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Perspectives Copyright 0 1994 by the Genetics Society of America Perspectives Anecdotal, Historical and Critical Commentaries on Genetics Edited by James F. Crow and William F. Dove Harvard, Agriculture, and the Bussey Institution J. A. Weir Division of Biology, University of Kansas, Lawrence, Kansas 66045 The genes are units useful in concise descriptions of the From a movement led by the physicist WALLACESABINE, phenomena of heredity. Their placeof residence is the chro- a GraduateSchool of Applied Science was organized in mosomes. Their behavior brings about the observed facts of 1906 to replace the undergraduate Lawrence Scientific genetics. For the rest, whatwe know about them is merely an interpretation of crossover frequency. In terms of geometry, School. As part of the reorganization in 1908, “Bussey” chemistry, physics, or mechanicswe can give them no descrip became a graduateschool for advanced instruction and tionwhatever. E. M. EAST (1926) research in scientific problems that relate and contrib Ute to practical agriculture and horticulture-this just at the time when the claims ofgenetics could no longer be ENETICS had its first influence in agriculture and ignored. In recommendingto President ELIOTthat W. E. G first achieved independent status in agricultural CASTLE’S work be transferred from Cambridge to the colleges. The main training ground was the Bussey In- Bussey Institution, Dean SABINEwrote, “The University stitution of Harvard University. has Professor Castle to start with; to lose him will be to By his will of 1835, BENJAMINBUSSEY left his extensive lose the best man in the country in genetics.” So instead farm (now the Arnold Arboretum) toHarvard to be held of going to Wisconsin or Yale, CASTLE moved hisanimals forever as a Seminary for “instruction in practical agri- to Forest Hills, soon LO be joined by the noted plant culture, in useful and ornamental gardening,in botany, geneticist EDWARDMURRAY EAST from the Connecticut and in such other branches of natural science, as may Agricultural Experiment Station. CASTLEand EASThave tend to promote a knowledge of practical agriculture, been thesubjects of recent Perspectives (SNELLand REED and the various arts subservient thereto and connected 1993; NELSON1993); see also the biographies by WRIGHT therewith.” The government of the University is also“to (1963) and JONES (1944). cause courses of lectures to be delivered there . and WILLIAMMORTON WHEELER, an erudite scholar who also to furnish gratuitous aid . to such meritorious rated artistry in taxonomy above all hisother powers, left persons as may resort there for instruction.” Texas in1908 to join the Bussey as Professor of Economic The bequest did not become available until 1861, and Entomology. He was appointed dean in 1915 when the the school did not open until 1871, following the con- Bussey was separated from the other schools of applied struction of a suitable building. The initial appoint- sciences, but in 1929 he resigned in favorof the more con- ments included, as instructor of farming, BUSSE~S genial atmosphere of the Museum of Comparative Zool- grandson-in-law, THOMASMOTLEY, who came with the ogy. IRVINGW. BAILEY, who came to the Bussey from the farm; FRANCIS STORER, brother-in-lawHarvard’s of Presi- defunct School of Forestry, acted as secretary. Between dent ELIOT,as professor of agricultural chemistry and 1912 and 1939, 44 doctorates in genetics were awarded. dean; andFRANCIS PARKMAN, famous the historian, as pro- Animal genetics had its roots in liberal arts, not ag- fessor of horticulture. riculture, with Johns Hopkins University as the early The resourcesfor the school were seriously de- leader. E. B. WILSON,the cytologist (Ph.D. 1881), T. H. pleted when a number of Bussey estate rental prop- MORGAN(Ph.D. 1890), E. G. CONKLIN(Ph.D. 1891), and erties were destroyed in theGreat Boston Fire of R. G. HARRISON (Ph.D. 1894), ostensibly students of the November 9-10, 1872. The meager income was aug- morphologist W. K. BROOKS,came under the influence mented by growing vegetables, cutting firewood, and of the physiologist H. NEWELLMARTIN and, except for CON- boarding horses. The school never did come up to WN, hrthered their studies inEurope. (MORGAN,who re- Harvard undergraduate standards,which were not all tained his interest in embryology,was the only one to be- that high, at best. come a member of the Genetics Societyof America.) Genetics 136: 1227-1231 (April, 1994) 1228 J. A. Weir At Harvard, C. B. DAVENPORT(Ph.D. 1892) and W. E. breeder, whereas H. C.MCPHEE (EAST’S student) became CASTLE(Ph.D. 1895) were students of EDWARDL. MARK, Chief of the Animal Husbandry Division of the U.S. Bu- whose doctorate in 1876 from the University of Leipzig reau of Animal Industry. P. W. GREGORYwent to Davis, was under RUDOLPHLEUCKART, an early proponent of the where he investigated the genetics of cattle, including need to combine morphology and physiology. Their the- size inheritance,and alsoworked with rats. EDWARD ses, based on morphological studies, were published in LIVESAYbecame professor of animal husbandry at West the Bulletin of the Museum of Comparative Zoology. Virginia. NELSON WATER^ went on to a career in poultry Both DAVENPORTand CASTLEhad farmbackgrounds and research. Four of CASTLE’S23 doctoral candidates spent later worked with laboratory and farm animals. DAVEN- their entire careers after graduation working with eco- PORT, with a B.S. degree in civil engineering, brought nomic species; four others were so engaged over a pe- quantitative methods to biology but turnedhis attention riod of years. EAST’Sstudent, W. R. SINGLETON,studied to eugenics and administration. CASTLE, whoassisted coat color inheritance in horses in collaboration with DAVENPORTas a student and succeeded him at Harvard CASTLE.The first generation of geneticists were able to when DAVENPORTdeparted in 1899, had little interest in pursue problems in pure and/or applied research. eugenics and nonein administration, but they remained CASTLE’Sown favorite specieswas the rat. He hadcome close friends, and DAVENPORT,through control of Car- to distrust the conceptof gametic purity on finding that negie funds, provided assistance to CASTLEat the Bussey. the Mendelizing guinea pig characters, polydactylism, Both men were interested in heredity even before 1900, long hair, and rough coat were not exactIy the same so their grasp of Mendelism, after its rediscovery in 1900, when extracted from crosses. To test his hypothesis, he was not immediate because they carried a considerable undertook aselection experiment with hooded rats and baggage of seemingly conflicting evidence. by 1914 had rearedand studied the color pattern of over After the rediscovery of MENDEL’Slaws, there ensued 25,000 rats. With great persistence he defended his hy- lively debate about the extentto which the laws applied pothesis in the face of mounting criticism-some rather to animals, including man.Naturally it was the conspicu- heated. EMERSONand EAST’Sclassic paperon maize, ous characters that attracted attention at first, but, as the showing that quantitative charactersare inherited in Men- inheritance of more and morecharacters was shown to delian fashion,appeared in 1913,a year beforethe comple- behave according to simple Mendelism or oneof its eas- tion of CASTLE and PHILLIPS’monograph (1914).The con- ily understood modifications, attention shifted to the cept was debated in Bussey seminars, with CASTLE and EAST materials in their own right-on one hand,as sources for holding opposing views. CASTLE was not convinced until he new problems; on the other, as a new approach to the performed the critical experiments suggested by WRIGHT. solution of old problems, such as the limits of selection His retraction appeared in the American Naturalist in 1919 or the consequences of inbreeding. (CASTLE 1919).In L. C. DuNN’swords, “Castlecured himself From 1901 to 1905, CASTLE,with the cooperation of of disbelief inthe integrity of the gene the hard way-by 15 four students in successive years, followedthe effects of years of arduous experimentation.” inbreeding in Drosophila melanogasterthrough 59 gen- It is noteworthy that bothCASTLE and EASTconsidered erations of brother-sister mating. After the move to the it the function of the professor to provide space, mate- Bussey, no one used Drosophila for thesis research, but rials, encouragement, and little else, with course work in it became the favorite material for Harvard undergradu- diverse areas only suffkient to broaden the students’ ate teaching in Cambridge, so the graduate students perspective. CASTLE’Sfirst graduate students, JOHN A. were familiar with its usefulness. They kept abreast of DETLEFSEN,who axrived in 1908 and worked with guinea developments and some, notably E. C. MACDOWELL,L. C. pigs, and EDWNCARLETON MACDOWELL,who arriveda year DUNN,and S. C. REED, later used Drosophila for research. later and worked with rabbits, receivedtheir doctorates in Although he emphasized the importance of labora- 1912. They did not share CASTLE’S skepticism concerning tory animals in the study of inheritance, CASTLE retained the multiple-factor theory, and MACDOWELL’Sthesis, pub- an interest in farm animals. Between 1940 and 1961 lished as a monograph (1914),carried a prefatory note (after retiring to Berkeley) he published 17 papers on by CASTLE:‘Wile not entirely sharing his views, I have coat colors in horses, and while at the Bussey he pub- tried not to bias his judgment either for or against the lished on inheritance in sheep and dairy cattle and on multiple-factor hypothesis which he adoptsin this paper. various aspects of animal breeding. But to avoid misunderstanding, I wish to say that in my After graduation, a number of CASTLE’Sstudents and own opinion the theory of the purity of the gametes has two of EAST’Sworked with economic animals. JOHN not been established.” CASTLEwas aware that the top DETLEFSENworked with rats, mice, and cattle at Illinois. signer of the thesis is not the only or even the chief source SEWALLWRIGHT went to the Bureau of Animal Industry, of guidance.
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