All the individual coloured (non-white) hairs are the same colour without shading or lighter
variations. a* "Agouti". Each hair is "shaded" i.e. banded with lighter/darker colour as in such animals as wild
rabbits. Sable and tan as in German Shepherds and Collies. The main colour is a mixture of different colours and shadings. a' Tan points, as in such breeds as the Dobermann and the Rottweiler. Dark saddle with tan markings as in such breeds as the German Shepherd or the Beagle. This gene deals with the amount of colour in the individual coloured (not white) hairs and where they are distributed on the body. Since GSPs have normally only one colour which is not shaded, they are most likely to be "AA", having gained "A" from each parent. It should be noted, however, that the German GSP Standard allows for the "gelber brand", which has been translated as "a slight tendency to sandy colour around the muzzle or feet". I here is some documentation of tricoloured GSPs described (Burns, 1952) as having markings "a la Dachshund". Presumably, such dogs were "a'a'", and their normal-coloured parents were both "Aa"'. In addition, I have seen many GSPs with a band of '. lighter coloured hairs near the ends of their (docked) tails. These hairs are shaded, agouti fashion, leading . me to speculate that such animals are exhibiting one of the "A" series recessives.
2. Gene locus "B" (black): There are only two alleles at the "B" locus:
B Black ;
The black allele "B" is a simple dominant over "b" (liver). There are no intermediate colours, i The GSP can be "BB" (homozygous black), "Bb" (heterozygous black) or "bb" (liver).
3. Gene locus "C" (colour vs colour paling):
There appear to be four alleles at this locus. They are:
C Colour, allowing full expression of the inherited colour.
c Albinism (complete). No colour.
Cch "Chinchilla" or partial albinism. This allele has little or no influence on black but liver animals may show direct influence by being paler in colour. Dr Little uses the example of the Chesapeake Bay Retriever, but the same variation in depth of colour could apply to the GSP. c'1 White coat, with dark eyes and nose as in the Samoyed. The GSP is likely to be "CC", but one could speculate on the possibility of some very light liver
animals being "Ccch or even cch, cch).
4. Gene locus "D" (dilution):
The two alleles at this locus are:
D Dense pigment, as in black or liver coloured dogs. d Dilute, as in blues and greys.
Since the effect of "dd" on liver is to cause the grey of the Weimaraner, and the effect on black is to cause blue, then it appears that the GSP has only "D" in its' makeup, and is normally "DD". I was interested to note in an American magazine that some Weimaraner fanciers were attempting to gain recognition for "blues" as well as greys. One wonders at the veracity of the pedigrees of such dogs, for to have obtained blue coat colour, black must have been used.
5. Gene locus "E" (extension):
There are four alleles at the "E" locus. These are:-
Em "masking", as in the black face mask found in such breeds as the Boxer or the Great Dane. E Dark pigment over the whole coat without "masking". ebr Brindle. e Red or yellow.
It appears that the GSP carries only the "E" allele of this series. It is, however,of interest to remember that early breed history describes the original breed colours as including red. Acceptance of the Schweisshund as being an ancestor would support this, since the Schweisshund varieties include both red and brindle dogs. It is surprising, therefore, that reds or brindles do not appear in GSP litters even if only on very rare occasions.
6. Gene locus "G" (greying):
There appear to be two alleles at the "G" locus:-
G Greying, where the coat becomes progressively lighter, with maturity, as in such breeds as the Poodle or the Kerry Blue Terrier, where a black (or liver) baby coat is, on maturity, blue (or chocolate fawn).
g Normal, no greying.
This effect must not be confused with "greying"
associated with age, or the paling described earlier.
GSPs appear to be only "gg" (normal).
7. Gene locus "S" (spotting .. . i.e. patching):
There are four alleles at the "S" locus. These are:-
S Self (solid) coloured. Small amounts of white on
chest, toes, and/or tail-tip can be present. s' "Irish" spotting. White markings on the
extremities, i.e. legs, neck, tail. This pattern is
commonly seen in such breeds as the Basenji and
the Boxer. sp Piebald spotting. Patches of solid colour on the
head and body, as in the Pointer and most spaniel
breeds. sw Extreme-white piebald. Solid colour is present
only on the head and/or the base of the tail.
The GSP appears to have all four alleles. It can have combinations of these, too numerous to mention. The overlap which occurs can make it difficult to decide which alleles are present, particularly in the case of a solid with very long "socks" and a great deal of white underneath, versus a very heavily marked "Irish spotted" animal. It is important not to confuse the term "extreme-white piebald" with those GSPs which do not have ticking in their white areas. The "S" gene controls the size and location of those areas of (solid) colour which are present at birth, and not those which appear later.
8. Gene locus "T" or "R" (ticking or roaning);
The inheritance of these factors is not as well documented as are those already mentioned. Dr Little postulates two alleles at the "T" locus. These are:-
T Ticking. t No ticking, i.e. clear white.
This simple explanation makes no allowance for the enormous variation in the amount and distribution of coloured hairs which appear in the white areas of dogs carrying genes for ticking or roaning. Such breeds include Dalmatians, Coon Hounds and Australian Cattle dogs as well as GSPs whose coats can be as lightly marked as the former or as heavily as the latter breed. Although the English (and Australian) GSP breed standard considers ticking and roaning to be variations of the same phenomenon, Dr Little distinguishes between them as follows:-
"Ticking"-pigmented flecks on a white
"Roan"-a mixture of coloured and white hairs resembling the type of coat colouring described as "silvering" in rodents.
Dr Little queries the existence of a separate gene for roaning, but has no proof of such. From my experience of breeding and observing GSPs, I believe that, rather than a "T" locus for ticking, there is an "R" locus, with three alleles as follows (Fig. 12.4):-
R Roaning. This is incompletely dominant to and
can be found in combination with:-r' Ticking or spotting (as in the Dalmatian, or
lightly marked GSPs). r Clear white.
The majority of "ticked" GSPs show a combination of ticking and roaning and could therefore be "Rr"'. "White" GSPs with extremely sparse ticking could be "r'r". It is interesting that in the GSP as in many breeds of dog, there is a common tendency to have the greatest density of colour on the back and upper body, with more white to be found on the extremities.
A further gene series, involving Merle, a colour/pattern found in Collies and Harlequin Great Danes, is of no consequence to GSP breeders, since it is not present in the breed. I have not included it in this series, for that reason.
OUTCOMES OF MATINGS, IN REGARD TO COLOUR AND
PATTERN (BLACK vs LIVER AND SOLID vs NON-SOLID ("TICKED")
The following chart lists all the possible combinations of the nine basic genotypes for colour and basic patterning in the GSP. The basic patterning may be either solid or non-solid. The amount of colour in the white portions of the coat (i.e. whether it is ticked, roaned or clear white) and the distribution of the solid areas (i.e. whether the dog has a "saddle" and/or patches or no patches, or whether a solid has a white chest or toes or no white at all) are not represented in the chart.
For the nine possible genotypes, see the illustration. Although that illustration and those which follow, employ "piebald spotted" animals to represent non-solid ("ticked"), one could substitute "Irish spotted" or "extreme white piebald" animals and the result would be the same, for all those versions of non-solid ("ticked") involve different alleles of the same gene "s", and all are recessive to "self "S" gene for solid coloured coats. Whether "solids" have no white in their coats, or have patches on the chest and/or lower legs is not important either, as it does not affect the outcome. In my experience, homozygous solids tend to have no white at all, but since heterozygous solids can also be devoid of white, it is necessary to know the genetic background of an animal in order to classify it correctly. Since there are only three genotypes, (liver/non-solid ("ticked"), heterozygous solid liver and homozygous solid liver) and six possible combinations (see chart) which do not involve one parent being black, only the first six apply to American GSPs. Four of these are illustrated. The last nine involve a parent which is homozygous for both black and solid colouring. Since such animals are rare and not favoured by breeders, no such combinations are illustrated. Combination 15 has been included, for although it is rather unlikely that anyone would mate two solid blacks, this combination provides the greatest number of possibilities. The percentages given represent the average outcome, so individual litters may vary widely from the average. Nevertheless, the possible outcomes are absolute, i.e. no other legitimate outcomes can occur from the possible combinations. For example, one can never obtain a solid from two ticked GSPs
nor a black from two liver-coloured ones (it is most important to remember the distinction between "liver" and "solid" here, since many fanciers refer to solid livers as "livers"), The abbreviations used in the chart are as follows:
Ivr = liver (colour), bik = black (colour) tkd == "ticked" (non-solid, pattern) sol = solid (pattern) het = heterozygous (i.e. one dominant and one recessive allele) horn = homozygous dominant. . . n.b.
n.b. Because the alleles for liver and for non-solid ("ticked") which are recessive must be present in the homozygous state in order for them to appear, I have not specifically referred to them in the abbreviations. If an animal is non-solid ("ticked"),
or if it is liver coloured, it can be assumed that it is homozygous for the genes for that pattern or colour. If, however, it is solid or black, it could be either homozygous or heterozygous for the pattern or colour, hence the extra classification in the chart.
A marker (#) denotes those combinations which are illustrated, with drawings of parents and offspring. Please note that sex has no significance here. Either parent could be of the colour/pattern used in the illustration.
ADVANTAGES/DISADVANTAGES OF INBREEDING
Inbreeding tends to reduce the variation between,
and increase the prepotency of, individual members of a family group. Unfortunately
it also tends to reduce size, fertility and general viability, i.e. less pups
are born and fewer survive, and those which do survive tend to be smaller
and finer boned than the average for the breed. The lower birth and survival
rates may be due to the fact that lethal recessive genes are combined in the
inbreeding process, and some puppies die in utero and are re-absorbed by the
dam, or are born with internal defects which cause their premature death.
Inbreeding also "brings to the surface" defects which are carried by recessive alleles. It must be emphasised that inbreeding does NOT cause such defects, it merely reveals them. The number of defects revealed by inbreeding varies, of course, according to the number of defects carried by the parent stock. It was interesting for me to note that the most important secondary sire in the German Shepherd Dog "Frankie" breeding programme, was Orkos ofLongworth UDT, described by Ginsburg as a descendant of the Fortunate Fields strain (Pfaffenberger, 1963). "Longworth" was the affix of L.C.Brackett, a famous American GSU breeder who was a great exponent of inbreeding (Willis, 1977). Fortunate Fields produced an inbred strain of superior GSDs (Pfaffenberger, 1963), so perhaps Orkas of Longworth's genes had already been rid of some of the undesirable defects which inbreeding can reveal. The use of Orkas may therefore have helped in limiting the number of defects revealed by the "Frankie" inbreeding programme. Although it is always distressing for breeders to produce defective animals, their very presence is a useful indicator of which recessive defects exist in breeders' stock. This knowledge can then be applied in an effort to eliminate those particular defects from their breeding stock. Just as inbreeding reveals recessively carried defects by producing animals homozygous for recessive alleles, it also produces animals homozygous for dominant alleles. If these dominant alleles are responsible for desirable qualities, then the animal possessing those alleles will be prepotent for them, i.e. it will pass them on to all its offspring. For example, supposing a breeder wished to produce only solid coloured animals. The solid coat pattern is governed by the dominant allele "S", whilst non-solid coat-pattern is controlled by the recessive allele "s". Any dog with "SS" would be prepotent for solid colour, even if mated to a non-solid "ss" dog, for their offspring would be "Ss" that is, solid, carrying non- solid. If only solid-coloured dogs were mated together, thus eliminating all "ss" dogs, eventually all the dogs would be "SS", and the desirable solid coat-pattern would be "fixed" in the line. No doubt, this was the method employed by Irish Setter breeders in their quest to remove the original red-and-white coat patterning from the breed.
Unfortunately, the complete removal of recessive alleles is extremely difficult, and probably beyond the lifespan or pocket of any dog breeder. In spite of dozens of generations of selection for the single allele of solid coat colour, non-solid (red-and-white) pups still appear occasionally in Irish Setter litters, as would ticked pups, if solid coats were selected for in GSPs.
If the desirable factor was governed by a recessive gene, it would of course appear only in its homozygous form. Unfortunately, it seems to be a general rule in animal breeding, that the homozygous recessive state is less desirable than the heterozygous state, and many life or health threatening defects appear when two recessive alleles combine (Willis 1977).
The significance of this was brought home to me recently by a very experienced breeder of (English) Cocker Spaniels who detected in some of her dogs a lethal, inherited kidney disease. This appears to be governed by one or more recessive alleles. Due to the extensive use of carriers for breeding, the condition is now present in the breed to an alarming degree, some twelve years after the warning signs first appeared, and it is becoming increasingly difficult to obtain genetically healthy (non-carrier) stock. Although many of the genes controlling desirable factors are unknown, and probably work together in combination to produce such things as shoulder angulation, head type, bodily proportions etc, in the inbred animal whole groups of genes can be "fixed". Such an animal is more predictable as a breeding prospect. Such predictability will not occur in only one or two generations of inbreeding, but each succeeding generation will provide a breeder with important information to be used in the quest for improvement. It is important to remember that if one is fortunate enough to own an outstanding animal, fifty percent of its genes will be lost in each succeeding generation, unless a programme of inbreeding, aimed at preserving those genes responsible for the outstanding qualities of that particular animal is instituted.
GENES VS "BLOOD"
Many, if not most dog breeders refer to the inheritance patterns in their particular breeds as their "bloodlines", and what they pass on to their progeny, their "blood". Some even refer to certain dogs within a breed as having "good blood". Of course what is in fact passed from one generation to another is not "blood" but a particular combination of genes. Nevertheless, the word "bloodline" is widely used by breeders in many countries. The Webster's Dictionary (1956) provides the following definition of the word "bloodline": "a direct line of descent; pedigree; strain: usually of animals". Thus of the three possible definitions, only one ("a strain") implies anything other than random breeding. A pedigree can be the result of endless outcrossings (as are most human pedigrees). A direct line of descent can refer to a single ancestor appearing only once in five generations. The word "strain", however, describes a closely related group of animals, which has been developed via inbreeding. Thus the expression "good bloodlines" may refer to a successful strain (or strains). The word "blood", used, for example, in the statement: "Flossie carries the blood ofHeidi", refers to a particular animal ("Heidi") and its relationship to its descendant ("Flossie"). The statement, however, may not be true, even if "Flossie" is a direct descendant of "Heidi". Below are two imaginary pedigrees for a "Flossie" whose great granddam was "Heidi". There is a great deal of difference, however, in the relative likelihood of whether Flossie A or Flossie B actually "carries the blood (genes)" of Heidi. In Flossie A's pedigree, Heidi is only one of eight great grandparents, each of which could contribute only 50% of their genes to their offspring. In the case of Flossie B, however, who is inbred 2:2,3 to her, Heidi's gene contribution is multiplied. Thus whilst it could be said that both Flossie A and Flossie B were of Heidi's "bloodline", it is very likely that Flossie B will carry Heidi's "blood", i.e. her genes, and quite possible that Flossie A has none of Heidi's "blood" at all. Therefore, whilst open to misinterpretation and sometimes being quite misleading, the terms "blood" and "bloodlines" can be used to describe particular strains within a breed, or the likely presence of the genes of a particular animal in a pedigree.
SELECTION: THE CRITICAL FACTOR
Successful breeding is undoubtedly due in part at least, to good luck, but in the long term, it is also due to careful planning and careful selection. Regardless of a breeder's scientific knowledge, if he or she is successful in producing consistently high quality animals, it is the result of choosing the best breeding stock, and selecting the best of what they produce. Many people believe that all the animals in a litter have the same genes. For example I have heard it said "Why use that dog when you own the brother and using him will achieve the same result?". Those who have read this and other chapters will know by now why this is not true, for no dogs are so inbred that their genes are identical, identical twins are very rare in dogs, and if the brothers in question are the result of an outcrossed mating, the result could be almost as varied as if one used an entirely unrelated dog.
Some breeders, without utilising a high level of inbreeding, seem to have a remarkable level of similarity of type in their kennels. This may be due simply to the fact that they are able to select that particular type from amongst their pups, and choose only sires of that type for mating with their bitches. Stock purchased from such a breeder may not produce a high level of consistency in their offspring, and the type may rapidly be lost, simply because the new breeder is incapable of selecting the type from amongst his or her pups. It is of great importance that breeders make the effort to avoid kennel blindness, for this undoubtedly will hamper them in their selection process. The failure to recognise that serious faults are creeping in to one's erstwhile excellent stock can result in disaster, particularly if the kennel is small and there is little or no access to closely related stock which is free of the defect or defects in question. It must be remembered that even the most closely inbred strain produces entirely outcrossed animals in the first generation following an outcross, and if there are no suitable close relatives to breed back to, the benefits of a lifetime's careful inbreeding could be lost in a very short time. Of course it is true in breeding as in other human activities, "one man's meat is another man's poison", and a dog regarded as entirely unsuitable for breeding by one breeder, may be considered an ideal specimen by a rival breeder. This may be so particularly in the GSP, a breed in which there is, at present, no single universally accepted desirable type-Provided that high quality animals whose appearance, temperament and abilities can be accommodated by the Breed Standard are being produced, there is nothing wrong with breeders producing different types. In fact the rivalry engendered by the different types competing against one another, may be of benefit to the breed. This may also provide future breeders with suitable outcrossed animals to counteract deficiencies in their stock. Selection from an outcrossed mating Since outcrossing, as a breeding tool, is used to introduce a factor or factors in which one's stock is lacking, it is of great importance to select from the outcrossed litter, only those progeny which have the desired feature or features. There is no point in relying on a pedigree to provide the desired factor(s) in a recessive form, for the main object of outcrossing is to acquire dominant genes for further use. The very fact of outcrossing increases heterozygosity, so the desirable factor(s) can only be incorporated into a strain if the outcrossed animals are "bred back" to their close relatives.
It is important to take a long view when outcrossing, and not lose heart if (as is often the case) type and quality in the outcrossed litter are very varied. Most breeders are happy to produce a litter which contains a single outstanding animal even if the rest of the pups are of low quality. If, however, consistent quality over the long term is what a breeder seeks, he or she is not likely to achieve it with constant outcrossing. An outstanding outcrossed animal is less likely to be a consistent producer than an outstanding inbred one. Selection from an inbred mating Because the process of inbreeding tends to produce smaller, finer and possibly weaker animals, the pups selected from an inbred litter should be well up to size, with good substance and a vigorous constitution. Any undersized, weak or defective animals should be eliminated from the breeding programme.
When using inbreeding as a breeding tool, progress will be made only if rigorous selection, that is, keeping and breeding only from the best, and discarding the unsuitable, is undertaken. An outstanding animal, which is inbred and is prepotent for its good features, can be of incalculable value to a breeder and to the breed as a whole. Such an animal is KS & Aust Ch Elk v.Hege-Haus. Elk was a leading sire in Germany in the eighties. KS & Aust Ch Elk was inbred to the v.Wasserschling "A" litter (Axel, Anni and Adda) and Imme v.Hanstein appears twice on his sire's side, although as mentioned earlier, Imme, on one side only, may have been of little significance to Elk. Axel v. Wasserschling, the dog which above all others, provided the breed with the necessary characteristics to revive it after World War II, appears in Elk's pedigree more than any other ancestor. Since many of the top German bitches with which Elk was mated also owed to Axel, the value of Elk as a sire was increased, for their progeny were also inbred and as a result, were consistent in type.
A RECIPE FOR SUCCESSFUL BREEDING
Breeders, when they first embark on a breeding programme, are usually motivated by the desire to breed dogs just like their beloved first one. In order to be successful as breeders in the long-term, however, they need to develop the ability to evaluate critically the faults and virtues of that beloved dog, to discard or minimise the faults, and to capitalise on and improve upon the virtues. They should also build up in their minds, a picture of the ideal GSP, as a goal to be sought after. Of course, no dog is perfect, but the quest for perfection is made more difficult without a clear idea of what that perfect dog should be.
The best way to achieve success without losing all that one started with in a couple of generations, is to inbreed, using high-quality close relatives of the best dog available, and selecting the breeding stock from the best of their offspring. It must always be kept in mind that defective pups will be born in the best of litters, and a breeder must be prepared not only to discard them but to ensure, for the good of his or her reputation and for the future of the developing strain, that they are not bred from.
When faults appear as they invariably will in an inbred strain, the breeder must be prepared to recognise them, discard the affected individuals and if necessary, resort to an outcross. A-preferred option in this case would be to use a dog which is itself from a strain excelling in the area in which the breeder's own strain is weak. The resulting outcrossed progeny should then be crossed back into the original strain to ensure that the benefits are not lost in future generations. Consistency in breeding usually comes from judicious inbreeding and always from a rigorous selection process. Patience and the ability to cope with disaster are also essential ingredients for long-term success as a breeder, for disappointments abound. It is also important to collect information on as many as possible of the animals which appear in the pedigrees of one's breeding stock, and of their relatives, both good and bad. A title may not reflect the breeding worth of an animal. It must be judged on its ability to contribute the particular characteristics needed or deemed desirable in one's breeding programme. Many breeders store enormous amounts of such information in their heads, but it makes sense to keep written records too, for much can be forgotten, and much can be useful later which did not seem worthwhile remembering at the time. Therefore the keeping of written records (or if available, computerised data storage) is recommended. In addition, photographs can be very useful, although it must be remembered that these can be deceiving, and if supplied by the owner, usually show the dog only in its most flattering light. Therefore it is useful to obtain a variety of photographs of dogs which are of particular importance to a breeding programme, and of the breeding stock at various stages of their development. It is critically important to remember that one is breeding living, breathing animals, not pedigrees, nor show/trial records, nor glamorous photographs. Nothing beats a thorough knowledge of the stud-dog himself, and the offspring he has produced, to relatives and non-relatives of one's bitch, as a tool in assessing his suitability as a mate for her. Often it makes more sense to consider using the sire of a wonderful but untried young dog, than to use the young dog himself, particularly if his sire has produced more than one or two good ones. A sire is only as good as his offspring, and the son of a prepotent sire may not necessarily be prepotent himself. A professional approach is most important if one is to succeed as a breeder. One must take the responsibility for one's mistakes and failures, and make an effort to compensate the buyers of one's pups if, through no fault of the buyer, the pup proves to be unsuitable for the purpose for which it was purchased. One must also be prepared to help and encourage the buyers of one's pups to get the most out of them.
Perhaps most important of all, one must provide them with a pup which is properly reared, both physically and mentally and in the best of health with an appropriate worming and vaccination programme rigorously adhered to. In order for those carefully manipulated genes to reveal their maximum positive benefits, they must have been and continue to be in a good environment.
Stud-dog owners have their responsibilities too. Apart from keeping the dog in good health, he must not be overworked, with so many bitches accepted close together that his fertility will suffer. He must also be trained to do his job, so that the inevitable aggressive or terrified bitches can still be served, without risk to him. His owner must learn to diagnose precisely the readiness of a bitch for mating, in order to avoid costly and unnecessary failures, when bitches are sent or brought long distances to him.
If a series of matings reveals a dog to be a consistent producer of a serious defect, or the carrier of a lethal gene, a conscientious breeder must be prepared to withdraw him from stud duties. An extremely popular stud dog so affected, can wreak incalculable harm on a breed, as numerous examples in dog breeding will testify. In the long term, such a dog may leave breeders without viable alternatives to counteract the damage he has caused. It is unlikely that very rapid progress can be made in producing a consistent and successful strain within a breed, unless one is prepared to breed and discard a large number of dogs. The reason for this is the necessity for inbreeding to fix type and the inevitable setbacks that inbreeding provides. If an individual breeder is unable to retain a fairly large number of breeding animals, one alternative is to cooperate with a group of people willing to work together to produce a jointly-recognised desirable type. Another alternative is to embark immediately on a very close inbreeding programme, based on only one or two selected dogs. The selection of breeding stock must be made with more than just conformation or narrow performance requirements as criteria, for if general mental or physical health requirements are not given top priority, disaster can ensue. As Fox (1978) points out: "Gradual degeneration. . . is occurring in those breeds of dog where man does not mimic the processes of natural selection whereby normative structural, physiological and behavioural functions are maintained and stabilised in the species or breed variety over generations". Inbreeding, per se, is not primarily to blame for this situation. It is "overbreeding" or undirected selection (Fox, 1978) which is the major culprit.
The minimum requirements for real breeding success may well be as follows:
A. A long-term goal (probably ten years' effort is a minimum) and the ability to meet some fairly hefty expenses.
B. The collection of information, and the formation of a breeding plan (which must be Hexible, to allow for the inevitable setbacks).
C. Tenacity, patience and a constant effort to avoid developing kennel blindness (the inability to perceive the faults of one's dogs, and/or the virtues of the dogs produced by one's rivals).
D. A professional approach which leads to the establishment of a reputation for honesty and fair-dealing.
E. The courage to discard a favourite dog which will do the breed more harm than good, the self-confidence to cope with the back-stabbing and unfair criticism which accompanies success and last, but not least, a dash of good fortune.