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THE SCIENCE OF ANIMAL BREEDING IN

BRITAIN: A SHORT HISTORY


This essay is a simplified out comprehensive account of British
work in thc field of animal breeding from the earliest attempts, by
farrners, to the most recent scientifically controlled experiments.
DI'. F. H. A. Marshall, Fellow and formerly Vice-Master of
Christ's College, Cambridge, is a leading British expert on Agri-
cultural Physiology, and no Icss well known as an authority OIl
sexual physiology and the physiology of reproduction. From Ig05
lo Ig08 he was Lecturer on the Physiology of Reproduction at
Eclinburgh Univcrsity, and from 1908 te 1919 Lecturer on
Agricultural Physiology at Cambridge, where he bccarne Rcader,
which post he he Id until 19+3. From 1930 to 193+ he was Director
of the Animal Nutrition Instituto at Cambridge. From 1933 to
1935 he scrved on the Council of the Royal Society and was
awarded the Royal modal in 19+0. In 1927 he was sent to AIgeria
by the Ministry of Agriculture on a delcgation to report on Dr.
Voronoff''s gland-grafting experimenrs,
Dr. John Hammond is Reader in Agricultural Physiology at
Cambridge University. He is an authority on cattle breeding, and
the author of many papers on fertility, growth, milk and meat
production.
Sciencc m Britain
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BRlTI SH COUNCIL' S CODE N."'-'''IE : ANIMALS (ENGLISH)
All rights reserued
Prwed /" Grcat Britain b)' The S lo" /IlJpe Preu LId .( 0 Stop/u Pren t OlllPOflY ) . Roehester, Kent
Top leJt: ]. ass Eioart, dJ51-1933
Bol/om l ~ r l : Hri/!ialll Bateson; 1861-1926
'To]: right: Walter Heape, 1855-1929
Boitom right: Thomas Bailoui Wond, 186.9-1929
y t: i: "Jo
THE SCIENCE F ANIMAL
BREEDING IN BRITAIN
A Short History
BY
F. H. A. MARSHALL
C.B.E., Se.D.) LL.D.) F.R.S.

jOHN HAMMOND
M.A.) D.Se. F.R.S.
ILLUSTRATED
Published for
THE BRITISH COUNCIL
BY LONGMANS, GREEN AND CO.
LONDON NEW YORK TORNT

CONTENTS
. /
lFrontispiece
Fa cing page 6
" " 7
I NTRODUCTORY
Tnz P RE- SClENTlFIC P ER lOD
FRA)i CIS G ALTON
] . COSSAR E WART
\VALTER I-f EAP E
I:-i BATESON
G EN ETICS I N E DI NllURGII-F. A. E. CREW .
ANlMAL PHYSIOLOGY IN C\ MllRlDGE
BIBLIOGRAPHY
11.. 1.. USTRATIONS
J. COSSAR EWART
\ VALTER H EAPE
\ VILLIAM DATESON
THOM:\ S BARLOW "\VOOD
EVOLUTlON IN THE S HORTIJORN BREED
OF H ORNS AND FACE COLOUR IN SREEP
FOET,\ L ATROPI-IY IN PIGS
SHETU.ND-SmRE C ROSSES .
G ROWTH CRANGES IN THE PIG .
"
"
"
"
"
5
5
7
9
13
14
2 2
12
13
20
b
THE SCIENCE OF ANIMAL BREEDING IN
BRITAIN: A SHORT HISTORY
'Breeding is the greatest industry to which Science has never yet bc .:
applied.' These words were spoken to the Zoological Section of the
British Association by William Bateson in his presidenti al address to that
body in 1904. And he added, referring especially to the study of hercdity
and variation, that the practical economic value of such application 'will
be found of extraordinary use'. Bateson's words were endorsed and
extended by Walter Heape in a work on The Breeding Industry published in
1906, and he remarked that the study ofthe physiology ofbrecding would
be of equal importance when applied to such subjects as animal fertility
and the factors controlling it.
It may be wondered how in the absence ofbiological research as applied
to the production of domestic animals Britsh breeders obtained such
remarkable results. For it is undeniable that very great success was
achieved and in all classes of stock. It has been truly said that with the
exception of the Perchern and Belgian horses, the Holstein-Friesian and
Swiss cattle and the Merinos sheep, almost all the important classes of
domestic animals widely spread throughout the temperate zones of the
world are of British origino This is the case also of many othcr animals
which have become separate breeds, sch as the American Trotter, which
is based on the English Hackney.
The explanation of this success is twofold. In the first place the selection
of suitable individual animals for perpetuating their kind is very largely
an art and not a science, and the capacity to practise it depends upon
certain attributes of hand and eye which sorne men in this country have
possessed to a remarkable degree. And, secondly, in spite of what Bateson
2 The Application ofScience lo Animal Breeding
and Heape have said to the contrary, many of the mcthods adoptcd by
the breeder were essentialIy though unconsciously scientific.
THE PRE-SCIENTIFIC PERIOD
The first attempts to bring about improvement in domestic animals were
rnade by prvate individuals without any support from the Government
or from scientific or agricultural societies. Among the most successful of
these was Robert Bakewell, whose work led to the development of the
famous Longhom cattle (1750) and the Leicester sheep (1755). John
Ellman, who brcd the Southdown sheep (1778), and Charles and Robert
Colling, who founded the breed of Shorthorn cattle (1780), supply other
well-known instances. Their methods were those of what Darwin calIed
'artificial selection'; that is to say, they chose thc best animals they could
get for the purposes they had in view and then proceedcd to breed from
them, continuing the process in succeeding generations. In other words,
having decided on what types to aim at, they concentrated on those
herds and flocks whose individuals had the desired characters. This
usually involved clase breeding and even intensive inbreeding, which later
scientific work has shown in certain cases to be harmfuI. Essentially the
same methods were adopted by breeders of horses, and in the case of
the Thoroughbred the performance tests ofthe racecourse became the chief
guide in the selection ofanimals to breed from, and these werc duly recorded.
With cattle also it was soon seen that in order to maintain the irnprove-
ment that had been reached, it was necessary to prevent 'dilution of
blood' in strains that had been distributed about the country and con-
sequently were in danger of being contaminated by crossing with inferior
animals, To avoid this herd-books were instituted, and in these the pedi-
grees of the breeding animals which traced back to the original superior
individuals were all recordcd. The earliest of these herd-books was Coates'
Herd-book for the Shorthom Breed and the first volume was published in 1822.
The institution ofherd-books led to the formation ofbrecd societies which
issued thcrn, and in this way standards of quality were set up and it
became the aim of breeders to attain to these standards in the anirnals
which they bred. Moreover, the process of artificial selection was con-
tinued within the families of animals entered in the herd-books, and so
"
The Pre-scentific Perod 3
atternpts were made, and usually successfuIly, to keep and even to
improve the excellence of type already reached, Again, as cornmercial
requirernents changed, or even in response to the caprices of fashion, the
standards successfully set up also changed as a result of selection within
the strains. Thus, by the method of close inbreeding, the Booths (Thomas,
John and Richard), Thomas Bates and Amos Cruikshank in turn bred
new and different types of Shorthorn cattle,
Breeding to type was much assisted by the show-yards which soon carne
to be formed. The first of these shows was that of the Smithfield Club,
founded in 1798. This did much to develop early maturity to the great
advantage ofthe feeder and the butcher. The Royal Agricultural Society's
Shows were founded for breeding stock and this was followed by numerous
county and other local shows, and at all of these, as at the Srnithfield
Show, prizes, medals or other awards were made for the best animals,
and this practice still continues.
To assist in improvement, production and progeny tests have since been
instituted. The Smithfield Club has kept records of carease tests since
r895, the classes being judged by butchers to meet the changing tastes of
the public for quality in meato In a similar way the National Pig Breeders'
Association has carried out carease tests for bacon and pork. For dairy
cattle, rnilking trials have been held at the London Dairy Show since 1876,
as wcll as the Royal Agricultural Society's Show. SimilarIy, milk yields
were recorded for cows, the Ayrshire andJersey societies being among the
first to do this, and in order to get a uniforrn practice throughout England
and Wales, the Board of Agriculture orgaruzed a regular system; registers
were compiled of cows with consistent high production, butter-fat tests
were made, and other points such as quantity, quality and colour of butter
were recorded.
The 'progeny testing' of sires and darns was subsequently introduced
into the show-yards at the Royal and Dairy Shows and awards given for
the collected progeny of an individual bull; furthermore, registers of merit
for male anirnals which sired cows of outstanding merit have been pub-
lished by various societies. The systcm took origin in the practice of letting
out a young bull or ram to other breeders to ascertain the quality of the
progeny before using thern upon specially selected females.
4 The Application oj Science to Animal Breeding
The publication of flock-books for sheep and stud-books for horses of
different breeds very soon followed the institution of herd-books for cattle,
and the tests adopted have been in general very similar. For racehorses, as
already mentioned, there was the additional performance test of the race-
course, and Allison has remarked that the 'mercantile test' is the safest
guide to the successful breeding of the British thoroughbred. The impor-
tance of progeny tests has also been appreciated by horse-breeders, and
the 'figure-system' of Bruce Lowe and Allison was an attempt to apply
this test in a particular manner. It was claimed that in order to produce
thoroughbred horses of merit it was necessary for the parents to have
not only 'running blood' but also 'sire blood' (which was supposed to
perpetuare the power of getting winners}, and figures were assigned to
the various farnilies denoting, in order of merit, the numbers of winners
within these families. In brief, without a sufficiency of 'sire blood'
in the pedigree of a horse, the chance of its producing winners was
small.
In all the work briefiy dcscribed aboye, highly successfuI though it was,
there existed little, if any, consciously applied science, and this is the
justification of Bateson's remark, quoted at the opening of this account of
the history of scientific breeding. It is now undcniable that the breeding
of livestock was hampercd by prejudice as we11 as ignorance such as could
only be overcome by exact scientific tests and the application of acquired
knowledge.
Even at the beginning ofthe present century the physiology of'reproduc-
tion was still in its infancy, and such observations and experiments as had
been made were most uncorrelated and little regard was paid to them.
Nevertheless, the importance of precise scientific method was envisaged
by a few, and it was these men who star ted to make experiments and to
conduct inquiries on their own with little or no financiaI support. The
influence of Darwin and the other great biologists of the Iatter half of the
nineteenth century upon these men was very apparent, while the general
acceptance of the doctrine of organic evolution and the theory of natural
selection as a factor in the production of new species brought about in all
educated men a transformation of outlook which extended to breeders of
animals.
Francis Galton
5
FRANCIS GALTON
Sir Francis Galton (1822-1911), a cousin ofDarwin, founded in the last
years ofthe nineteenth century the biometrical study ofheredity. Although
he was rnainly interested in the study of human heredity, his law of
ancestral .inheritance-i-that the two immediate parents contributed
between them one-half of the effective heritage, the grand-parents ene-
fourth, and so on-had a large influence on stock-breeding. It was on this
idea that the Breed Societies constructed their extended pedigree system.
It is not without significance that the characters, such as stature, which
Galton investigated were those that also interested the stock-breeder. In
this method of research he was followed at a later date by Karl Pearson,
who, among other things, showed that the intensity of inheritance as
measured mathematically varied in different parts of the body; the
significance of this was not recognized until afterwards, when it was
realized that sorne parts are more susceptible to the nutritional enviren-
ment than are others.
J. C. EWART
Among other matters of both scientific and practical importance and
upon which animal breeders were apt to maintain views since proved to
be partIy or entire1y wrong, were the questions relating to inbreeding,
prepotency, the inheritance of acquired characters, telegony (and, in its
cumulative form, saturation), and maternal impressions. One of the first
to undertake extensive work upon these matters was Cossar Ewart (1851-
1933), Regius Professor ofNatural History in the University ofEdinburgh.
About the year 1895 he acquired land at Penycuik, Midlothian, and here
he established a private experimental farm for animal breeding and began
a long series of investigations in Genetics, in which he was a pioneer.
Since the time of Darwin there was no ene, at any rate in Britain, who
had devoted himself to this kind of work, and it was both arduous and
expensve and requiring much patience and foresight. Sorne of the more
important work was done before the rediscovery of Mendel's Iaws of
heredity, referred to below, but by adopting such methods as were known
to him, Ewart successfully conducted a number of investigations upon
prepotency, cross-breeding, inbreeding and reversin, as well as upon
6 The Applicaiion 01Science to Animal Breeding
tciegony and similar real or supposed phenomena among horses of various
brceds and other domestic animals, He fully realized the potential
economic importance of such work, and was not slow to ernphasze the
desirability of applying scientific methods to the study of reproduction
in thc interests of the breeding industry.
Ewart's paper on 'A Critical Period in the Development of the Horse'
(1897), in which he showed the reason why mares tend to slip foal at or
about the sixth or seventh week of pregnancy (when the type of placenta-
tion or mode of attachment of the embryo to the mother is changing from
the primitive yolk-sac or marsupial type to the aIlantoic type which
characterizes most of the mammalia), and pointcd out the need for extra
vigilance on the part of the stud groom at this time, supplies an example
of a physiological discovery of practical significance, and Allison, the
Special Commissioner of the Sportsman newspaper, emphasized i15 impor-
tance in the pages of that journal.
Ewart's most famous experimenta were probably those relating to tele-
gony or the theory, widely entertained both by men of sciencc and by
practical breeders, that a previous sire may so ' infect' the dam served by
him as to impress certain of his characters upon her subsequent ofspring
by other sires. The theory of saturation was an extension of that of
telegony, and postulated that a dam by being servcd over a number of
years by a particular male had his characters so impressed upon her
that her offspring after successive pregnancies became more and more to
resemble the sire. Darwin himself appeared to accept the theory of tele-
gony, and Allison, in his book on The British Thoroughbred Horse, stated
that the truth of telegony was established fact and that it only rcmained
for scientific men to supply the physiological explanation. There can be
no doubt that nearly all breeders took the same view, which is even now
stiIl held by sorne.
The classical case ofthe supposed phenomenon, which is cited at length
by Darwin, was that of Lord Mortori's Arab mareo This m a r ~ , after first
being served by a quagga and producing a striped hybrid foal, afterwards
gave birth to an Arab foal as a result of mating with Sir Gore Ouseley's
Arab staIlion. This foal, which is figured by Darwin, had striped markings
that were said to resemble those of the quagga with which the mare had
EVOLUTlON IN THE SHORTHORN BREED
T op : Cow of 1840 by Reforma , Dam b)' Raby, Grandam b)' Sir Oliva , Centre: Dniry
Shorthorn Coto of 1938, Kn ells Elliot Fernleof 2/1d. Bottom: Beef Shorthorn COIV rif
1939, Scotston Laueuder Lass
r--- - - - ---, - .- - _....
-- --------,1

'., .
_..
' I!
!
< 1

lnlm ilancc l!f Horns and Face Colour in Sheet) fi om IVood, 19u9 )
IVlim a /JII ll n! black-faced bieed 1 j is a ossed unth 1I hornedu.hite-faced breed
( 2) spcckled-faced horned males (3' and polled fe male: (1) are produced.
When the latter are bred together, among other iohite-faced polled (G}
nnd blark-horned (7j ani mals are produced
J. C. Ewart
7
first mated. Ewart repeated the experiment, but the quagga as a species
having become extinct, he employed a Burchell's zebra stallion, and this
served mares of several different brceds, and got striped hybrid foals.
The mares were afterwards put to stallions of their own breed, but the
resulting 'subsequent foals' never showed any sign of having been affected
by the pr vious zebra sire, Ewart carried out other telegony experiments
with various species of animals and they were all uniformly negative as
regards any evidence of the occurrence of the phenornenon. Accounts of
this work, together with the results of other breeding experiments, are
given by Ewart in his book on The Penycuik Experiments (1899), in his
presidential address to the Zoology Section of the British Association in
1903, and in numerous papers contributed to scientific and agricultural
journals.
In 1913 the University of Edinburgh rented a farm at Fairslacks,
Midlothian, and here Ewart kept a considerable number of sheep of
different breeds and started experiments designed to improve the fleece,
This work brought him into contact with the woollen industries of Great
Britain and they realized the importance of the work, In 1923 he went
to Australia at the invitation of the New South Wales Govcrnment and
visited many important and some very outIying sheep stations. He then
proceeded to New Zealand and conducted similar work there. There can
be no doubt that these investigations lcd to the further recognition of
the economic importance of scientific breeding, In 1924 the Worshipful
Company of Woolmen in London struck a gold medal, which they
presented to Ewart in recognition of what he had done.
WALTER HEAPE
In the meanwhile Walter Heape (1855-1929), whose name is mentioned
at the beginning of this s h o ~ t history, had formulated and conducted
a scheme of inquiry into certain other matters connectcd with sheep,
and more particularly those dealing with feeundity. The inquiry was
conducted under the auspices of the Evolution Committcc of the Royal
Society, ofwhich committee Heape was an original member (1896). The
investigation was carried out with the co-operation of the Royal Agri-
cultural Society, and a full report was published in 1899. It dealt chiefly
8 Tlu Application 01Science to Animal Breeding
with the incidence of abortion and barrenness in the different English
breeds, and the causes were shown to be different, depending both upon
breeding and feeding, and to vary with local conditions. The general
fertility of the breeds was also compared and many useful suggestions of
genetical value to farrners were put for ward. At a later date a somewhat
similar inquiry was carricd out in Scotland by the Highland and Agri-
cultural Society (MarshalI, 1908). Among thc practical conclusions which
resulted from these studies was one on the effects of 'flushing' shcep ; that
is, supplying thern with extra corn, cake or turnips or putting them upon
superior pasture or a good new ley at the approach of the tupping or
breeding season. Sheep brcedcrs had already found that these practices
generally increased the fecundity of the ewes, but the matter was not
clinched until it was shown that under the conditions of flushing the egg-
containing follicles in the ovaries ripened more rapidly, and that at the
periods of oestrus or heat two or more eggs would often be discharged
rather than one, the final result being an increase in the number of lambs
born. Another matter of interest concerned the inheritance of fertility.
The evidence collected showed that rarns whi ch were born as twins and
not singles could transmit the fertility of their dams to the next generation
of ewes, and consequently that the practice ofbreeding from singles rather
than twins was one to be deprecated, notwithstanding that the rams which
were single lambs were often better developed owing to their having
obtained more nourishment from their mothers in the early days of their
lives,
Heape also published an important papel' on artificial insemination
(1897). This practicc, although there is evidence that it may have been
known to the Arabs, was first elearly demonstrated by thc Italian biologist
Spallanzani (1784) in the successful insemination of a bitch. But it was
not takcn much notice of until Heape called attention to it by publishing
the experiments by Millais, in which by this method he effected crosses
between Blood-hounds and Basset-hounds (two breeds differing consider-
ably in size). Heape al so emphasized its probable importance to horse-
breeders, qu oting the reports of the Royal Commission on horse-breeding
in which it was shown that the percentage of sterility in thoroughbreds
in any one year might be as high as 40 per cent. One of the earliest
Walter Heape
9
successes in inseminating horses was when the thoroughbred mare Sandi-
way was successfully impregri ated by injecting semen obtained frorn the
stallion Trenton and afterwards produced a first-class foal named Sand-
flake. The more recent deveIopments and applications adopted in the
practice of artificial insemination are referred to below.
Heape's best known and probably his most important work was his
memoir on The Sexual Season oj Mammals (1900), in which he gave a
comparative account of the oestruaI or breeding cycles in all the different
animals for which any data existed. This was soon followed by fuller,
more detailed accounts by others on the oestrual cycles of the sheep, the
dog, the ferret, and the rabbit, and in recent years similar studies have
been made for a large number of other animals.
Heape's book on The Breeding Industry has been mentioned aboye in the
opening paragraph. In this book, which contains much statistical informa-
tion, he stressed the great economic importance of animal breeding in
Great Britain and the heavy losses which were annually incurred through
failure to apply scientific methods to animal production for commercial
purposes. In particular he stressed the importance of breeders keeping
records so that their experiences should not be lost but be made available
to the community.
GENETICS IN CAMBRIDGE-WILLIAM BATESON
Another Cambridge man who played an important part in promoting
and taking part in the study ofbreeding was Wilam Bateson (1861- 1926),
whose first book entit1ed Materials for the Study oj Variation, by its insistence
upon the phenomenon of a discontinuity in organisms, gave a fresh orienta-
tion to the way in which problems of reproduction and evolution were
regarded. Then in 1910, with the disinterment ofGregor Mendel's Versuche
ber Pflaneen-Hybriden (originally published in 1865), there appeared the
clue for which Bateson had been seeking, and he and his co-workers in
Cambridge made much of it in' working out the inheritance of many
kinds of animals and plants. In the meantime, Bateson brought out in
Cambridge an English edition of Mendel's work in the book entitled
Mendel' s Principies qf Heredity (lg0g).
The original experimenta of Mendel, as is well known, were upon

10 The Application 01Science lo Animal Breeding


hybridization in peas, the two parent varieties initially seIected differing
from onc another in ane particular character. The hybrids produced in
cros sing were all similar superficially and resembled one of the parents
in the character in question, which was therefore called the dominant
character, the other character being known as recessioe. When the hybrids
were crossed among thernse lves, approxirnately one-half of the offspring
were found te> be identical with the parent hybrids (dominant hybrids),
one-quarter also superficially resernbled one of the original varieties (the
graudparent with the dominant character), while the remaining quarter
were like the other pure varicty (the grandparent with the recessive
character). Thus the pure dorninants and the dominant hybrids resembled
one another ourwardly, bu! they differed in their capacity to transmit
thc charar teri stic in question, since the pure dominants alone invariably
bree! true. The reccssives also always bred true. Mendel drcw the con-
clusion that in the hybrid the gametes (both sperrnatozoa and ova) were
of two kinds in respect of the characters they carried, or gave rise to,
and they were respectively identical with the two kinds represented by
the garneres of the original pure varieties. The differentiation of gametes
carrying diflercnt characters is the essential principIe in Mendel's theory,
the existence of dorninant and recessive characters, though often obser-
vable, being by no means universal.
Thc Mendelian coneeption of gametic differentiation is the principIe
underlying all thc later work of Bateson and his pupils. Among the earlier
investigations in Cambridge was th at of himself and R. C. Punnett, who
afterwards succeeded him, upon the blue AndaJusian fowJ. Keepers
of ths breed had always recognised the practical impossibility of obtain-
ing a pure strain. However carefully the birds were sclected they
invariably prcduced two sorts of 'wasters', sorne being pure bIack and
sorne white with irregular black marks or splashes. Bateson and Punnctt
wcre the first to supply the explanation. They found that in breeding from
a large number 01 blue Andalusan fowls, on an average half of the off-
spring were blue like the parents, a quarter werc black and a quarter
were splashed-white. They conscquently drew the conc1usion that the
mechanism of inheritance in the Andalusian fowl was comparable to what
Mendel supposed to exist in his hybrid peas. The gametes of the brced,
Genetics in Cambridge-s-William Bateson 11
instead ofbeing a1l similar and carrying the blue character (as one would
have supposed on Weismann's or any of the oldcr theories of heredity)
were of two differcnt kinds, those of the one kind carrying the black
character and those of the other being bearers of the splashed-white.
Such gametes, uniting by chance when the fowls mated together, gave
rise to three kinds of offspring, (me black-white (becoming blue actually
like the parents), one black-black, and one white-white, these appearing
(on the average) in the proportion of 2 : 1 : 1, according to the law of
probability. In this particular case of Mendelian inheritance, neither of
the two altcrnative parent characters (black and splashed-white) was
dominant and neither recessive.
It will be seen from the aboye that the Mendelian conception of
dominance gives a precise significance to the older iclea of prepotency
on which Ewart had been working in Eclinburgh. One of the earliest
workers to apply the Mendelian laws to farrn animals was]ames Wil son of
Dublin, who used them to explain the evolution of thc British breeds of cattle
and to elucidate the problem of 'bulldog' calves in the Dexter breed of
cattle, a matter which was later investigated indetail by CrewatEdinburgh.
Among other experiments carried on by the Cambridge School of
Genetics were those of T. B. Wood on the inhcritance of horns and face
eolour in sheep (Igog). By crossing Suffolk sheep with Dorset Horns,
Wood was able to superirnpose the complete hornlessness of the former
breed upon the white faces of the latter, but in sorne of the hornless sheep
the transmission was not 'purc' since certain individuals grew very short
horns or scurs, The experimcnts as a whole were, however, further evi-
dence that the principies of Mcndelian inheritance could be applied to
domestic animals.
Furthermore, in Iater experiments in which Merino rarns were crossed
with Shropshire ewes it was shown that sorne of the more important
characteristics of the body 01' carease (that is to say, the 'rnutton points')
rmght be transmitted to the third generation so as to appear in new
combinaticns in the cross-bred sheep. Thus, taking the four points, ' over
the shoulder', 'behind the shoulder', 'top of leg', and 'loin', which are
widely different in Merinos and Shropshires (being 'bad' in the former
and 'good' in the latter in regard to mutton production), segregation
12 The Application ofScience to Animal Breeding
appeared very clearly in animals of both the second and third generation,
the points being rcproduced in all possible combinations among the cross-
bred sheep. This case of apparent Mende1ian transmission was all the
more rernarkable in that the characters were not superficial, but deep-
seated and re1atcd to bodily conforrnation, each of them depending on a
numbcr of anatornical factors. It could not be said, however, that all the
characters were inherited 'pur', and in some animals they were definite1y
composite, being different frorn those of both the parent breeds (K. J. J.
Mackenzie and F. H. A. Marshall, 1917).
" Throughout the period there was no proper accommodation for keeping
animals [01' experiments in Genetics, and Ior the purposes of the investiga-
tions just referred to the premises used were Bateson's private garden
at Grantchester, the University Farrn, and Wood's private farm in
Norfolk.
In 1g08 the University of Cambridge created for Bateson a chair in
Biology, but he he1d this only until the end of IgOg, for in 1910 he moved
to Mcrton, Surrey, in arder to take up the directorship of the John Innes
Horticultural Institution, where his work on Mendelian inheritance
related exclusively to plants. Thc animal work at Cambridge was, how-
evcr, carried on by R. C. Punnett, who succeeded Bateson in thc chair
ofBiology. In 1912 a new chair in Genetics in Cambridge was insttuted
and to this Punnett was appointed, arrd a School ofGenetics with Govern-
ment support was founded with adequate buldings and accommodation
[al' thc srnaller animals, and much valuable work on poultry and rabbits
has been "done, notably the work on the inheritance of body sizc carried
out in conjunction with P. G. Bailey.
Together with M. Pease, Punnett produced a sex-linked breed of
poultry which would breed true-the Cambar, the cocks of which at
hatching had a much paler clown colour than the pullets. This was made
by combining the barred pattern of the Plymouth Rock with that of the
Campine. From this breed many new sex-linked varieties of the common
breeds of poultry are now being produced. This latter work was carried
out by Pease at the Animal Research Station-the fie1d station of the
Animal Nutrition Institute ofthe School of Agriculture. The ficld station,
consisting of laboratories, animal houses and sorne eighteen acres of land
Weight-gr-s.
/95
Foctal ,ltrojd])' in Pigs ( (rom Hammond, r.940)
The foeiusesJiom a rol!' sholcillg their ]eloiioe positiou
in the ulcrus (Inri thc weighl of each f oetus. Al thoug]
pregll(Ul(Y is not f m aduanced, uf the foe iuses
halle degtlleraler/ and oihers me about 11) die
Shetland-Shire Crosses ifrom [,jal/U/l & Ha mmond, 1938)
Thc maternal effcct on the sire of the foa! in reciproca! crosses betuieen the large Shire liorse ami small Shetlaud
pon)'." all to the samc sra/e. Top line-s-Parents: Shire stallionX Shetland mate, and Slnre mure X Shetlaiul
stallion. Middle line: Their respectioefoals al birth. Bouom tine: Thefo als al one montlt old
Genetics in Cambridge- William Bateson 13
situated on the University Farm, was built with the assistance of the
Empire Marketing Board in 1934. Pease, in conjunction with the Poultry
Rcscarch Station at Reaseheath, Cheshire, also developed inbred lines of
White Leghorns, which are proving themselves useful for mating to other
strains to 'produce good laying birds.
G E N E TIC s 1 N E D 1 N B U R G H - F. A. E. e R E W
In thc meanwhile, great progress was made in developing the scientific
study of animal breeding in Scotland. Shortly before the war of 1914-18,
and chicfly through Ewart's influence, the Board of Agriculture for
Scotland appointed a special committee to deal with this matter. After
being for some years in abeyance, this committee in 1919 beca me active
and appointed F. A. E. Crew director of research, and this position he
retained until 1944-, but he was necessarily absent on national service from
1939 onwards. As a result ofliberal benefactions, the scheme was greatly
enlarged, and a new and fiourishing Department of Genetics with a
considerable staff ofworkers was formed in the University of Edinburgh.
A chair was founded in 1928 and Crew became the first Buchanan Pro-
fessor of Animal Genctics, only resigning to take up a new professorship
in the university in 194+ During the period of his directorship Crew
published several books (Animal Genetics, The Genetics oJSexuality, etc.) and
he and his staff bctwcen thcm produced a very large number of papers
dealing with experimental work on animal breeding. Among these may
be mentioned the work of Finlay, who organized a Cattle Breeding Con-
fercnce at Edinburgh in 1924, of Roberts on the inheritance of colour
and other characteristics in sheep, of Calder on the coefficient of in-
breeding in herd-books, and ofBuchanan Smith and Donald on studies of
herd-books. A. Greenwood, who started work at Edinburgh on stuclies of
the physiology of development in the fowl, began in 1931 a genetic analysis
of the factors affecting egg production in a strain of Brown Leghorns. He
separated out, by selection and inbreccling, lines which showed different
characters, such as intensity of production, large and small egg size, large
and small body wcight, etc.-work preliminary to combining the desirable
characters in one strain.
14 The Application 01Science to Animal Breeding
Research students from various universities throughout the world took
up temporary residence in Edinburgh and worked in the Genetics depart-
ment, availing thernselves of the new buildings and accommodation at
Nether Liberton on the outskirts of Edinburgh, and these contributed to
the output of work, which grew in volume until about the time of the
out break of war. The Imperial Bureau of Animal Genetics was established
at Edinburgh in 1932 to collect and disseminate the knowl edge on Animal
Breeding; the late Deputy Director, ]. E. Niehols , has also published a
book on Lioestock Improoement, which describes the relation of heredity to
environment.
ANIMAL PHYSIOLOGY IN CAMBRIDGE
The work done in Cambridge, described or alluded to aboye, was
almost exclusively carried out in conneetion with the Zoological Depart-
ment of the University or, as has been shown, in the newly created Sehool
of Geneties; it deaIt mainly with the problems of heredity and variation,
the study of whieh had becomc a new braneh of biologieal science. But
in 1908 a series ofinvestigations was started in conneetion with the rapidly
expanding Sehool of Agriculture in Cambridge, and it remains to give
sorne account ofthe historyof this movement which was more particularly
concerned with physiological research into the functions of the repro-
ductive organs and its practical application to the control of animal
breeding. Heape had left Cambridge before 1908, but his influence still
persisted, and the inception of the scheme for research on the functions of
the generative organs and the fertility of the domestic animals and other
cognate problerns must be traced to his writings.
In 1 gog Professor J. N. Langley, realizing that ordinary laboratory
accornmodation was quite insufficient, formed a committee, of which he
became chairman, to deal with the acquirement of land and buildings
which could be used for researches on the animals of the farm. On this
committee the Schools of Pathology, Parasitology, Biochernistry, and
Agriculture, besides Physiology, were represented, and it was agreed that
these departments should act jointly in the matter. As a result land was
eventually acquired and contributions from each of the departments
agrced upon, and in this way the Field Laboratories of the University of
Animal Physiologs in Cambridge 15
Cambridge were started. In 1919, after the terminati on of the German
war, therc was foundcd in association with thc School of Agriculture an
Institu te of Animal Nutrition, and thi s contained a sub-department to
deal wi th reproduction and allied subje cts in farm animals, and a director
of thi s sub-depa r trncnt (F. H. A. Ma rshall ) was ap pointed. The agri-
cult ural section of the Field Laboratori es then became sepa ra ted offfrom
the other sections, but the work on the ph ysiology of animals was still
continued for sorne yea rs at the old headqu arters on thc road leading out
of Cambridge towards El y. In rgQ3, with the formati on ofa new University
Department of Animal Pathology, the whole of thi s land carne to form
par t of the new department, which had its own professor ; and those who
continued to work there were exclusively occupied with the study of
animal diseases. In the mcantirne, the work on physiology was tr ansfer red
to Huntingdon Road in proximity to the Universi ty Farrn , and it is still
being continued in enlarged premises, These deveIopments occurred in
response to an increasing realizaticn of the practical importance of such
studies to the farmi ng community and to the country and ernpire. The
Empire Marketing Board gave the gr eater par t of the finan cial support,
while the Rockefeller Trust, by thei r generosity in providi ng endowment s
for teaching, assisted indirectly by r elieving the demands upon oth er
funds which were drawn upo n t o suppor t research. The Development
Commission and the Ministry of Agri culture provided the working expenses
(and later, aftcr its formation) with guidance from the Agricultura!
Rcsearch Coun cil. In the meantime, thc original buil dings of the School
of Agriculture were more than dou bled in size.
Mu ch work on physiology was also done in the buildin gs of the School
of AgricuIture itself where smaller animals could be kept Ior purposes of
experiment, whil e sorne of the animals on the University Farrn werc
utilized from the first. Moreover, in conneeti on with the sarne work, visits
were paid to other farms for the pur pose of animal study as wcll as to
bacon factories at Cal ne and Du nmow, where special investigations were
carried out in association with the work in Camb ridge.
The earlier work on thc ovary as an organ of internal secret ion and its
infiuence on the periodi city of the oestrual al' sexual cycle was carried out
in Edinburgh, and evidence was adduced that the sexual glands controlled
16 The Application 01Science to Animal Breeding
the changes which occurred in the other organs of the body in association
with the processes of breedng. It had formerly bcen supposed that some
of the se changes might take place in the absence of the essential organs
of reproduction, but this was definitely found to be crroneous. Several
workers confirmcd the hypothesis ofBorn and Fraenkel (in Germany) that
a structure called the corpus Iuteum, which is formed in the ovary after the
rupture of the follicle and the discharge of the egg, played an important
part in prcgnancy and was a factor in the development of the udder. This
work was extended in Cambridge, and complete studies of the physiology
of the oestrual cyc1e and pregnancy in various animals were carried out
there (see Marshall, 1922; Marshall and Halnan, 1935; Harnmond, 19'27).
Among other matters of interest it was conclusively proved that by
spaying (ovariotomy), the removal of the ovaries 'Nas sufficient to prevent
the recurrence of heat 01' cestrus, the extirpation of the uterus 01' ' bed'
being unnecessary, and that spayed sows tended to fatten faster and better
than 'open' 01' unspayed ones. Moreover, the mammary tissue in spayed
pigs did not develop, and consequently the black pigment in the neigh-
bourhood of the teats in black 01' black and white breeds 01' pigs was not
forrned, since it is normalIy confined to the mammary tissue. In this way
the condition of the belly-piece known as 'seedy-cut', which is due to
such pigment, could be avoided, 'seedy-cut' being deprccated by bacon
manufacturers as it disfigures the meato K. J. J. Mackenzie was one of
those who took a leading part in these investigations,
Another of the earliest studies in Cambridge was one on fatness as the
cause of sterility in domestic animals. It was known that very fat animals
did not come in season in a marked way and that their heat periods were
often irregular, and that the animals might even fail to breed altogether.
Nevcrtheless, the value offat animals was such that it induced the farrner
to fatten them to so great an extent that they might be rendered ternpor-
arily 01' even permanently sterile. The Cambridge inquiries dealt with
the physiological condition of the breeding organs of such. animals and
revealed the reasons why they were unabl'e to brecd, thus confirming
the experiences of those practical farmers who deprccated the practice of
undue fattening. In general, it was shown that whcrcas the supply of good
food raises the nutrition and increases the bodily vigour and fertility, over-
Animal Physiology in Cambridge r 7
feeding and laek of exercise lead to exeessive storage of fat and reduction
of vigour and fertility (see Marshall and Hammond, 194-5).
An investigation was also condueted upon low fertility in pgs which
normally have large litters. It was shown that reduced fertility in many of
the pigs was caused by the atrophy of the foetuses during pregnancy and
generally in early prcgnancy (Hammond, 1914). The results "vere corn-
pared with those found in sorne strains of rabbits, in which the foetal
atrophy was found to be a reeessive genetie character (Hammond, 1928).
Among the more irnportant work done during the succeeding decade
was that of Professor T. H. Bissonnette, who carne from U.S.A. and pro-
ceeded to work on the part played by light in bringing animals into a
brecding condition. His experiments showed that ferr ets could be brought
on heat and made to breed by subj eeting them to artifieiallight or ordinary
elcctrieal illumination in the winter, when the anirnals are norrnall y in the
non-breeding condition. His rnethod was to supply the ferrets with the
illumination at the end of the day, thus extending the period of light to a
lcngth comparable to that which they normaUy experience at the coming
of spring. A large body of experimental and observational evidence has
sincc shown that a very considerable number of other vertebrate animals
rcact similarly to light. The practice adopted by the poultry industry of
subjeeting hens to artificial illumination in order to promete egg-laying is
probably an applieation of the same principie. Later experiments showed
that non-luminous ultra-violet irradiation applied to ferrets may be even
more effective than light. .
Further, fundamental investigaton of the factors which control fertiliry,
so important to the whole bre eding industry, was now actively pursucd.
It was shown that the number of eggs ripened by an animal depended on
sorne substanee cireulating in the blood and not on the structure of the
ovary itself, for when one ovary was removed the same number of eggs
were shed as in an animal with two ovaries (Asdell, 1924-). This substance
~ was later found by workers in U.S.A. and Germany to be a secretion from
the anterior pituitary gland, and a substance with the same effect could be
obtained from the blood serum of pregnant mares between the forty-fifth
and ninetieth days ofpregnancy. Experiments still in progress have shown
that these substances can be used, by injection a few days before service, to
18 The Application 01Science lo Animal Breeding
produce twi ns or triplets (according to the amount injected) in beef cattle
(Hammond, Jm., and Bhattacharya, 1944). These substanees are abo
being used to make sheep breed out of season and so breed three erops of
larnbs in two years, as well as to make goa ts kid in the autumn months
and so add to the suppli es of' winter rnilk.
The causes of sterility among pedigrcc stock are many. In ordcr to
investigat e these in thoroughbred horses, an experimental stud was start ed
in 1935 at Newrnarket with the assist ance of the Thor oughbred Brecders' .
Associa tion and the Agri cultural Res ea 'eh Council with funds dcrived
frorn the Bctting Control Board. Mar es which had been sterile for three
ycars werc taken in, studied by F. T. Day, F.R.C.V.S.,. and when got
in foal were returned to their owners, This work had to be discontinued
at the outbreak of war in 1939, but not befar e sever al of th e main causes
of sterility had been determined and methods for che ear ly diagnosis of
pregnancy developed . The Iatter proved most uscful during the war when
supplics of pre gna nt mare ser um hormone were obtained from wild Welsh
mountain ponies.
In 1922 a section of the Institute was devoted to the study of malc
fertility and sterility, The earlier work of Heape (see aboye) on art ificial
insemination was extended particularly with respect to the problern of
preserving the life of the sperm outside the body by control of tempera turc
-the optimum being 4:oF.-and th e respi rat or y proccsses (vValto n,
1930) . Successful long-distance transport of semen wa s achi eved in 1927,
ra bbi t spe rm being sent bet ween Cambridge and Edinburgh by post.
La ter , when methods developed in U.S.S.R. for collecting semen from
the larger farm animals became available, long-distance transport of r am
semen to Poland (Walto n and Prawochenski, 1936) and bull semen to
Holland (Edwards, Walton and Siebenga, 1937) was achi eved in co-
opera tion with scientists in thosc countries. Ano ther aspect of this work
was in its applica tion to the mas s production of th e best genetic typ es
among farm animals, parti cularly among dairy cattle in which there is no
means of judging the genetic value of the bull unt il his first daught ers
come int o milk, at whi ch time he is at least six years old. Calves Irom such
bulls preven for high milk production can now be produced at the rate of
about a thousand a year, in place of the thirty to fifty by natural rnatings.
Animal Physiolog in Cambridge 19
Earlier statistical studies of the wastage among dairy cows, by H. G.
Sanders in Cambridge and by N. C. Wright and others in different arcas,
showed sorne 20 per cent were culled becaus e of low yields, while furthcr
studies by J. Edwards showed how thi s was due lai gely to t he unknown
genet ic qu alities for milk production of the bulls uscd in the industry. The
applica tion of artificial insemination from proven hulls should therefore
have consider able economic advantages.
The first steps towards a similar extension of the reproduct ive powers of
good genetic stock on the female side were begun in 1929 by G. Pincus, a
visiting National Rescarch Council Fellow from Harvard Universit y, who,
fol1owing work initiated by Heap e, cultivated th c fertilized eggs of th e
rabbit outside the body and later successfully transpl anted thcm into
other does, By the use of anterior pituitary hormones (see aboye) he was
able to ob tain up to fifty or more fertilized cggs at one time for this purpose.
These experiments have yet to be extended to farm animals, in which the
possibilities are great.
I n breeding for milk, one of the difficul ties met with in the sclection of
the dairy cow by milk records is in determini ng how far the environrnenta l
conditions affect the yield given. A statistical investigation of the factors
affecting milk yields was published by H . G. Sanders in 1928, and thi s
enabled 'correct ion factors' for age, dry peri od, service period, etc., to be
made. This investigation led to a study of the way in which the ud der
grows to produce milk. Observations on heifers in calf for th e first time
showcd that a sudde n increase in th e growth of the mil k-secr et ing tissues
of the udder occurred at th e twentieth wee k (Hammond, 1927). Lat cr
experirnen ts by Wallace showed that this growth could be augmented
considerably by a hi gh level of feeding in the later stages of pregnancy.
It was found, too, th at the stimulus for growth carne from che rnical sub-
stances--oest rogens- produc ed in th e pl acenta and circul ating in the
blood. Synthetical1y produced chcmical substances wi th an actiou similar
to the ocstrogens were produced by Prof. E. C. Dodds, Middlesex Hospital,
London. J. Hammond, J nr., and F. T. Day, using these substances by the
method of tablet implantations, developed in 1\. S. Parkes' s laboratory at
the National Medical Research Institute, Hampst eacl, wer e abl e to obtain
growth of the udder and milk yields of up to 30 lb . per day witb a normal
20 The Application 01Science to Animal Breeding
lactation period in heifers which had never been pregnant, Experimenta
on similar lines in cattle and goats wer e also conducted at the National
Institute for Rcsearch in Dairying at Reading by S. J. FoIley and his
co-workers. These experiments indicated some of the basic ph ysiologicaI
factors which have to be taken into account when breeding for such a
complicated character as milk production.
The size and vigour of the young at birth playa material part in the
survival rate of all animals and especially of sheep, in which parturition
frequently occurs under exposcd conditions. Studies on the effects of l.
high planc of nutrition in the later stages 01' pregnancy in increasing the
birth weight and vigour ofthe lamb by Hammond (1932), Verges (1939 b)
and Wallace (1944) suggested a remedy for the large losses which occur
at or shortly after lambing time owing to the birth ofweak lambs. Similar
results were also obtained by W. Thomson and A. H. H. Fraser in experi-
ments at the Rowett Research Institutc, Aberdeen.
Early experiments with rabbits had shown that the weight 01' the
individual young decreased as the number in the litter rose, and this
raised the problem of how far the nutritional environment of the mother
could affect the ultimate size and development of the young. Reciprocal
crosses, by means of artificial insernination, were made between the largest
horse (Shirc) and small est pony (Shetland), and it was found that there
was a maternal, in addition to a genetic, influence of the mother not only
on the birth weight (see illustration) but also on the ultimate size which
crosses attained (Walton and Hammond, 1938).
The experirnents made by Mackenzie and MarshaIl ( 1917) on the
inheritance of mutton qualities in crosses between Merino and Shropshrc
sheep had shown the necessity for a more accurate definition of meat
qualities in animals before much progress could be made in brccding for
specific meat qu alities. This led to a study of the growth and deveIopment
of the mutton qualitics in the shcep by P. G. Bailey, a geneticist, unfor-
tunately killed in the war of 1914-18; A. B. AppI eton, a comparative
anatomist from the Cambridge University Department of Anatomy; and
]. Hammond, a physi ologist from the Institute of Animal Nutrition; and
the results, published as a book (Hammond, 1932), present an account
of the scientific principIes which form the basis of meat production in the
Crowlh Changas in the Pig (1 ,0111 Hauunond, '940)
Each animal ir icduced lo the same headsirr. il s IlI/ imbrooed breed like lhe
Mi ddfe Wltite grnws up, the fJfofJOrli oTl 01 loin lo head and nrrl: increases
greaI0'; but an unimprooed ~ ) ' P e JUe/1 as ihe Wild B OIl /" g rolt'J up uiithout
mucli cliange in bod)' proportions, 'Top: Fceiu, of 2 months. 211d fine:
Middle WhiteJ ioeek old, Jj/b . 3 rd tille: Middle IVhite-Ij ioeeks
su. JOO lb. Bottom: Wild Bom- adlllt , alioul300 lb.
Animal Physiology in Cambridge 21
animal. This was followed by analytical studies of the meat qualities of
differcnt breeds of cattle, pigs and sheep, made by visiting research workers
from other countries (Hirzel, 1936; Hammond and Murray, 1937; Plsson,
1939-40), and derived from carease tests at shows and selected animals
from brecders. Later, experiments on the effect of the nutritional leve!
during the growth period in pigs (McMeekan, 1938) and in sheep (Verges ,
1939 a) showed how important a high nutritional level is for the full
expression of the genctic characters of the animal for meat purposes. AH
these results suggested the conclusion that, for meat purposes, thc nutri-
tional environment in which selection of breeding stock 1S made is of thc
highest importance for satisfactory progl'ess. 1t is only under such condi-
tions of high nutrition that the full expression of the genctic characters
can occur and so progress can be made by selection. If the environrnent
is not optimal for the development of the character for which sclcction
is being made, then the development of the character is limited by the
environment and not by the genetic constitution (Hammond, 1940). Thus
man has the power of directing the course of evolution, not only of
his domes tic animals, but also of himself, by the environment, nutritional
and otherwse, which he himself creates for the rearing of thc next
generation.
22
The Application ofScience lo Animal Breeding
BIBLIOGRAPHY
Further details concerning work quoted abooe can be obtained Jrom the Jollowing
publications, listed (excepting Jor the jirst reference under authors' names:
Animal Brccding Abstracts, 1933-47, Imperial Bureau oj Animal Breeding and
Genetics, Edinburgh.
AscleJI , S. A., 1924. Sorne effects of lateral ovariotomy in rabbits, British
Journal oj Experimental Biology, I.
Bateson, 'vV., 18g4. Materials for the Study of Variation, London.
--, 1909. Mendel's Princ ipies oj Heredity, Ca mbridge.
Bissonnette, T . H., 1932. Modifi cations of mammalian sexual cyeles.
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Calder , A ., 1927. The rol e of inbreeding in the Clydesdale breed of hors es.
Proceedings of the Royal Society oj Edinburgh , 47.
Crew, F. A. E. , 1925. Animal Genetics, Edinburgh.
- - , Ig27. The Genetics oj Sexuality in Animals, Cambridge.
Darwin, C., 1859. The Origin oJ Species, London.
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Day, F. T., 1940. Clinical and experimental observations on reproduction in
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Donald, H. P., 1940. Breeding pol icy in relation to performance testing in pigs.
Empire J ournal oJ Experimental AgricuIture, 8.
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--, Walton, A., and Siebenga, F., 1938. On cxehange ofbull semen between
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Ewart, J- C. , 1897. A Cr ical Period in the Deoelopment oj the Horse, London.
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Galton, F., I88 3. Inquiries into Human Faculty and its Deoelopment, London,
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Hammond,j., 1914. On sorne factors controlling fertility in domestic animals.
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- - , [927. Reproduction in the Cow, Cambridge.
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breeds of pigs. J ournal if Agricultural Science, 27.
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McMeckan , C. P., 1940. Growth an d development of thc bacon pig, wit h
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--, 1922. The Physiology of Reproduction (znd edi tion), London.
- - a nd Halnan, E. T. , 1945. Physiology of Farm Animals (3r d edition), Cam-
br idge.
-- an d Hammond,]. , [945. Fertility and animal breeding (6th edi tion),
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Nichols, J . E., [944. Liuestock lmprooemeni, Edinburgh.
Pl sson, H. , [939-40. Meat qu alities in the sheep, wit h special refcrence to
Scottish brecds and erosses. J ournal uf Agricultural Science, 29 and 30.
Pearson, K., 1900. The Grammar of Science, London.
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Pincus, G. , 1930. Observations on the living eggs of the rabbit, Proceedings of
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-- and Bailey, P. G., 1914. On inheri tance of weight in poultry. Journal of
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Sand crs, H. G., 1928. The variation in milk yields caused by season of the
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The Application 01Science to Animal Breeding
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