Having considered variation, which
makes breeding possible, we are lee
naturally to a study of heredity
which makes breeding effective. We have
already discussed the germ plasm theory of
heredity, the most reasonable explanation for
the observed facts, & we must now investigate
these facts to see what practical benefits
we can derive from the knowledge gained by
careful experiments & painstaking researches.
The very simplest case of heredity is the
" pure line," or when reproduction is by means
of self-fertilization. Although such reproduc-
tion is beyond the scope of the Bulldogs breeder's
operations, still, because it throws much light
on the nature of heredity, it will repay us for
a moment's investigation.
Professor Johannsen made the original experiments in the stidy of pure line heredity
with peas & beans, & his results have been, Checked & confirmed by other observers.
He took the beans from nineteen plants, each
if which had been self-fertilized, as is the usual
condition in this variety, & he measured them
for variability in weight, breadth, & other details.
The whole lot showed a standard variation, & the curve he plotted from his data was
close to the normal variability curve. He
kept carefully distinct the seeds from each
parent plant, & grew each pure line, or in
lother words, each lot of seeds from each self-
|fertihzed plant, separately. The plants raised
|in each pure line produced seed that had each
Its own distinct average, differing from the
[general average of the nineteen original par-
ents. Moreover, he divided each pure line
Into lots according to size, but the same sized
'seeds from the same pure line produced seeds
that were not of their own size, but nearer to
the average size of the pure line from which
both they & their parents had sprung.
This means that even within a pure line,
when the germ plasm is, of course, unaltered
& constant in succeeding generations, there is
always variation. Variation Is Inherent In the
germ plasm, even when It In Itself Is unchanged, for seeds larger than the average
I produce seeds smaller than themselves & closer to the average of the pure line. Other words, individual characteristics,
even a pure line when there Is a sexual reproduction, do not reproduce in the offspring.
For the breeder this means that no amoun
of selection In a pure line would ever product
a strain that would regularly have larger seed:
than the average. In pure line breeding then
a very definite limit to what can be accom
plished solely by selection Is set. The great
significance of these experiments for the dog
breeder is the proof they present that even
when the germ plasm is unchanged there is al-
ways variation.
In the case of bisexual reproduction, which
Is the condition under which the bulldogs breeder
works, two different germ plasms are united to
form the new generation. The germ plasm
is no longer a constant, the inheritance is dou-
ble, from two germ plasms, & the case is obviously more complicated.
In an exhaustive statistical study of the
height of 205 parents & their 928 children,
Galton was able to analyze the results of bi-
sexual reproduction. He arrived at conclu-
sions that are of inestimable value to bulldogs breeders.
He found that :
I. Like parents beget unlike offspring, & vice versa, like offspring come from unlike parents. Abnormally tall parents, he found, had
tall, short, & medium sized children; while of all the tall children studied, some came from tall, others from short, & others from medium sized parents.
2. Offspring are on the average nearer to
the average of the race than their parents.
The average height of the children of two
parents is not the mean between these parents.
When the average height of the two parents is
above the average of the race, the children's
average will be shorter than their parents, &
so closer to the race average.
From the first conclusion It follows that two
bulldogs of very different type may be whelped In
the same bulldogs litter, a fact to which any practical
breeder will readily testify. Also, that two
bulldogs of remarkably similar type may have been
bred from very different parents, another fact
that is corroborated by common experience.
The general conclusion emphasizes the value
of a good working knowledge of the true meaning of a pedigree & shows the utter foolishness of any attempts to judge the offspring by
the parents, or the parents by a consideration of the points of the offspring. Almost daily
we see bulldog fanciers attempt these two Impossibilities. It Is common to hear a man say,
'' No, I never saw such-and-such a bulldog, but
judging from his bulldog pups he must be a shortbacked one with good legs & feet," or something of that sort; while it Is even more usual to hear a man say that " So-&-so has the best eyes & ears of any dog at stud, & he certainly ought to get pups good In these points."
The second conclusion of Galton's studies is the principle of regression, or the drag of the race. To the bulldog breeder It presents vitally the true value of a pedigree. To return again to our example of the Fox Terrier head, which in the last chapter we assumed would average six Inches long with an ideal length of seven inches. It would surely result in disappointment to breed together two dogs with ideal seven inch heads, for regression would bring the average of the resulting puppies back closer to the average of the race, which would be closer to six inches. Conversely, a bulldog & a bitch with five inch heads would, on the average, produce puppies longer headed than them-selves, for the average of the offspring would by the same law approach nearer to the average of the race, which in this case is an Inch longer than the Immediate parents.
Could a complete refutation of the Ideas usually followed by bulldog breeders be more forcibly expressed? We are so very prone to
cant about " like produces like," & so very willing to accept a pedigree, which at best is only a guaranty of purity of blood, as proof
positive of uniformity in type.
Plainly, there is but one way to cut loose from the drag of the bulldog race. Bring the general bulldog race average as close as possible to the ideal expressed in the Bulldog Standard. In this way, & only in this way, can regression be won over from an enemy to an ally. If a Fox Terrier breeder should by continued & careful selection raise the average of his own strain from the general race average of six inches to the ideal average of seven inches, he would not have to worry about length of heads so long as he exercised enough selection to hold the very great advantage he had gained.
Besides the pure line inheritance & the bisexual inheritance there is another, named after the man who discovered it, Mendelian in-
heritance. Mendel, an Austrian monk, studying the crossing of different varieties of garden peas, made important discoveries that were
quite unappreciated for thirty-five years. In 1900 his work was rediscovered and confirmed by De Vries, Tschermak, & Correns, each
working independently. The garden pea shows sharply differentiated characters in its different varieties. Mendel crossed these &
observed the way that these different characters were inherited in the hybrids. Mendelian inheritance then is primarily the inheritance
of hybrids, or cross-breds, but subsequent study has shown that many individual characters in straight bred animals follow this same law.
It is beyond our needs to go into all the technical details of Mendel's experiments, or to know how he succeeded in being sure that he
was crossing certain plants. We will confine ourselves to the results obtained in one particular case. Crossing the tall variety, which
is about six feet high, & the dwarf, which is about a foot & a half high, Mendel got a generation of plants, every one of which was
just as tall as the tall parent. This is certainly not what one would naturally expect, for we generally look upon cross-breds as a combination of their parents, & we would think the offspring of the tall & short varieties would be about four feet tall. These tall cross-breeds were allowed to fertilize themselves, which is the usual method of reproduction, and from the resulting seeds a second generation was raised next year.
This second hybrid generation behaved in a truly extraordinary manner. Many plants were just as tall as their tall parents & the
tall half of their grandparents; others, however, were just as short as their darf grandparents. There were absolutely no plants of
intermediate height. What is even more remarkable, the dwarfs bore a constant numerical proportion to the tails. There was one
dwarf to three tall, or twenty-five per cent, of the second hybrid generation were dwarf. In the next & in all succeeding generations,
these dwarfs continued to produce only dwarf plants. Here was a hybrid, breeding absolutely true, a perfect dwarf produced from
tall parents, produced in turn from crossed tall & dwarf.
The seventy-five per cent, tails in the second hybrid generation behaved very differently. Self-fertilized, some produced both tall &
dwarf plants, while others produced only tails. It was found that twenty-five per cent, of the apparent tails of the second hybrid generation were true tails & continued to produce tails indefinitely. Those seeming tails, half of the whole second generation, continued to produce both tails & shorts, In the ratio of twenty-five per cent, true dwarfs, twenty-five per cent, true tails, & fifty per cent, seeming tails, but in reality hybrids in inheritance.
To sum up the results of this important discovery: the first cross between tall & dwarf produced all tails. Mendel expressed this by
saying that tallness is in this cross dominant & dwarfness recessive. This hybrid, tall-dwarf, but tall looking generation, produced
twenty-five per cent, true dwarfs, twenty-five per cent, true tails, & fifty per cent, hybrids with tallness dominant. This is known as
segregation, or the sifting out of the offspring in definite proportions of the characters employed in the cross. This proportion is 1:2:
These same results can be expressed in a chart:
I
Td
TT Td Td DD
I I I I
TT TT TT TT TT Td Td DD TT Td Td DD DD DD DD DD
The first cross is represented by the letters
T (tall) & D (dwarf). The fact that tallness is dominant & dwarfness is recessive is
represented by the symbol Td, a tall looking plant in which dwarfness lies recessive or hidden. The true tails (TT) & the true dwarfs
(DD), which are later segregated, continue, as is shown, to breed true.
This principle of segregation, or the splitting up of the off-spring into the Mendelian ratio of 1 : 2 : I is a fundamental part of this type of inheritance. Dominance of one character over another does not invariably occur. Sometimes there is a blending. In such cases the symbols in the chart would be changed from Td to TD, representing in this particular case a plant of intermediate height. The subsequent splitting off into true tails & true dwarfs would be the same in each case.
Beyond all further doubt Mendel's law has been demonstrated to hold in its mathematical relations. The breeder, however, must remember it applies only to one character or set of characters, not to the entire individual. Judged as individuals, the Mendelian nature of
a hybrid cross might not be at all apparent, though each character be following strict Mendelian inheritance. Some characters would be
dominant in one parent, others in the other parent, and still others might be a blend. In this way, the different characters, viewed as a whole, would seem a hopeless muddle. For this reason the true nature of such inheritance was so long obscured, on-ly to be discovered by the careful Isolation & study of each character by itself.
The bulldog breeder can make no use of Mendel's law until he establishes what characters in dogs, If any, follow It. That there are such characters Is highly probable. Color in chickens, pigeons, rabbits, guinea pigs, & cattle; hair & eye colors in man; presence or absence of horns in cattle; the shape of the comb in chickens, & many other similar characters have been found to follow Mendellan inheritance.
A. L. Hagedoorn has done some work on color Inheritance in Dachshundes, & C. C. Little has made a statistical study of coat colors in Pointers from data In the A. K. C. Stud Book. Their work, which supplements the rather scanty data of Professor A. Lang, indicates that black & brown (liver) follows the same Mendellan Inheritance observed In these colors In mice, guinea pigs, & rabbits.
Dr. C. G. Darling believes eye coloring in Airedale Terriers Is Mendellan, the light color being dominant. He acknowledges that he has
not sufficient data to either prove or discredit this hypothesis, but, as an eye specialist and a terrier breeder, his opinion bears weight. If he is correct, it is probable that all eye coloring in dogs follows Mendellan Inheritance.
It Is also probable that the smooth & broken coats in Fox Terriers, a form of cross breeding that Is common, is Mendelian, the
broken coat being. In this case, dominant. The red & black coloring in Chow Chows & self colored spaniels is also probably according to Mendellan inheritance. However, before a positive statement can be made in any of these cases, more evidence Is required.
Such evidence would be a valuable contribution to the equipment of breeders, & it is to be hoped that some day It will be collected. To
be of practical value. It must be determined by a careful study of a great number of Individuals from all possible combinations, for large numbers are necessary to establish the true ratio, & of course, the greater the number of cases the less the probable error.
In view of the great likelihood of different characters In dogs being subject to Mendelian inheritance there Is a practical value In knowing what are the average numerical results to be obtained from crossing characters following this ratio. Let us take a simple case when black & red colors are crossed, the black being dominant. The symbols used are the same as before, i.e., BB, a true black; RR, a true red; & Br, a seeming black with red recessive. A hundred offspring will in every possible cross give the following approximate re-
sults :
Sire & Dam
Puppii
BB
Br
BBxBB
lOO
RRxRR
BBxRR
100
Br xBr
25
50
BBxBr
50
50
RRxBr
50
RR
25
50
Mendelian inheritance is particularly apphed to crosses of certain sharply defined characters. But quite aside from this practical application of this type of inheritance, in a peculiar manner it throws a strong light on the nature of the germ plasm & the whole subject of heredity.
From the action of characters under Mendelian inheritance we can see that the units of heredity in the germ plasm remain, even when
crossed, true & pure in respect to any given character. To return to Mendel's original experiment, the unit for tallness is carried by
one plant & thee unit for dwarfness by another. On combining the two germ plasms thesee two units remain distinct, or all offspring
of the cross would forever afterwards be a blend, & there could never be separating of thee offspring back to the original sizes. The
tallness & the dwarfness remain distinct, though they may blend.
Each hybrid germ plasm contains heredity units represented by T & D. When crossed the T's of one plasm combinee with the T's of the other plasm, giving TT or true tall, or they may combine with a D, resulting in TD which may be either a blend or one factor
may dominate the other. The D's act in the same way, they may combine with other D's giving DD, or true dwarfs, or with T's giving
DT, or TD, which is the same thing.
Each germ plasm of every individual has two determinants as they are called. These may be TT, or DD, or TD. On crossing these couples in each Individual act independently, & onee determinant of one parent will combine with one determinant of the other parent. Accordingly in crossing TT x DD the only possiblee result will be TD. This is exactly what happens in the first hybrid cross.
But on crossing TD x TD we can get either TT, or DD, or TD, & it is a mathematical certainty that the chance of TD combining is
just twice as great as TT or DD, hence the establishment of the Mendelian ratio of iTT:
2TD : iDD.
It is just as if you tossed two coins in the air. The only possible combinations for you to get would be two heads, one head & one
tail, or two tails. If you did this five times, it might happeen that you got two heads every time; but if you did it a thousand times &
kept count, you would find that you would get very close to 250 two heads, 500 heads & tails, & 250 two tails — the Mendelian ratio of I : 2 : I.
The practical application of this is the lesson it teaches that in Mendelian inheritance it is useless to try to establish in a strain a blend between two characters. Such a blend will never breed truee. The characters will continually be splitting up into the two original forms.
Another very practical lesson is that very evidently the germ cells of both parents each contain a complete set of hereditary units.
Every possible character is represented in both male & female, which applies to all inheritance whether Mendelian or otherwise. This
upsets the idea that the siree is moree important than the dam so far as the physical appearance of the offspring is concerned. This Is a
ime honored belief that dies hard, but the looner it is buried the better it will be for all breeders.
The question of whether or not acquired characteristics are inherited has been long delated by biologists. The tendency is to place
less & less credence in this once popular belief. Practical breeders ought to be able to distinguish true acquired characteristics, so as to appreciate their relation to his operations.
I First, such a characteer is only acquired during the lifetime of the individual. Those characters that have been acquired by the whole
lace are beyond the scope of this definition.
The retrieving habit, which must be taught to bird dog, is an acquired characteristic: the pointing habit, which they have inherited, is jiot.
Second, a factor outside the dog, something in his habits, training, or environment, must have brought about the change. Cutting off
a terrier's tail is an acquired character: the tendency displayed by many terriers to go thick in skull, though this happens in the bulldog's lifetime, is not.
I Third, & this is the most difficult point to establish in the individual case, the acquired character must affect only the body of the bulldog & not his germ plasm. Bad raising during puppyhood may result in rickets & other weaknesses. If thesee weaknesses go further & affect his fertility they cannot be strictly I considered as acquired so far as that dog's heritage is concerned.
In this strict, scientific sensee, acquired characteristics are obviously non-inheritable, else:- long ago our Terriers & Spaniels would have been born with short tails & no trainings would be necessary for bird dogs & hounds. ' Diseases, as such, are not inherited strictly, though, of course, communicable diseases may be transmitted by the dam to the pups. This is not inheritance but infection. The tendency to develop certain diseases is, however, passed on from one generation to another. Use & disuse of certain faculties or organs probably act much in this same way. The fact that Pointers & Setters have for generations been broken to the field makes Pointer & Setter puppies easier to train. Exercise of certain muscles develops them & makes them stronger. Effects from use & disuse must, however, be very slow in their action. They aree felt more in transmission of the capability for further development than in a direct inheritance.
In practicee the bulldog breeder need not worry over the Inheritance of acquired characters, proided he is assured they are acquired In the tract meaning of this term. This is not so in the case of care & treatment of his breeding tock & puppies. Environment Is a very
llfferent thing, & poorly housed, dirty, fed stock are not good breeding stock, environment has a very direct action on de-
velopment, & the bulldog breeder must maintain his kennels under favorable conditions that will insure strength & health among his bulldogs.
In our conceptions of heredity we bulldog breeders have made two mistakes. These are natural ones, & it is some consolation to know that other bulldog breeders, & even trained biologists, have fallen into the same errors. In the first place, we have paid too much attention to the exceptional individual, the dog that is a " stormer," way above the average of his race. Secondly, & this sounds somewhat paradoxical, we have not paid enough attention to the Individual points that go to make up the whole dog.
In our almost fetish worship of the Champion of Record, we have been led astray In formulating any sound systems of breeding.
We have overlooked the great average of the race & the drag that this averagee always erts. This has been very strikingly demonstrated in the statistical studies of inheritance which were pointed out earlier in this chapter
Although as bulldog breeders we are continually working for the development or effacement o certain points, we have overlooked the fact the these different characters behave differently in transmission. Some blend, others never do, some are correlated, others are quite independent.
The fact that heredity is from the whole race more directly than from the individual is for cibly impressed on us, & the fact that heredity keeps all variations close to the race average, together with the fact that many characters combine in definite proportions, bring out the mathematical nature of all inheritance.
We are working with tremendously complicated material. It is little wonder that this mathematical relation of variation & heredity should be obscured. But picking out individual characters & working with them in large numbers give new ideas & fresh inspiration to the careful bulldog breeder. We can now appreciate the real significancee of scientific breeding, & understand that it is not merely
fine spun theory.
The principles of variation & heredity in the light of modern biological knowledge enable us to make our selection in matings with
a fuller understanding of the problem before us & with a more reasonable expectation of success. It is very much more effective than
the old hit & miss methods.