Techniques of Hybridization of Poultry D O N C.
WARREN
Bureau of Animal Industry, Regional Poultry Breeding Project, Lafayette, Indiana (Received for publication July 11, 1949)
INTRODUCTION
I
It is true that considerable work has been done upon techniques of hybridizing poultry by a few experiment stations, notably Iowa and Minnesota (Waters and Lambert 1936; Shoffner 1948a, 1948b; Wilson 1948), but much of the philosophy guiding procedures must come from corn breeding and to a degree from swine breeding. These are the two fields in which greatest effort has been made toward the solution of such problems. It is in corn breeding that the greatest contributions have been made but there are some obvious limitations in poultry breeding which did not impede the development of hybrid corn. First of all, the difference between self-fertilized plants and bi-parental animals in the rate at which a high coefficient of inbreeding may be attained has a significant influence on procedures. In plants one generation of selfing gives an inbreeding coefficient equivalent to that of three generations of sibbing, which is.the standard procedure
The vast amount of study by plant breeders on methodology of utilization of heterosis should yield information of value to poultry breeders, and provide them with sign posts for expediting prog59
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in animals. This makes a wide difference in the time and cost of development of inbred lines but may have the merit of reducing the impact of inbreeding and making possible the salvaging of lines which might otherwise be lost by too rapid deterioration. Another significant difference between plant and animal breeding is the ease with which controlled matings may be made. In plant breeding hand pollination may become a laborious and expensive procedure which limits its practice, while in animals such controlled matings involve only a choice of males going in the breeding pen and constitute regular commercial practice. Although the tendency has been to attempt duplication of the procedure of the corn breeders, the techniques finally adopted by poultrymen may differ widely. The decisions of the plant breeders regarding their practices have been largely influenced by availability and economy of techniques in their field which may or may not have application in poultry breeding. One advantage in the manipulations of the material of the animal breeder over those of the plant breeder is the availability of proved individuals. The plant breeder dealing with annuals has not proved sires or dams for exploitation. In the maintenance of inbred lines the plant breeder has a great advantage in being able to store his seed for years.
T IS not the purpose of this discussion to elaborate on the merits of hybrid poultry. While there may be grounds for entering upon such a discussion, the major problem facing the poultry industry is to obtain a prompt evaluation of the possibilities of the utilization of hybrid vigor. The exceptional progress made in plant breeding through the utilization of this phenomenon has forced upon the poultry industry the decision regarding the acceptance of hybrid poultry before it is in a position to form a sound opinion.
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ress. It should be kept in mind, however, that differences in materials of plant and animal breeders may demand widely varying procedures for attaining the same end. ORIGIN OF INBRED LINES
It remains a moot question among plant breeders as to the value of testing of
DEVELOPING INBRED LINES Animal breeders have been inclined to favor a slow rate of increase of the inbreeding coefficient combined with high selection pressure for the establishment of inbred lines (Waters and Lambert 1936). This greatly extends the period of development of inbred lines and makes it possible to carry fewer lines with the same facilities. Some poultry geneticists have held that this is the only method by which inbred lines of poultry can be expected to survive. This philosophy prevailed at the initiation of the cooperative swine breeding project ten years ago. Today opinions are being expressed by some of the leaders of the swine breeding program that progress might have been greater if they had accelerated the inbreeding to the highest degree possible and screened lines more rapidly. Following this lead, some poultry
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Since interest in the development of inbred lines of poultry has recently been aroused it may be well to discuss problems related to the establishment of such lines. Of the many inbred lines now in the stocks of agricultural experiment stations too little attention has been paid to the quality of stocks giving rise to them. There are at hand some outstanding commercial strains of poultry and greater utilization of them should be made in initiating inbred lines. Not only should the best performing stocks be used but at least sufficient family testing should be done to locate superior individuals with which to start inbred lines. The history of plant breeding indicates that diversity of genetic origin is an important factor in the success of combining inbred lines (Sprague 1946b), so even a large number of inbred lines of relatively common origin are probably not too promising with regard to combinability. This limitation emphasizes the value of cooperative effort on the part of agricultural experiment stations, thus providing a wide range of stocks as a source of combinations. Experience in corn breeding would indicate that the recent ruling of the Federal Trade Commission that the term "hybrid" may be used for poultry without qualifications only where inbred lines from two or more breeds or varieties are combined will probably not handicap the program since such combinations are the more promising, but provisions may eventually need to be made for a broader interpretation.
stocks for combinability before inbreeding is started (Jenkins 1935, Sprague 1946a). The excessive cost of establishment and maintenance of inbred lines in poultry might justify the use of early testing even though it is of questionable value to corn breeders. If nicking of different breeds and strains of poultry has any prediction value as to the subsequent combinability of inbred lines derived from them, then we already have available to us a considerable body of information on the subject as a result of research and commercial crossbreeding. If testing before inbreeding has prediction value for combinability it would show where to concentrate efforts on inbreeding. The successes, such as Austra-Whites in the Midwest and in the Barred broiler crosses in the East, may be indications as to vantage points for launching programs of inbreeding. First there is need for critical evidence regarding the relationship combinability before and after inbreeding.
HYBRIDIZATION OF POULTRY
There is no information as to the degree of inbreeding necessary for hybridization in poultry. It should be recognized that there is no magic about the practice of inbreeding since it merely tends to produce stocks, the combinations of which give repeatable performances. It also tends to establish lines of diverse genie constitution. Both corn and swine breeders seem now of the opinion that inbreeding can be less than thought necessary a few years ago.* It is true, however, that the success of hybrid corn is based largely upon uni* Personal correspondence.
formity of performance resulting from homozygosity attained by inbreeding. What is needed is a high concentration of specific genes rather than homozygosity for all genes. It is possible that this may be accomplished by techniques other than inbreeding. Therefore, in poultry where inbreeding is so costly other possibilities must not be overlooked. Corn breeders have found that certain weaknesses can be tolerated in inbred stocks (Sprague 1946b). In fact, a number of them cannot be avoided. Some defects in the inbreds are not reflected in their hybrid offspring. If the same situation holds in poultry breeding, then a classification of the economic traits of the fowl in this respect would indicate where greatest selection pressure should be applied when developing inbred lines. Where dominance or epistacy plays an important role in the inheritance of a trait, superiority of expression in the inbred will be of lesser importance. In traits which show the additive type of inheritance, selection pressure will be of considerable value in the development of inbred lines. COMBINING INBRED LINES
With even as few as a hundred inbred lines the identification of inbred line combinations which do well is a tremendous task. In this connection the phenomenon of sex-linkage confronts the poultry breeder though not a problem to the plant breeder. The necessity of making reciprocal crosses virtually doubles the task. Perhaps a proper utilization of the available knowledge of sex-linked factors in poultry may help to reduce the number of reciprocal matings required. The corn breeders have used the cross of inbred lines on open-pollinated stocks for purposes of gaining a preliminary evaluation of available inbred lines (Sprague 1946b). If such a short-cut proves reliable in the produc-
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workers are starting a large number of inbred lines (brother-sister matings) at one time, eliminating most of the lines after one generation of inbreeding, and repeating this program two or three years as means of obtaining promising inbred lines. This procedure is based upon the thinking that any lines which show heritable defects or any lack of vigor at the start are likely to become worse and eventually reach a condition which would render them useless for hybridization purposes. By such early screening it is hoped that effort will be limited to inbred stocks which will have higher survival possibilities under the rigors of inbreeding. Owing to the expense of carrying inbred lines of poultry any wastage which can be prevented by early and rigid screening would seem desirable. Such screening would fit more satisfactorily into a scheme of intense inbreeding. Slower inbreeding with higher selection pressure is more likely to produce viable inbreds but since chance is a factor in finding superior combinations there is the question whether such a program will provide a wide enough source of germ plasm for making combinations. Both the slow and rapid methods of developing inbred lines are being followed and it may be well that this is true.
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OTHER METHODS OF UTILIZATION OF HYBRID VIGOR
There is developing a school of thought that perhaps the conventional methods of production of hybrid corn have resulted in considerable wasted effort (Hull 1945). It is proposed that utilization of heterosis may be accomplished without the usual laborious procedures of inbreeding and testing for combinability. This would involve developing through selection composite stocks with high specific combinability for a previously chosen tester stock. If this method has possibilities in the
poultry breeding program it would avoid the somewhat discouraging picture ahead of those wishing to take advantage of hybrid vigor. Poultry investigators cannot ignore the possibility that there may be available among the many strains of poultry some two which have the genetic constitutions that, when brought together, may have stimulatory effects comparable to those of combinations of inbred lines. If such combinations were available they would greatly reduce the cost of utilization of heterosis. If such combinations were found, and they are more likely to be found in closed flocks, then further selection for improved combinability between the two stocks might be practiced and perhaps the use of a limited degree of inbreeding to improve the uniformity of results. The chances of finding such successful combinations are probably better in poultry than in corn where cross pollination is such a problem.
PROBLEMS
There are many questions regarding hybrid vigor in poultry that are now facing investigators. Several of these have already been mentioned. Others are the manner in which inbred stocks may be maintained when a satisfactory degree of inbreeding has been reached. Can such lines be carried long in mass matings? Can single crosses used in double crosses be carried in mass matings without a deterioration of their stimulatory effects? Probably the greatest need at the present time is some prompt means of obtaining an evaluation of possible techniques. Some rapidly breeding form such as mice or Drosophila offers definite possibilities in obtaining dependable answers more promptly than might be obtained from the slower breeding domestic forms.
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tion of hybrid poultry it will greatly reduce the problem of testing for combinability. It would detect the more promising inbred lines of poultry and then the usual technique of making all possible combinations among a limited number of the better combining inbreds would serve as a basis for prediction as to how the lines will combine in double crosses. There are certain limiting factors, particularly color inheritance, which will have a bearing on the methods of combining inbred lines. It probably would not be desirable to have crosses which throw more than one plumage color unless the resulting hybrid chicks were sold in their respective color groups. Proper use of the dominant white plumage color factor will help to reduce variability in plumage color. The value of dominant white for this purpose emphasizes the need for a heavy variety with this factor for development of inbreds to be combined in producing brown-egg hybrids. A knowledge of the inheritance of morphological characters of chickens will help to indicate the methods of combining inbred lines of the various breeds and varieties for the production of a hybrid which is likely to gain public acceptance.
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N E W S AND N O T E S
Shoffner, R. N., 1948b. Performance of "incrosses" among lines of Leghorns. Poultry Sci. 27: 683. Hull, F. H., 1945. Recurrent selection and specific Sprague, G. F. 1946a. Early testing of inbred lines combining ability in corn. Jour. Amer. Soc. Agron. of corn. Jour. Amer. Soc. Agron. 38: 108-117. 37: 134-145. Sprague, G. F., 1946b. The experimental basis for Jenkins, M.T., 1935. The effects of inbreeding and of hybrid maize. Biol. Rev. 21: 101-120. selection within inbred lines of maize upon hy- Waters, N. F., and W. V. Lambert, 1936. Inbreeding brids made after successive generations of selfing. of the White Leghorn fowl. Iowa Agr. Exper. Sta. Iowa State Col. Jour. Sci. 9:429-450. Res. Bui. 202. Shoffner, R. N., 1948a. The reaction of the fowl to Wilson, W. O., 1948. Egg production rate and fertility in inbred chickens. Poultry Sci. 27:719-726. inbreeding. Poultry Sci. 27: 448-452. REFERENCES
News and Notes (Continued from page 51)
research assistant in the Poultry Department at the University of Massachusetts. NOMINATIONS FOR T O M N E W M A N
PUBDUE NOTES
M E M O R I A L AWARD
Wilson Henderson (B.S. '32, M.S. '41 Univ. of British Columbia; D.V.M. '47 Toronto) was appointed Associate Professor of Veterinary Science and assigned to poultry diagnostic and research projects in September, 1949. He was formerly at West Washington Experimental Station (Puyallup). A. H. Johnson (B.S. Rhode Island State '49) has been appointed Inspector for the Indiana State Egg Board and National Plan Inspector for the State Poultry Association. He began his duties with the Egg Board in July and was transferred to the Association in October, 1949.
Russell H. Hinds, Jr. (B.S. Univ. of Maryland '49) has been appointed Extension Poultry Specialist for southern and western Maryland.
The Council of the Poultry Association of Great Britain has established the T o m Newman Memorial Award, consisting of a handsome medal and sum of money, to be awarded annually to the person making the most important contribution in poultry husbandry research work during the preceding year. Nominations of candidates in the United States for work done in 1949 are invited and should be sent to Dr. Morley A. Jull, University of Maryland, College Park, Md., secretary of the U. S. Consultative Committee. T h e other members of this committee include: Dr. F . R. Beaudette, Rutgers University, New Brunswick, N . J. Dr. L. C. Norris, Cornell University, Ithaca, N . Y., and Dr. R. E. Phillips, Iowa State College, Ames, Iowa.
MASSACHUSETTS N O T E S
J A M E S E. R I C E M E M O R I A L L I B R A R Y
Edgar W. Spear (B.S. University of Massachusetts '49) has been appointed
The Poultry Industry of the United States and Canada is raising $25,000 to
MARYLAND
NOTES
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Poultry Science Association, to the retiring Editor—Thanks for a job well done! H. D . B.