Chapter XXI
SHORT-HAIRED BIRD DOG
Taken from Inheritance in
Dogs with Special Reference to the Hunting Breeds, by Ojvind
Winge, published by Comstock Publishing,
(NOTE: This study was conducted using Volumes 55 to
67 of the Danish Studbook; therefore, some of the terms are not what we are
accustomed to. For example, “self-colored” is a term now called “solid”; “speckled” is now
called “ticked” in most countries, while “mottled” is a term that is now known
as “patched”. You may also wish to
refer to chart at the conclusion which gives explanations for the various gene
designations.)
That
character which has given this breed its name is the short hair lying close to
the body which is due to the dominant factor K. The whole breed has thus KK.
Furthermore the dogs always possess EE
and are therefore never red or yellow.
On the other hand, B may or
may not be present. B + E
gives black animals; E alone, brown.
The
breed includes self-colored (TT or Tt)
and mottled (tt)
individuals. In addition, the white
parts produced by the tt-forumula
may be pure white, rr or dappled speckled (RR or Rr). The S
gene is sometimes apparent; if it is present alone in tt dogs, it produces in them
small, distinct black spots in the white areas; if it occurs together with R, the dogs become dappled-spotted with
small darker spots. The presence of S is difficult to establish in the
dark-dappled R animals. Finally, in brown dogs, both a darker brown
and a lighter brown, presumably a recessive shade, are found. The gene Z,
so named by Steiger (1936), is found in nearly all
Shorthairs. If it is lacking and zz occurs, the
brown coat is modified to a cinnamon color. Thus, in the offspring of 90368 Freja of Helhoejgaard (brown) and
64829 Flok (brown-speckled with brown markings) nine
brown and two cinnamon-colored puppies occurred.
“Bicolor” markings occur only rarely in the Short-haired
Bird Dog, as mentioned by Buchwald (1945), who designated the markings as being
“a la Dachshund.” The father of the famous
35447 Bob (Koege) thus had what the Studbook calls
“hound markings.” 31845 Hestehavens Rap was described in the Studbook as
“brown-speckled with brown spots. Tricolored head. Hound markings.” A brother to Rap had also, “Tri-colored head. Hound
markings on cheeks and ear,” while five others from the same litter
were brown-speckled with brown spots. Of
the parents of this litter, the Studbook merely states that the father was
“brown-speckled with spots”; no color is given for
the mother. From the standpoint of
inheritance it is not surprising that the bicolour markings, which are of a
recessive character, and which can therefore be concealed, may turn up in some
dogs.
All the
short-haired dogs in Volumes 55 to 67 of the Danish studbook formed the basis
for his investigation.
As is
known, there are fewer black Shorthairs than brown, and fewer self-colored than speckled.
(Some animals in the studbooks are called dappled, but there are many
more designated as speckled; in the following, we will use the most frequently
employed designation – speckled.) Hence
it follows that self-colored black are rare, even
when one correctly considers as self-colored those
animals that have a white or speckled chest spot or a little white or speckling
on the paws. As previously mentioned, T animals (especially Tt) often have
these localized markings, which are quite apart from the concept of “white
mottling.” Actually, many self-colored Shorthairs have these chest
and paw markings.
In the
thirteen volumes of the Studbook there occurs only a single completely black
breeding animal, the bitch 51966 Toboels Svarta and only three black offspring individuals – 55948 Palle of Horndrup and 60336 Toboels Rubbi, both of which,
however, had a speckled chest spot (the spot shows the presence of the R gene), and 64433 Mona of Tungepil, which had a small white swirl on the chest. Only a very few black-and-white-moddled are found, but many black-speckled – that is,
black-and-white-mottled with the gene R
– are listed. By far the greatest number of Shorthairs are brown, and, of these, most are
speckled, so that the most common formulas are bbEETtRRKK and bbEEttRRKK. Since the brown-speckled dogs are often
rather dark, even in the speckled areas, it is not often very apparent that
they are all actually mottled of brown and speckled parts. The self-colored
brown parts are often confined to a spot above the base of the tail, the ears,
and a single spot on the body.
One
finds also that speckled dogs are sometimes designated “speckled” and sometimes
“speckled with brown markings.” This is
due to inadequate description. All
speckled Shorthairs have self-colored areas; none are
speckled over the whole body.
In
itself, inheritance in the Short-haired Bird Dog is of a very simple type, and
for this reason we selected this breed as an example of one, two, and three
gene inheritance in Part One.
Black X
black is naturally able to give black or brown offspring, for BbEE can
segregate bbEE,
but brown X brown can never produce black offspring. However, in the volumes of the Studbooks
examined, not a single case is found in which two black have been mated
together. Had it occurred, a segregation
of 3 black : 1 brown would presumably have occurred,
for all black (or black and speckled) Shorthairs are certainly heterozygotes, BbEE, and not homozygotes, BBEE.
If
black is crossed with brown, and if the black is BbEE and the brown bbEE, 50 per cent
of each color should occur in the offspring. The sparse material from this crossing shows
8 black : 7 brown.
The
gene for mottling (white-mottling) is recessive, tt, which means that self-colored can segregate out mottled (to which category all
speckled also belong), but mottled (including speckled) cannot segregate out
self-colored.
An
examination of the parents and offspring in the Studbooks shows the data given
in Diagram 35.
Diagram 35
|
|
Self-colored
brown (with or without chest spot) |
Speckled |
Brown-and-white-mottled |
Total |
|
Self-colored brown X self-colored
brown (with or without chest spot) |
188 |
18 |
5 |
211 |
|
Self-colored brown X speckled |
165 |
118 |
6 |
289 |
|
Speckled
x speckled |
(10) |
373 |
19 |
402 |
|
TOTAL |
363 |
509 |
30 |
902 |
In this
summary the crossings with brown-and-white-mottled dogs are not taken into
consideration, for these are often doubtfully described. They are called “brown-and-white-mottled,”
“brown-spotted,” “brown-mottled and speckled,” or “brown-mottled with brown
splashes,” and so on, and it is not always possible to form a very clear
picture of the actual appearance of the dogs.
Since the true speckled (those which have more than just a chest spot or
speckled paw) are, however, always white-mottled, tt, with regard to heredity, the
inheritance of white mottling can very well be tested by examination of them.
Diagram
35 shows the following: self-colored brown X self-colored
brown gives predominately self-colored brown and only
a few speckled, which shows that many self-colored
brown must have the formula TT. Had they all been Tt, a fourth of the offspring
should have had tt
– that is, brown speckled or brown-and-white-mottled. Here, however, only 23 out of 211 have tt, which
indicates that about 30 per cent of the self-colored
have TT, while the rest have Tt. In the crossing together of self-colored brown there should then be segregated out 26 with tt from a total
of 211. As previously mentioned, the
difference between the brown-speckled and the white-mottled is that the former
contains the R gene and the latter, rr.
The
cross, self-colored brown X speckled, also shows that
several self-colored brown have TT, for more than half of the offspring are totally brown. According to the segregation counts, for
every 124 Tt-dogs
there are 41 with TT or 25 per cent
of all self-colored brown dogs.
Speckled
X speckled (tt
X tt),
which properly indicates a cross between dogs that are speckled or dappled with
large brown spots, should give nothing but speckled, tt (or brown-and-white-mottled,
when R is lacking). It is seen that there are 10 dogs that are
described as “brown,” and these therefore should not occur. The presence of 10 out of 402 is naturally
not more than what would be expected through error; and that mistakes exist in
the statements becomes quite clear: In
the first place, 4 of the 10 are from one litter; this has probably had a
father other than the one quoted. In the
second, the remaining 6 are all from the same bitch, 51054 Lyna-Jesa
of Havrevaenger, which is designated “brown with
speckled markings.” It must be assumed
that the bitch had only a speckled spot on the chest and perhaps on the paws,
and it therefore should be considered as self-colored
brown (with T).
Among
the brown-and-white-mottled offspring in the table, some are not clearly
designated, for six are called “brown-mottled,” one “light-brown-mottled,” and
two “brown spotted.” Of the speckled,
two are called “brown-speckled with white markings,” but it should be noted
that small, almost completely white, spots on the forehead or chest stripes may
be present in speckled dogs. These dogs
are therefore assumed to be correctly classified.
To
examine the offspring from brown-and-white-mottled X brown-and-white-mottled
with the help of the studbooks involves, as suggested, uncertainty because one
cannot always know just what a definite color
designation covers. As a rule,
“brown-mottled” may well be assumed to designate brown-and-white-mottled, and
there can be no doubt that the designation “brown and white” may also cover
it. If the offspring of dogs from these
two categories are examined, there is found a total of thirty-two
brown-and-white-mottled, together with one “brown-mottled-speckled” and one
“brown with a gray chest and gray
paws”; these two should not occur, for mottled without speckling (ttrr) should not
be able to segregate out mottled with speckling (ttR); but since, as mentioned,
the designation of the parents is uncertain, one must be content with the fact
that, according to the studbook, nearly all individuals have that color which they theoretically should possess.
A
strange individual is that discussed by Buchwald (1945) – “black-brown” 28465
Bruno-Hubertus-Rosengaarden. The studbook calls him “dark brown with
speckled forepaws.” There seems
therefore to be no doubt that the dog did not look black but brownish. Nevertheless he had given black offspring
with several brown bitches, which is inadmissible in Shorthairs. With the brown 25745 Nora Kalbyris
he produced the black 31881 Svarta of Vesterlund; with the brown-speckled 29544 Ketty-Claushoej he gave among others, 30737 Pan of Noerlund, which was black with a speckled chest; and with
28839 Wanda-Claushoej, which was brown with speckled
forelegs, he gave 30138 Pikas of Langaa,
which was black with speckled paws.
Bruno-Hubertus
has the pedigree shown in Diagram 36.
|
Diagram 36
|
It is
easily seen from the pedigree that Bruno-Hubertus cannot have Tycho as a father, as is stated. Bruno-Hubertus’ mother, as well as his stated
father, had the factor for white mottling, tt, in agreement with the fact
that their parents had also possessed it.
Bruno-Hubertus was, on the other hand, dark brown with speckled forepaws
and therefore was “self-colored”; he had a T, which neither of the dogs stated to
be his parents possessed. Since there is
scarcely a possibility that the statement with regard to the dog’s mother can
be incorrect, the only solution remaining is that it had a father other than Tycho. This is not
only a possibility but a necessity.
Bruno-Hubertus was actually self-colored (with
speckled forepaws), which also appears from the fact that he has produced self-colored offspring (30737 Pan of Noerlund)
with brown-speckled bitches – e.g., 29544 Ketty-Claushoej.
A
closer examination of this mysterious Bruno-Hubertus reveals that there has
always been some suspicion concerning his origin on the part of some of the
professional breeders. The dog was Pointerlike and much more slender and fast than the
Short-haired Bird Dogs were at that time.
This is clearly revealed in a photograph of the dog. Taking everything into consideration, there
is no doubt that Bessie Claushoej was mated to a dog
of another breed, and that therefore genes were introduced which resulted in
Bruno-Hubertus being self-colored instead of
brown-speckled and in receiving a coat of a divergent nature – a brown-black color, which in inheritance is black, but which become
brownish because hereditary units of unknown nature were introduced by the
irregular relation.
Let us
attempt to calculate the genetic formulas in some Shorthairs on the basis of
the statements in the studbooks. We
shall choose the black-speckled 64323 Lis Mona, whose
pedigree is given in Diagram 37.
Since
all Shorthairs constantly have the factor for the brown color
present, they must have EE; otherwise
red or yellow animals would frequently occur in the bred, which
is not, however, the case. The
consequence is that at least one of the parents of each black Shorthair is
black, while in the Setter, Pointer or Spaniel a black dog may occur from the
crossing of brown with red. Since, in
addition, all mottled individuals among Lis Mona’s
ancestors are speckled, it is likely that they all
have RR.
Lis Mona’s father, 55948 Palle of Horndrup, was black with a speckled chest. Since only the chest was speckled, while he
otherwise was not so, he had T in his
formula, for if he had had tt, he would have been also speckled over large areas on the
body. He must therefore have TT or Tt. Since one of his parents, the mother, 42680
Lady II, was black-speckled and accordingly had tt, her son must have received a t from her, and consequently Palle had Tt in the formula. We
know now that he was EETtRR. Finally, as to the B gene, he must have had Bb,
for only his mother, Lady II, was black, while the father, 40903 Gelsteds Rino, was brown; from
him, accordingly, he must have received a b. Thus Palle’s
formula is completely worked out. On the
maternal side all are brown-speckled, so that all the animals here must have bbEEttRR.
From
the formulas it appears that Lis Mona and her
possible full brothers and sisters should theoretically show a segregation in
the ratio, 1 BbEETtRR
(black with speckled chest) : 1 BbEEttRR (black-speckled) : 1 bbEETtRR (brown
with speckled chest) : 1 bbEEttRR
(brown-speckled). Nothing is to be
found, however, in the studbooks on Lis Mona’s
brothers and sisters.
The
nose in the Shorthair is brown in brown dogs and black in black dogs.
|
Diagram 37
|
The
various types of Short-haired Bird Dogs have the following formulas:
|
bbEEttrr |
1. Brown-and-white-mottled; the white pure
white (Plate III, figure a) |
|
bbEEttR |
2. Brown-and-white-mottled; the “white” parts
brown-speckled; consequently, brown-speckled with brown spots (Plate III,
figure e) |
|
bbEETrr |
3. Self-colored
brown, possibly with a pure white chest spot and white paws (Plate III,
figure c) |
|
bbEETR |
4. Self-colored
brown, possibly with a speckled chest spot and speckled paws (Plate III,
figure d) |
|
BEEttrr |
5. Black-and-white-mottled; the white pure white
(Plate III, figure b) |
|
BEEttR |
6. Black-and-white-mottled; the “white” parts gray-speckled; consequently gray-speckled
with black spots (Plate III, figure g). |
|
BEETrr |
7. Self-colored
black, possibly with a pure white chest spot and white paws |
|
BEETR |
8. Self-colored
black, possibly with a gray-speckled chest spot and
gray-speckled paws (Plate III, figure f) |
A
formula as, for instance, no. 3 thus indicates that these dogs have bb and EE, and either TT or Tt, together with
rr.
There
is also the gene S which causes small
dark spots in the white parts of white-mottled dogs. A dog of the formula bbEttrrS is, accordingly,
brown-and-white mottled with distinct, small brown spots in the white; such
dogs are common (Plate III, figure h).
Finally,
brown dogs can be light or dark brown.
The latter shade seems to be dominant.
(Note: the above dogs are labelled as:
|
(a) Brown-and-white-mottled
(bb E tt) |
(b) Black-and-white-mottled
(B E tt) |
|
(c) Self-colored brown with a white chest spot and white paws (bb
E T rr) |
(d) Self-colored brown with a speckled chest spot and speckled
paws (bb E T R) |
|
(e) Brown-speckled
with large brown spots (bb E tt R) |
(f) Self-colored
black with a gray speckled chest spot (B E T R) |
|
(g) Gray-speckled
with large black spots (B E tt R) |
(h) Brown-and-white-mottled
and with small brown spots caused by the belton
gene (bb E tt S) |
The
essential rules of inheritance thus are:
|
Brown
X brown |
always
gives brown, never black, offspring |
|
Black
X black |
can
give either nothing but black offspring, or 75 per cent black and 25 per cent
brown |
|
Black
X brown |
can
give both black and brown, as a rule 50 per cent of each color;
rarely nothing but black offspring |
|
Self-colored X self-colored |
can
segregate at most 25 per cent white-mottled or speckled |
|
Pure
white-mottled X pure white-mottled |
gives
nothing but pure white-mottled |
|
Speckled
X speckled |
Never
gives self-colored, but either nothing but speckled
or at the most 25 per cent pure white-mottled |
|
Self-colored X speckled |
Gives
either nothing but self-colored or 50 per cent
self-colored and the rest speckled or pure
white-mottled |
|
Light
brown X light brown |
Gives
nothing but light brown offspring |
|
Dark
brown X dark brown |
Gives
either nothing but dark brown offspring or 75 per cent dark brown and 25 per
cent light brown |
|
Light
brown X dark brown |
Gives
either nothing but dark brown offspring or 50 per cent dark brown and 50 per
cent light brown |
Owing
to incomplete designation of the various shades of brown in the studbooks it
has not been possible to verify the last three assertions.
The
following is a list of the hereditary factors (taken from Chapter XIII,
Inherited Factors (Genes) in Dogs) listed in alphabetical order, according to
their designation:
A.
Epistatic red factor, which produces a red color in dogs possessing the factors for a black or a brown
coat.
B.
(Yellow) red factor. aaB-dogs are red or yellow-red, tan-colored.
C.
Self-colored-coat factor, in contrast to
a series of color patterns:
cbi (“bicolour”), which produces two
colors in both black and brown dogs – i.e., lighter,
red-yellow-brown, tan-colored in black dogs (black
and tan) and corresponding, but usually more yellow, markings in brown dogs
(liver and tan). [It is sometimes stated
that red dogs can have bicolored markings; they may
thus be called “red and lemon.” Mrs. G. Pouw-Neumann of Voorthuizen,
csa (“saddle”), which limits the black coat to a
saddle-shape or covering on the back, as in the Airedale Terrier, while the
rest of the animal is tan.
cbr (“brindle”), which limits the black coat to tigerlike, black cross stripes on a lighter tan-colored background, as in certain Boxers and Great
Danes. In addition, these animals always
have a black nose, and the black color is extended to
form a mask.
cma (“mask”), which limits the black coat to a mask,
while the rest of the dog is red-yellow, tan-colored.
Ct, which is dominant over all the others and produces
total blackness in dogs possessing the character for black clor. The gene is epistatic
to the wild-color gene G.
There are thus six alleles in this series, and they are dominant
over each other in the following order:
> csa
Ct > C > cbi > cbr > cma
I have observed a hybrid between an Airedale Terrier with csa and
a Boxer with cbr. It was distinctly wire-haired and brindle
without “saddle,” which shows that cbr is dominant to csa. It is not known how csa behaves with
respect to cma.
D.
“Dilution” factor, the gene for normal color
formation. The dog with dd receives a
greatly diluted color. D
is therefore a recessive dilution gene.
Black dogs with dd
become “blue”. See also gene Z.
E.
Brown factor, which in itself produces a brown coat – lighter or
darker chocolate brown or liver brown.
The tip of the nose and the balls of the feet are brown. E +
B produces a
black coat through a complementary effect.
F.
Albino gene; F is
essential for coloring. Dogs with ff
are practically white even when, according to the remaining gene combination,
they ought to be colored. In ff
animals all colors are bleached to such an extent
that they are practically invisible.
Only the coat color is affected, while the eye
color and nose color are
unaffected. True albinos with red eyes
are rare and are due to another recessive albino-gene.
G.
Wild-color factor, which produces in dogs
with the factor for black (B + E) a wolf color
and in brown dogs (E), a brownish
wild color, i.e., the colored
hair becomes white in the middle.
Shepherds often have the G
gene. The gene is hypostatic to
H.
“Harlequin” factor, the gene that produces in the Great Dane, Dunkerhound, Dachshund, and others the well-known harlequin
pattern – irregular, large or small, roundish black
spots on a white or gray background in dogs
possessing the factor for a black coat (B
+ E).
The gene in a double dose causes deafness, blindness, and so on (see
page 45).
I.
“Interaction” factor, which exerts an effect in dogs having cbicbi,
as the bicolour (black and tan) character is not exhibited when I is
lacking. Therefore black and brown dogs
that have cbicbi,
do not become black and tan, or liver and tan, when the animal possesses ii.
Neither are the dogs with the tricolour factor tricolored
if they lack I. Black dogs with cbicbiTii
(see gene T) are black; black dogs
with cbicbiTI
are black and tan. White-mottled dogs
with cbicbittii
are black-and-white-mottled, but when they have cbicbittI,
they are tricolored.
K.
Short hair is dominant over k, long hair. Probably there are more alleles and also
modifying genes to be found which cause varying degrees of short-hairedness and long-hairedness.