88 THE AMERICAN SOCIETY OF ANIMAL PRODUCTION GENETICS IN PRACTICAL STOCK B R E E D I N G BY E. G. RITZMAN New Hampshire Agricultural Experiment Station The a r t of animal breeding has gone through a process of slow evolution which is characterized by three distinct stages of advancement. Since the earliest days of history mankind has been t r y i n g to control in some measure or other the transmission of desirable Or undesirable physical characteristics or functions. No other science has been more inviting to our innate curiosity, b u t its history also shows that no other line of investigation has been more baffling. As a consequence a large p a r t of the earlier work was based on general observation supplemented by no small degree of speculative assumptions. The simple f a c t that "like produces like" was one of the first of these general observations and its general t r u i s m was so striking t h a t it has held the stage since the early records of history. Practically all the progress in animal improvement has been due to intelligent application of this principle. As animal f o r m s and functions were improved and artificially bolstered up it soon became a p p a r e n t that the exceptions to the rule were quite common, but it took y e a r s before it became recognized that the g e r m plasm of an animal was not necessarily identical with its somatoplasm. The first really marked scientific advance along this line was b r o u g h t about by the Weisman and Darwin School of investigators in pointing out t h a t the carrier of h e r e d i t a r y qualities, the germ plasm, represented an ancestral composite which did not necessarily conform to the somatic expression of the carriers of that germ plasm. On the basis of this new philosophy was built the first a t t e m p t at scientific analysis and control of h e r e d i t a r y influences. The means by which this was carried on consisted in a mathematical analysis of the probability with which an animal could t r a n s m i t his own physical characteristics b y exact m e a s u r e m e n t of those t r a i t s together with those of his ancestry. Galton's law of ancestral heredity was, therefore, merely a modification of the original rule and, while it improved the method o f analyzing the probable potential t r a n s m i t t i n g THE AMERICAN SOCIETY OF ANIMAL PRODUCTION 89 qualities of an animal, it still left the control of h e r e d i t a r y qualities dependent on a wide margin of chance. Mendel also began essentially as a biometrician, but his unique method of attack b r o u g h t forth the key to the situation by showing that h e r e d i t a r y factors are unit characters capable of combining and segregating in definite proportions. Mendelism m a r k s the first step in the possibility of controlling the hereditary qualities of our domestic animals, but its validity rests on the premise that the parent stock must be pure with regard to the t r a i t under consideration. It is essentially a process of selecting f r o m any source of parentage the characteristics desired to be united in the first cross or F1 generation. This F1 is of course a crossbred c a r r y i n g gameticaUy all the desirable as well as the undesirable traits of the two parents. Its somatic development m a y be desirable or undesirable. In fact, while animal breeders have often resorted to crossing on the ground t h a t the first cross was physically superior, the somatic development of the F1 is scientifically u n i m p o r t a n t since it is highly heterozygous. The important function of the F1 is that when inbred to itself it t u r n s out to be a P a n d o r a ' s box from which all sorts of new combinations issue, some pure in character and some mixed. In our domestic livestock the term purebred is in a sense very misleading. Since most of our pure breeds have been built up f r o m a combination of blood, they are all relatively heterozygous with regard to certain of their t r a i t s especially those relating to conformation. Nevertheless, inbreeding tends to intensify these traits so that the germ plasm will more nearly represent the soma of its host. Now, remarkable as it m a y seem, our improved breeds have unconsciously been produced by Mendelism, the original improvers like Bakewell, Collins, and others having possessed a high degree of intuition in selecting the dominant individuals t h a t issued f r o m the P a n d o r a ' s box. More recent achievements like the famous Corriedale are nothing b u t the result of inbreeding hybrids, since it is a fact t h a t in some of the most noted foundation flocks of this breed in N e w Zealand no outside blood was used for periods involving six to eight generations. While it has always been supposed that such 90 THE AMERICAN SOCIETY OF ANIMAL PRODUCTION intense inbreeding would finally be injurious, all the more noted improvers of livestock seem to have been able to avoid such misfortune by intelligently discarding the potentially harmful inbred individuals. The guiding principle by which they determined the desirable from the undesirable was not given to posterity. It was probably largely unexpressed intuition since they were ignorant of the laws of dominance and recessiveness, but it is more or less certain that the old school of improvers deemed it essential that physical or somatic perfection involved also a superior expression of character, suggestive of strength in the male and feminine gentleness in the female. Now the general history of improved breeds is much the same; usually a mixture of quite unlike blood was used to furnish that combination of traits which was desired, the next step being usually inbreeding to maintain the standard, incidentally, however, intensifying the germinal determiners. Then there followed a general dissemination and demand for such improved stock beyond the available supply, which of course resulted in bringing in much new blood, curtailing selection and, therefore, generally weakening the breed by establishment of a large number of subfamilies developed along different blood lines. Practical breeders have not recognized sufficiently the importance of concentrating along a chosen blood line to intensify t h e germinal determiners notwithstanding the fact that the breeding of outstanding producers repeatedly illustrates this point. Much of the more recent genetic investigations on the breeding of show animals emphasizes the important part that concentration of specific blood lines has played in producing winners. This simply supplies the proof to the progress of breed history in general that prepotent individuals can be produced only by concentration of germinal factors representing similar somatic or functional characteristics. While genetics has thus already accomplished much of practical value in analyzing the causes that have produced successful animals in the past, its chief effort and contribution is along more basic lines of tying up the physical and functional traits of animals with the influences or causes that bring about variation between individuals. Cytological research has 'already THE AMERICAN SOCIETY OF ANIMAL PRODUCTION 91 t h r o w n much light on the p a r t played by the chromosomes in the transmission of h e r e d i t a r y qualities, a number of which, such as color, horns, ear length, fecundity in poultry, etc., have been assigned a definite genetic constitution controlled by presence or absence of a definite number or combination of genetic factors t h a t conform to the law of segregation. While the practical application of genetic factors or rules by which the transmission of somatic t r a i t s can be controlled thus apply for the most p a r t to traits which have no g r e a t economic significance, there is some evidence that functional traits as well as conformation m a y eventually be correlated with t h e i r tangible unit factors or d e t e r m i n e r s when the analysis of the soma is made on the basis of skeletal units, that is, on single bone dimensions. Since conformation as it is now treated in practice is a complex combination of physical traits, its hereditary control can hardly proceed beyond the generalized rule that "like produces like" until the practical stockman is given a more 9specific education on those physical and functional t r a i t s which act independently in a h e r e d i t a r y sense where close correlation in h e r e d i t a r y behavior does not exist. F r o m this point of view the linking up of the men who are doing this fundamental pioneering in genetics with those of us who are daily called upon for advice that essentially demands the practical application of fundamental laws is of historic significance. The contact is stimulating to the geneticist and it is essential to f u r t h e r progress in practical animal breeding. THE NEXT ANNUAL DINNER In keeping with a custom established by the Society an annual dinner and meeting will be held in honor of one of its members, whose achievements in the field of animal industry have been noteworthy. The dinner and meeting for 1923 will be held Sunday, December 2. Only those who have attended on f o r m e r occasions can look f o r w a r d with a full appreciation of this coming event.
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