The career of Leon Jacob Cole is a beautiful example of the fortunate matching of an historic opportunity with appropriate talent. The signal contributions of Dr. Cole to the early applications of genetics to animal and plant improvement were well documented 50 years ago by his contemporary peers (H. L. Russell, E. W. Lindstrom and W. C. Coffey, ASAP, 1939) at the presentation of his portrait to the Saddle and Sirloin Gallery by the Society. One of his famous students, Ivar Johansson (1961), later summarized Dr. Cole's accomplishments for readers of the journal Genetics. A brief biography of Dr. Cole also was prepared for the Dictionary of Scientific Biography, Supplement II in 1985 (Chapman, 1989). This effort seeks to remind the present generation of animal breeders and their students of "our roots" from the perspective of two former students of Dr. Cole, who are now themselves retired.

Born in 1877 and reared as a "city boy" in Allegany, NY, before the rediscovery of Mendel's laws, "L. J.'s" early scholarly interests in animals and plants led him to prepare well for his later key contributions to the training of geneticists during the critical embryonic stages of this science, particularly for its applications to improvement of agricultural animals and plants.

After work experience on farms during pre-college summers, he attended Michigan Agricultural College in 1894-95 and 1897-98, and then followed with an A.B. in Biology from the University of Michigan in 1901 and a Ph.D. in Zoology from Harvard in 1906. Cole's abilities were recognized early by his appointment as Assistant in Zoology at Michigan from 1898 to 1902, and as Austin Teaching Fellow at Harvard from 1903 to 1905. He also was chosen as a member of the Harriman Expedition to Alaska in 1899 and of the Yucatan Zoological Expedition in 1904, a summer investigator at the Bermuda Biological Station in 1903, at the Tortugas Station in the Gulf of Mexico in 1906 and at Woods Hole Marine Labs during other summers from 1901 to 1906. These experiences provided invaluable stimulation from contacts with eminent ornithologists, mammalogists and botanists. As a 2~yr-old university student in 1901, Cole first published on the leg-banding of birds as a method for studying their migration. He guided organization of the American Bird Banding Association in 1909 (McCabe, 1979) and is recognized as the father of this endeavor. As a graduate student in 1905, Cole published a monograph for the Bureau of Fisheries on German carp in the United States (Cole, 1905).

With this rich background in animal biology, Cole became the 29-yr-old Chief of the Animal Breeding and Pathology Division at the Rhode Island Experiment Station in 1906. There he began his lifelong marriage to Margaret Belcher Goodenow and his research on the genetics of pigeons, which he continued as Instructor in Zoology at Yale from 1907 to 1910. Then came the unique opportunity for a key role in the early development of the new field of genetics and its applications in agriculture.

In 1910, Dean Russell of the University of Wisconsin, College of Agriculture, realized that the time was ripe to begin serious consideration of potential application of the embryonic science of genetics in the improvement of animals and plants. There was some initial difficulty in convincing the University President and Board of Regents that a Department of Genetics should be located in the College of Agriculture rather than in Letters and Science. But, with support from Regent and Ex-Governor Hoard, approval was obtained for a Department of Experimental Breeding, later changed to Genetics. Establishment of a single new Department of Genetics was judged more likely to stimulate development of genetic applications to both plant and animal improvement than would scattering this work among such departments as animal, diary or poultry husbandry, agronomy, horticulture and entomology. However, this approach would require effective working relations with the separate departments, a challenge which young Dr. Cole proved especially fitted to meet.

The genius of Dr. Cole's contribution to the genetics of plant and animal improvement was the broad scope of his interests and his ability to relate to and develop cooperation with others working with particular species. Dr. Cole's personal research continued to be with genetic traits of pigeons, such as plumage color, sex-linkage, sex-ratios, immunology, fertility, physiological and anatomical defects (Cole, 1910). This led to other research on the genetics of inbreeding, pathology, disease resistance and physiological development in poultry. In 1912 he also initiated basic research on genetic segregation and heterosis for contrasting traits in crosses of Angus with Jersey and Holstein breeds of cattle. Although results were interesting, the cattle crossbreeding experiment mainly illustrated the need for much larger experiments and for the study of variation in the general population of cattle breeds (e.g., Cole and Johnson, 1948).

Dr. Cole initially supervised graduate students in both plant and animal genetics. In 1919 he added a coworker, E. W. Lindstrom, to handle plant genetics. He was succeeded by R. A. Brink in 1922 when Lindstrom became Chairman of Genetics at Iowa State College. As the program grew, an amazingly broad spectrum of graduate students flocked to graduate training in genetics at Wisconsin. At this point in his career, Dr. Cole's earlier associations with W. J. Beal and the pioneering work of E. M. East and G. H. Shull sparked his interest in development of inbred lines and hybrids for commercial corn production. This work was initiated in collaboration with E. W. Lindstrom and was later developed by R. A. Brink and N. P. Neal. The earlier associations with Frank R. Lillie, H. S. Jennings, S. J. Holmes, Raymond Pearl, E. L. Mark, G. A. Parker and W. E. Castle also undoubtedly were most helpful in meeting the challenges of developing a highly diversified research and training program in basic and applied genetics. Although not himself trained in quantitative genetics, Cole was keenly aware of its potential importance for the improvement of economic characters in animals and plants, and he encouraged his students to study the work of Sewall Wright, and, later, Jay Lush and others in this emerging field.

The general approach of the "Chief' (the affectionate tide used by his students) was to develop cooperative research with the departments of Agronomy, Horticulture, Entomology, Poultry and Dairy and Animal Husbandry and to invite staff and graduate student participation. These research programs included genetic improvement of corn (Brink, Neal), potatoes (Rieman), clover (Smith), poultry (e.g., Martin, Jull, Maw, Asmundson, Jaap), dairy cattle, pigs, beef cattle and sheep (e.g., Lush, Craft, Yapp, Warwick, Turner, Chapman, Dickerson), bees (Laidlaw), pigeons (W. F. Hollander) and fur animals (Shackelford). One personal example was the cooperation with Prof. Humphrey, Chairman of Animal Husbandry, which involved one of the authors' (GED) Ph.D. analysis of University dairy herd records and DHIA records, as well as his later staff function in Extension dairy cattle breeding schools, preparation of the Wisconsin Bull Book of dairy sire and dam progeny tests by the Dairy Records Office, and in experimental studies of inbreeding effects in dairy cattle at Wisconsin State Institutional herds. Many of the later leaders in plant and particularly in animal genetics were among the 62 Ph.D. and 61 M.S. students trained during 30 years under Dr. Cole's leadership.

The department's program was expanded in 1930 to include the genetics of immune systems, beginning with pigeons and then extending to other species, under the leader-ship of M. R. Irwin (e.g., Irwin and Cole, 1945). Dr. Cole's earlier interest in possible methods of sex-control (Lush, 1925) was renewed in 1930 (Cole and Johansson, 1933; Cole et al., 1946b). In 1933, the genetics of reproductive physiology was added under the leadership of L. E. Casida. Work of the genetics of color phases in foxes and mink was begun by Dr. Cole after retirement as chairman of the department in 1939, in conjunction with a graduate student, R. M. Shackelford, who continued this work in the department at Wisconsin (e.g., Cole and Shackelford, 1946a). Another student, A. B. Chapman, developed and continued the collaborative research on the genetics of dairy cattle, beef cattle, swine and sheep, in addition to basic research on the genetics of growth and reproduction in laboratory rats. The Wisconsin group played a key role in establishment of the Regional Dairy Cattle Breeding Research Project ~C-2) and has participated actively in the Regional Swine, Beef Cattle and Sheep Breeding Re-search Projects. Thus, upon retirement, Dr. Cole had established and staffed a very broad research and graduate training program in both animal and plant genetics, in cooperation with other university animal and plant science departments (Brink, 1958).

Dr. Cole's approach to training placed more emphasis on the student's interest and capacity than on his own lecturing skills. The topics for weekly seminars were chosen carefully to challenge preparation by both the student speaker and the audience, and vigorous critical discussion was encouraged. Shared reporting of current literature also was assigned. Course programs for candidates in the animal genetics area were broad, including in-depth plant and animal genetics and statistics, plus biochemistry, nutrition, physiology, anatomy, endocrinology, and economics, as required to supplement the student's earlier background. Effective reading knowledge of science literature in two foreign languages also was required; a skill now generally supplanted by computer skills. These features led to a highly diverse group of staff and students, but one kept unified by the interchange of information and ideas at the single weekly seminar and by occasional circulation of the "Genotype" newsletter to both current and alumni students of the department. Dr. Cole was always helpfully but unobtrusively interested in the personal welfare of his students. Even though his "old school" background led him to discourage marriage until training was completed, he gracefully accepted the marriages of his students and commended their spouses for their support. Mrs. Cole (Daisy) also played a most supportive role in the social life of Cole's students, and in the rearing of their two children, Margaret Valeria (Mrs. Frank Husting) and Edward Goodenow.

Because of his broad interest and associations in the fields of biology and agriculture, Dr. Cole was called upon for a variety of wider responsibilities. In 1923, U.S. Agriculture Secretary Wallace requested a one-year leave of absence from the University for Cole to serve as Chief of the Husbandry Division in the Bureau of Animal Husbandry. In that capacity, Cole was able to stimulate more fundamental research at the branch stations of the Division, add key staff, such as Jull in poultry and Howe in nutrition, and expand the basic work of Sewall Wright in quantitative genetics and the animal breeding efforts of McPhee. He thus contributed to the strong scientific research emphasis that has characterized the Animal Husbandry Division and its successors.

In 1925, Cole was elected Vice-Chairman, and then Chairman in 1926-27, of the Division on Biology and Agriculture of the National Research Council, as well as Chairman of its Committee on Animal Breeding, to stimulate and coordinate breeding research. The benefits resulting from such contracts among workers on common research problems undoubtedly have contributed to later development of Regional Research Committees. He also represented agriculture on the Board of Biological Fellowships of NRC in order to assist promising young men in obtaining added scientific training. He served as President of the Wisconsin Academy of Arts and Sciences (1924-27) and of the Genetics Society of America (1940) and as a member of the Editorial Board of Genetics (1935-48). He also served on the Advisory Committee and the Council of the American Genetic Association and on the Council for the American Naturalist. In the 1 920s, Dr. Cole was invited to serve on an advisory committee for Margaret Sanger's Birth Control Clinical Research Bureau, and is credited with providing information concerning potential positive eugenic benefits of birth control. Recognition's included the honorary D.Sc. from Michigan State College in 1945 and the 1939 portrait hanging in the Saddle and Sirloin Club Gallery at Chicago (now in Louisville).

Leon J. Cole, for those who knew him personally as his students or colleagues, will always be remembered for his gentle and self-effacing encouragement when helping in the selection of significant research problems, in urging thorough study of related literature, careful planning and conduct of the research, and during the period of interpretation of results and preparing papers for publication. Above all, he taught us to use our own minds and abilities-he did not constantly monitor specific progress or provide close supervision. Yet, he did provide the essential opportunity and environment for self-growth in knowledge and experience. Dr. Cole's legacy to the field of animal genetics was primarily the first diverse U.S. core of well-trained geneticists ready for breeding research with all livestock species as well as for the training of the next generation of geneticists. He also left a strong Genetics Department at Wisconsin, equipped for research and graduate training in animal and plant genetics, immunogenetics and physiology of reproduction, in cooperation with appropriate other university departments. "L. J." was the right man at the right time to facilitate early application of genetics to animal and plant improvement in the United States. ---

¹Roman L. Hruska U.S. Meat Anim. Res. Center. ARS. USDA, A218 AnS, Univ. of Nebraska, Lincoln 68583-0908

²Depts. Of Meat and Anim. Sci. and of Genetics, Univ. of Wisconsin.

Received November 14, 1988

Accepted November 14, 1988

Literature Cited

ASAP, 1939. The Cole Dinner. In: Proc. Of 32nd Annu. Mtg. Of Am. Soc. Anim. Prod. 448-466.

Brink, R. A. 1958. A history of the Department of Genetics. Dept of genetics. Univ. of Wisconsin, Madison. (Mimeo).

Chapman, A. B. 1989. Leon Jacob Cole, In: Am. Council Learned Societies (Ed.) Dictionary of Scientific Biography (Suppl. II). Charles Scribner's Sons, New York (In press).

Cole, L. J. 1905. The German carp in the United States Rpt., U.S. Bureau of Fisheries 1904:523-641.

Cole, L. J. After 1910. Bound reprint files of Cole's publications. Dept. of Genetics. Univ. of Wisconsin, Madison, WI.

Cole, L. J. and I. Johansson. 1933. Sex control again. J. Herd. 24-246.

Cole, L. J. and I. Johansson. 1948. Inheritance in crosses of Jersey and Holstein Friesian with Abdereen-Angus cattle. Am. Nat. 72:145.

Cole, L. J. and R. M. Shackelford. 1946a. Breeding mutation mink and foxes. Wisconsin Agric. Exp. Sta. Spec. Bull.

Cole, L. J., E. Waletzkey and R. M. Shackelford, 1946b. A test of sex control modification of the acid-alkaline balance. J. Hered. 31:501.

Irwin, M. R. and L. J. Cole. 1945. Immunogenetic studies of cellular antigens: individual differences between species hybrids. Genetics 30:439.

Johansson, I. 1961. Leon Jacob Cole. Genetics 46:1.

Lush, J. L. 1925. The possibility of sex control by artificial insemination with centrifuged spermatozoa. J. Agric. Res. 30:893.

McCabe, R. A. 1979. Wisconsin's forgotten ornithologists: Leon J. Cole. The Passenger Pigeon 41:129.