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JAY LAURENCE LUSH January 3, 1896May 1, 1982 BY ARTHUR B. CHAPMAN A Y BEAU RE N C E ~ U S H macle the following autobiographi- ~J Cal statement in ~ 967~: I was born in a log house on a farm in southwestern Iowa tSham- baugh], the second of six children.... My father was born in Canada. His parents were brought as children from southern England. My mother's father came from northern Ireland, but her mother was of old American stock, Scotch and Scotch-Irish in origin. In our home we read many books of the kinds which were still consid- ered classics around 1900 to 1910. Although money was scarce, we always had enough to eat, plenty to read, and clothing enough to keep warm. I went to an ungraded country school and entered a high school in Kansas at the age of 11. At Kansas State Agricultural College (now Kansas State University) I majored in animal husbandry. Mathematics was easy but not intriguing. History, physical geography, geology and parts of chemistry and biology were most interesting. I was active in debating. About 1914 I got my first intriguing glimpses of genetics. Also I encountered several interesting, friendly and challenging professors, mostly in biology or some ,. . . . at Its app lcatlons. After receiving the B.S. degree in 1916, I taught agriculture and chemistry in a Kansas High School for a year; then returned to KSU for my Master's degree and an apprenticeship in agricultural research. My first contribution to science was an article printed in the./ournal of Heredity 12:57 - 71 in 1921. This was what I then thought was worth publishing from my Master's degree. I spent nine months in the Air Force immediately after receiving the ' Autobiographical statement, National Academy of Sciences, 1967. 277
278 BIOGRAPHICAL MEMOIRS M.S. degree and I was commissioned as a Second Lieutenant in the reserve in February of 1919. I installed the Smith-Hughes program of agricultural instruction in another Kansas High School in the early part of 1919. I went to the University of Wisconsin in June of 1919 to do more graduate study In genetics. Immediately after finishing my Ph.D. work there t1922], I went to the Texas Agricultural Experiment Station at College Station, Texas. An important bit of Lush's personal history for the follow- ing year 1923- was his marriage to Adaline Lincoln. Mrs. Lush, a second cousin once removed of Abraham Lincoln, is a truly remarkable person. She gracluated from high school at the age of thirteen and from the University of Arkansas at sixteen; she then earner! a master's clegree at the University of Chicago at age seventeen. In an aware! speech given when Lush was being honoree] at the Poultry Breeders Rouncitable meeting in 1969, the speaker, Arthur Heisclorf, made this remark about Mrs. Lush: "l think she is the person who has been the secret catalyst twho] has sparked Dr. Lush onto the accomplishments he has made." To this tribute should be aciclect how important a role she has taken as a gracious host- ess ant! "foster mother" to countless students. She also found timeand had the abilityto teach French, German, Italian, Latin, and Spanish to private pupils; to concluct a number of trips to Europe; and to be active in a number of organiza- tions. She was namect Iowa Mother of the Year in 1963. Dr. and Mrs. Lush have a daughter, Mary Elizabeth Hausrath, a son, David Alan, and seven surviving grandchil(lren. Lush's history, as recounted by him in 1967, continues below: For more than eight years I did research in animal husbandry [in Texas]. Most of that pertained to animal breeding, but some of it was in other areas of animal production. The necessities of the research drew me fur- ther into biometry. In January of 1930 I came to Iowa State University (then Iowa State College) as Professor in the Department of Animal Hus- bandry to do research and teaching in Animal Breeding.... All of my
JAY LAURENCE LUSH 279 work has hinged around finding ways to apply genetics more efficiently in improving animals and plants. For these purposes I used many biometrical tools developed by others and for myself made a few minor innovations in those. Most of my discoveries were small ones, usually growing out of some actual problem in application. These are put together in some 200 research papers and in my textbook, Animal Breeding Plans F1937] which has sold more than 22,000 copies. It is currently being printed in its fourth language [Spanish; earlier in Polish, Portugese, and Rumanian]. Perhaps the most important single paper was one in the American Naturalist in 1947 entitled "Family merit and individual merit as bases for selection." In 1972 a symposiums was held in his honor. All the pa- pers presented except one by Lush himself, "Teaching Ani- mal Breeding ant! Training Graduate Students" (1973), were by Lush's former students or one-time colleagues at Iowa State University. These papers reflect the high esteem in which Lush was hell! as research worker, teacher, and hu- man being by those who knew him best. The deep insight ant! extensive coverage given by these papers to Lush's life ant! contributions to his chosen field have led me to quote extensively from them in this biographical memoir. A former student, R. R. Shrode, introcluced the sympo- sium and captured the essence of Lush's contributions: In effect, the field of Animal Breeding is a program of intellectual "linebreeding" to Lush. It is with tremendous professional respect and personal affection for our honoree that we dedicate this Symposium to our friend and teacher, lay L. Lush, who has contributed more than any other individual, directly and indirectly through his many students, toward the continuing evolution of Animal Breeding from an art into a science.3 2 The Animal Breeding and Genetics Symposium in honor of Dr. Jay L. Lush, sponsored jointly by the American Society of Animal Science, American Dairy Sci- ence Association, and Poultry Science Association, Virginia Polytechnic Institute and State University, Blacksburg, Virginia, July 29, 1972 (1973). Copies of the sympo- sium proceedings may be obtained from Business Office, American Society of An- imal Science, 309 West Clark St., Champaign, Illinois 61820. 3 R. R. Shrode, "Introduction Why We Are Here," in Proceedings of the Animal Breeding and Genetics Symposium in Honor of Dr. jay L. Lush, American Society of
280 BIOGRAPHICAL MEMOIRS An Iowa State University colleague, A. E. Freeman phrased it this way: As problems arose and could be defined in a mathematical or statistical sense, he and his students found answers to them in a way useful to im- proving domestic animals. The emphasis on breeding plans did not pre- clude interest and work on problems of a more theoretical nature. He clearly contributed to problems of almost purely theoretical interest, at least at the time; but it is safe to say that most of this work was started by seeing an actual problem arise that generated the germ of an idea for the theoretical work.... Dr. Lush's special interest in animal breeding was definitely aroused in 1914 by the teaching and enthusiasm of E. N. Went- worth Esee Lush's obituary for Wentworth (1962) and his response during the dedication of the Jay L. Lush Auditorium at Iowa State (1974~] who was later his major professor for the M.S. degree fat Kansas State Agri- cultural College].... flush] continued his graduate training at the University of Wisconsin under the direction of Dr. L. I. Cole. Though it may now seem a bit strange, Dr. Lush was a physiologist. His Ph.D. thesis was "The possibility of sex control by artificial insemination with centrifuged spermatozoa" (Lush, 1925~. He didn't succeed in this venture, but neither has anyone since. His interest in measurement and use of statistical tools was clear in this work. The data were arrayed by expected sources of variability, cor- relation coefficients were computed and probable errors were used to help determine if associations were real. Also, he fit normal curves to distribu- tions of sperm head length measurements and tested these for goodness of fit. So, even as a physiologist, Dr. Lush's interest in measurement and statistics was evident.4 G. E. Dickerson, a former colleague at Iowa State Univer- sity, referred in his symposium paper to the influence Sewall Wright's work had on Bushes biological and statistical think- ~ng: Animal Science, American Dairy Science Association, and Poultry Science Associa- tion, Blacksburg, Virginia, July 29, 1972 (1973), p. iii. 4 A. Freeman, "Genetic Statistics in Animal Breeding," in Proceedings of the Animal Breeding and Genetics Symposium in Honor of Dr. jay L. Lush, American Society of Animal Science, American Dairy Science Association, and Poultry Science Associa- tion, Blacksburg, Virginia, July 29, 1972 (1973), pp. 1, 2, 3.
JAY LAURENCE LUSH 281 How packed with meaning this subject fInbreeding and Heterosis in Animals] is for animal breeders! And how greatly our understanding of the potential usefulness of inbreeding and heterosis in animal improve- ment has expanded during the last four decades as a result of the research, writing and teaching of Dr. Jay Laurence Lush! While Dr. Lush was busy at Texas A&M from 1922 to 1930 publishing studies of inheritance and performance evaluation, he must also have been studying Sewall Wright's interpretations of the U.S.D.A. inbreeding and crossbreeding work with guinea pigs (19221.5 This seems clear from his 1927 paper clarifying the limitations of "percentage of blood" in describing genetic likeness, partic- ularly among collateral relatives and from the subsequent series with his students and collaborators on the amount and kind of inbreeding, occur- ring during breed development in cattle, sheep and swine (1932 to 1939, 1946), using the technique of Wright and McPhee (1925~6 for sampling random lines of ancestry. When Dr. Lush arrived at Iowa State in 1930, earlier experiments with full-sib inbreeding in swine at Iowa and elsewhere had been discontinued due to loss of fertility. However, Wright's theoretical analyses and some results with guinea pigs (1921,7 1922S) had indicated that selection might be able to offset unfavorable effects of milder inbreeding and that inbreed- ing was a powerful tool for creating genetic diversity among lines. This led Dr. Lush to initiate an experiment in 1930 comparing intense and mild linebreeding in pigs, with concurrent individual and progeny test selection. During this same period (1933), Lush's famous bulletin on linebreeding was published. It eloquently stated the case for subdivision of breeds into many lines, each mildly linebred to carefully selected ances- tors, with continuous elimination of the poorer ones and recombining of better ones, closely paralleling Wright's (193119 ideas on optimum popu- lation structure for evolution.~° 5 S. Wright, "The Effects of Inbreeding and Crossbreeding on Guinea Pigs," U.S. Department of Agriculture Bulletin, lO90(parts 1 and 2, 1922); 1121 (part 3, 1922). 6 S. Wright and H. C. McPhee, "Approximate Method of Calculating Coefficients of Inbreeding and Relationship from Livestock Pedigrees," I. Agric. Res. (Washington, D. C.), 31 (1925): 377-83. 7 S. Wright, "Systems of Mating," Genetics, 6(1921): 111-78. ~ S. Wright, "Effects of Inbreeding and Crossbreeding," 1090. 9 S. Wright, "Evolution in Mendelian Populations," Genetics, 16(1931):97-159. t° G. E. Dickerson, "Inbreeding and Heterosis in Animals," in Proceedings of the Animal Breeding and Genetics Symposium in Honor of Dr. jay L. Lush, American Society of Animal Science, American Dairy Science Association, and Poultry Science Asso- ciation, Blacksburg, Virginia, July 29, 1972 (1973), pp. 54-77.
282 BIOGRAPHICAL MEMOIRS Freeman also pointed to Sewall Wright's influence on Lush's thinking: "Lush commuted tin 1931] to the University of Chicago to audit Sewall Wright's course in Statistical Ge- netics and other Zoological courses there. The influence of this training and these visits with Dr. Wright on Dr. Lush's teaching and research is evident." Lush said, at the Poultry Breeders Roundtable in 1969: "Those were by far the most fruitful ten weeks ~ ever had." R. A. Fisher's work was also caller! on frequently by Lush, as Freeman states: "Before about 1930, the primary statistical tools used in animal breeding were correlation anc! regres- sion methods. R. A. Fisher lecturect at {owe State through the summers of 1931 and 1936. Fisher's work greatly ad- vanced the knowlecige and use of statistics. Dr. Lush was unique in combining the work of both Fisher and Wright to solve animal breeding problems." In what specific areas of animal breeding were Lush's con- tributions macle? Freeman notes the following: Many of his early papers explored husbandry problems; others con- sidered the mode of inheritance of qualitative traits; some were concerned with measurement and description of economically important traits; and others are clearly a start of current-day animal breeding theory. In the early years, the first three types of papers were more numerous than the last type. Of course, this changed. As Dr. Lush was confronted with ques- tions that stemmed from practical problems, he tried to answer them from the existing knowledge in classical genetics, plant breeding or allied theory. If existing knowledge did not yield an acceptable answer, he used statistical methods to better describe problems or relations existing between traits, then put this statistical description into a form usable by the breeder.... Many of Dr. Lush's publications from 1926 to 1930 could be described as developing and using more accurate ways to measure quantitative traits.'3 Lush undertook studies using records collected on swine, dairy cattle, beef cattle, sheep, goats, poultry, and honeybees. " Freeman, "Genetic Statistics," p. 4, 5. |2 Ibid., p. 5. |3 Ibid., p. 3, 4.
JAY LAURENCE LUSH 283 In 1930 he also initiated an experiment on "closed-herd" se- lection in dairy cattle as well as the one on selection and in- breeding in swine. Both of these long-term experiments pro- vided data for many M.S. and Ph.D. theses and resulted in major contributions to the field of animal breeding. R. W. Touchberry, another former student, gives a de- tailed discussion in his symposium papery of "some of the pertinent points in a few of what ~ [Touchberry] consider to be his LLush's] most important papers." For those who are familiar with the terminology of genetics, statistics, and ani- mal breeding, Touchberry's paper provides a summary of many of Lush's contributions to the field. ~ will attempt to give the essence of these contributions. There is one paper (Lush, 1947) that serves well as a pro- totype for many of his papers. It is also the one that Lush considered his "most important single paper." It can be used to illustrate his way of thinking about a problem and how that approach leads to a solution. The study began, as did so many of Lush's projects, with a practical problem: "how much attention ought to be paid to the merits and defects of litter mates when choosing boars and gilts to use for breeding." The problem developed into the more general one of asking how much a population mean would be changed by selecting on individual performance alone versus selecting on family merit alone versus selecting on a combination of the two. How did Lush approach this and similar problems? He started with the fundamental principles of genetics; then, by invoking a deductive argument, he gave them effect through the use of the tools of population genetics (discontinuous classes, qualitative differences) and biometrical genetics (con- '4 R. W. Touchberry, "The Life and Contributions of Dr. lay Laurence Lush," in Proceedings of the Animal Breeding and Genetics Symposium in Honor of Dr..Jay L. Lush, American Society of Animal Science, American Dairy Science Association, and Poul- try Science Association, Blacksburg, Virginia, July 29, 1972 (1973), p. 89.
284 BIOGRAPHICAL MEMOIRS sinuous distributions, quantitative or measurement differ- ences). In this American Naturalist paper, Lush points out: "The process of selection consists only of predicting the breeding value Genetic make-up or transmitting ability for the trait under consideration of each individual which Is being considered and then keeping it or culling it on the basis of that prediction. If the same fraction of the population must be saved but there is a choice of bases on which selection may be macle, then the difference in results depends only on how accurately each inctividual's breeding value can be pre- clictect from each of these bases."~5 In comparing the three bases of selection in this paper, Figure ~ is usect as a graphic way of looking at the interre- lationships between the variables. The arrows in this diagram react from "cause" to "erect," and the value attached to each one is clefinect as a path coefficient (stanclarcI partial regres- sion coefficient) by Sewall Wright, who developecI this pro- ceclure.~6 The bictirectional arrows refer to simple correla- tions between the variables. In this diagram Pj stancts for the phenotype Observed measured value) of an individual, i; Y. the average phenotype of a family; Gi, the breeding value of an incliviclual (average effects of the genes it contains Fish- er's "expected value"~71; W. the factors other than Gj that af- fect each Pi in a family the same way but may diner from family to family; Ui, the factors other than Gi that affect Pi but that are no more alike for members of the same family than they are for individuals that belong to different families; . 15 J. L. Lush, "Family Merit and Individual Merit as Bases for Selection," American Naturalist, 81 (1947) :243 - 44. t6 S. Wright, "On the Nature of Size Factors," Genetics, 3(1918):367-74; "Corre- lation and Causation," Journal of Agricultural Research (Washington, D.C.), 20(1921):557-85; "The Method of Path Coefficients," Annals of Mathematical Statis- tics, 5(1934):161-215. ]7 R. A. Fisher, The Genetical Theory of Natural Selection (New York: Oxford Uni- versity Press, 1930).
JAY LAURENCE LUSH Uj ,' G - 9 /T' `, 2 1\ r 9 n rGP g rG t;12 1 +(n-1)r p 1 + (n - 1 It] rpy= ~ 1+(n1)t t = r =92r+r2 285 \ - A FIGURE 1 Path coefficient diagram of biometrical relations involved in mass, family, and combination selection. Source:.T. L. Lush, "Family Merit and Individual Merit as Bases for Selec- tion," part I, American Naturalist, 81(~1947~:246.
286 BIOGRAPHICAL MEMOIRS and r, the correlation between the breeding values of mem- bers of a family (Wright's relationship coefficients. The algebra to which this approach leads has been spelled out by Wright (see note 16) in a form that relates correlation coefficients to their path coefficient components. Some of the pertinent correlations in terms of path coefficients are given below the diagram in Figure I. The correlations (rap, ray, and rot) reflect the relative progress expected under the three methods of selection, "where ~ is the index or most probable breeding value of an individual, as estimated from the optimum linear combination of attention to its own phe- notype and attention to the average phenotype of the family to which it belongs." The phenotypic correlation between members of a family is clenotect by t. As an example of the use that can be macle of these cor- relations in terms of their path coefficient components, let's look at the make-up of rap and ray. If selection is practiced on family average (Y) alone, the progress ma(le would be expected to be ~+(n-I)r/~/nt!+(n-I)t] times as rapicl as mass selection (selection on incliviclual performance); that is, rely = gEl+(nI)r]/~/ntI+(n-~)tJ versus rap= g, where g = the path of influence from G to P or the square root of heritability (the portion of the phenotypic variance due to genetic differences between inclivicluals), and n = the num- ber of inctividuals in the family. By inserting the values for g, n, r, and t that apply in a particular case, the difference in predictive value for transmitting ability from Y and P be- comes evident. As a graphic representation of the relative effectiveness of the three methods of selection, Lush (1947) gives the fol- lowing diagram (Figure 2) for a case where n = 21. The progress from mass selection is represented! by a level plane '8 S. Wright, "Coefficients of Inbreeding and Relationship," American Naturalist, 56( 1 922) :330-38.
JAY LAURENCE LUSH ~p3044 ~ ~/^ ~ / /// /~ r 287 - 7.0 -6.0 -5.0 -4.0 nip -3.0 -2.0 - 1.0 O. .8 '~ 0.98 o 'No Upper surface = combination selection Mass selection Family selection Case of n = 21 FIGURE 2 Relative effectiveness of the three methods of selection at all combinations of r and t when families contain 21 individuals. Source: }. L. Lush, "Family Merit and Individual Merit as Bases for Selec tion," part I, American Naturalist, 81~1947~:256. __11 ~~ 111CL11~ I! l~ ~ l~ ~! at height I.0 above the base and where all combinations of r and t are given. The above gives the basic approach used in this paper anct many others by Lush, but the full substance of this study goes much further. Reaclers of Lush's papers were usually made , aware <~s they are in this one) of the danger of accepting conclusions without taking into account the role playecl by chance, the frequent need for making simplifying (possibly oversimplifying) assumptions in order to Granola with ~ problem, ant] the errors likely to be made by semantic a~-- guments that are not supported by experimental evidence or by quantitatively evaluated deflective arguments. -r -- ~~ - - - _ _ A__ _w ~,1 `~1~ vv lLll O.
288 BIOGRAPHICAL MEMOIRS As an illustration of one of the caveats stemming from the above in the paper uncler discussion, he points to this possi- bility: If the actual effects of a gene substitution upon P are sometimes larger and sometimes smaller than the average eject of that gene substitution in that population, depending upon what other genes are present, the dif- ferences between the actual and average effects are termed epistatic or dominance deviations. These give rise to "special" breeding values. That is, they cause some matings to produce better offspring and some to pro- duce poorer ones than would be expected if one knew the kind of offspring those same parents would produce if mated to a representative sample of the whole population. Most of the dominance and epistatic deviations from the additive scheme are included in U. but a small part are in W. He also points to the neecI to keep in mind other factors: the fiducial limits of the estimates; the role that mutations might play; the effect of selection in the same or in a clifferent direction within the population; the need for experiments to check on the theory involved in this work; ancT "the naive view, repeatecIly disproven but still often inferred to be axi- omatic, that family, breect, and race are unimportant, or even unreal, unless the families, breeds, or races are so distinct that they clo not overlap at all." Lush showed statistically that "family selection is most superior to mass selection when fam- ity members resemble each other least; i.e., when the families overlap widely in their phenotypes and t is therefore low!" As is usual in Lush's papers, he not only answers the spe- cific questions asked but expands the answer to encompass much more. In this case he states the solution in terms of intercIass correlations and then translates the solution into analysis of variance and intracIass correlations in the hope that the "biometrical relations may be clearer." These trans- lations have uncloubtecIly helped students understand the equivalent meanings of two important ways of stating the solution statistically. Furthermore, he provicles, in regression form, the equa- · .
JAY LAURENCE LUSH 289 tions for predicting breeding value of the incliviclual basecl on its own phenotype and its family average. He discusses uncler what conditions negative attention is pair! to family average and the elect inbreeding wouicI have on the results. Family selection for "all-or-none" characteristics, as well as characteristics ctistributed continuously, is clearly cliscussecI. A number of other qualifications ant! special conditions are also mentioned. And finally, the implications of these conclu- sions for man are given consideration. This is a paper in which the conclusions and interpretations make a contribu- tion to genetics, to animal and plant breeding, to statistics, and to sociology and anthropology. Many of his other papers also have a significant bearing on problems in several fielcis. The other papers summarized by Touchberry,~9 which fol- low the same pattern as the one just cliscussect, macle major contributions to animal breecling by clarifying problems in- volving progeny tests, inclivi(1ual performance, pedigree in- formation, heritability, selection, ant! the role player! by chance. Some of the sources of information used by Lush in his research were the records from private farms enrolled in the Iowa Cow Testing Association, on animals registered in the breech associations and on poultry of the Kimber Poultry Farm. These proviclecl an insight into the genetic and envi- ronmental sources of variation in economically important traits uncter commercial conditions. Lush was also associated with an Atomic Energy Commission research project on the . ~ ,% . . . . . . . genetic ettects ot ionizing rat latlon in swine. Lush's view on teaching is given in the paper "Teaching Animal Breeding ant! Training Graduate Students," which he presented at the symposium in his honor in 19~73. His introductory remarks in this paper reflect an attitude of his that was clearly evident to his students in his teaching. '9 Touchberry, "Life and Contributions," pp. 89-104.
290 BIOGRAPHICAL MEMOIRS Immediately on completing my Ph.D. degree, I did research for more than 8 years, with almost no interruption for teaching. I'm glad it hap- pened that way. If I had taught the same course as much as three times in succession, using the available texts and my graduate notes and all the rest of what I thought I knew, I would surely have come to believe those things myself so firmly that the errors among them could scarcely have been corrected by any amount of subsequent experience. As it was, the cattle and sheep and goats talked back to me. Having no papers to grade or class rolls to call, I listened. Usually the animals were saying something like: "Most of the things you think you know may be true in principle but you have many of them out of all proportion to their actual importance. When you draw a conclusion, you often overlook circum- stances which, if you considered them properly, would upset your recom- mendations badly." Trying to solve these apparent inconsistencies drove me, whether I wished it or not, in the direction of measuring more accu- rately the factors in the problems. I was always needing to be surer of how the various factors interacted in any whole operation we might be consid- ering. Lush had a penchant for those apposite sayings that are so helpful to students trying to think through what is meant by some of the more esoteric concepts of genetics, statistics, and animal breecting. In my symposium paper, ~ have quoted a number of these. One that ~ fount] myself using quite frequently in my teaching because of the effect it had had on me is one that he user! because "Some of us think we have seen signs that many populations Lbeing selected] clo not actually change fas] rapidly fas we think they should]: Heritability may not be as high as we think. Selection may not have been as intense as we think. Perhaps the rate of 203. L. Lush, "Teaching Animal Breeding and Training Graduate Students," in Proceedings of the Animal Breeding and Genetics Symposium in Honor of Dr. Jay L. Lush, American Society of Animal Science, American Dairy Science Association, and Poul- try Science Association, Blacksburg, Virginia, July 29, 1972 (1973), p. 78. 2] A. B. Chapman, "Selection Theory and Experimental Results," in Proceedings of the Animal Breeding and Genetics Symposium in Honor of Dr. lay L. Lush, American Society of Animal Science, American Dairy Science Association, and Poultry Science Association, Blacksburg, Virginia, July 29, 1972 (1973), pp. 42-53.
JAY LAURENCE LUSH 29 progress actually is substantially as much as it should be." Dr. Lush then points out that "we ought to think of the old adage that when one is asked to explain how it is that witches can turn milk blue, the first thing is to find out is whether witches really can turn milk blue!"22 His success as a teacher and director of graduate students (26 who earned the M.S. anct 124 a Ph.D. uncler his direction is admirably presented in Touchberry's symposium paper, which he concludes by saying: As an advisor of graduate students, Dr. Lush was highly respected and admired. He was firm without being threatening and he got his points and message across without raising his voice or using profanity. He was a warm and friendly person with a tremendous respect and tolerance for stu- dents.23 Lush's influence on animal breeding around the word has been enhanced greatly by the wide distribution of his classical book Animal Breeding Plans ( 19371. His equally authoritative mimeographed notes, "The Genetics of Populations" (1948), have also played a major role in the thinking of animal breed- ers who were lucky enough to have them. He playecl a major role in establishing anct was an active participant in the regional laboratories for animal breeding research joint ventures of cooperating states and the U.S. Department of Agriculture. He was also instrumental in the formation and guidance of the National Poultry Breeders' Roundtable, an organization of commercial poultrymen and academic staff that meets annually to discuss research in ge- netics and in animal and plant breecling. The meeting in 1969 (Eighteenth Annual Session, May 7-~) was helct in his honor. 225. L. Lush, "Summary (Symposium on Selection, Chicago, November 1949)," fournalofAnimalScience, 10(1951):18 - 21. 23 Touchberry, "Life and Contributions," p. 104.
292 BIOGRAPHICAL MEMOIRS Lush acted in an advisory capacity to these and many other organizations, both foreign and domestic. (He traveled extensively abroad ancT served as an advisor on animal breed- ing in a number of countries.) As a result, he was responsible for a profound change in the approaches to animal breeding research ant! practice in many countries. Lush received a number of honors and awards, one of which was to clecticate to him the Iowa State University Au- clitorium in 1973. His remarks on this occasion (1974) illus- trated his humility and include a typical "Lush" analogy: "~:'m Proust of this occasion, of course, although ~ am uncomfort- ably aware that others cleservecl the honor more. Also, ~ know that sheer luck had much to do with the things ~ dicI accom- plish. In some ways they resemble the occurrence of an ava- lanche in the mountains. For an avalanche to occur at all requires some snow, of course, but the small event which ac- tually triggers the avalanche mightjust as well have happened a hunctred yards to the right or to the left, or it might as well have happened yesterday or not until day after tomorrow! An avalanche is contagious in that once it starts, it jars things loose for hundreds of yards around." A fitting ending to this biographical memoir is the sym- posium statement of Touchberry: "He tEush] has defined the problems of genetically changing farm animals in a logical, biological, quantitative and economic way. Further, he has shown how genetics and mathematics can help in solving problems of animal breeding. In cloing this he has benefi- cially affected the lives of many. Thus, to me, it seems fitting to say that, rather than having followed a profession, he has, for the past 40 years, led a profession."24 24 Ibid.
JAY LAWRENCE LUSH HONORS AND DISTINCTIONS HONORARY DOCTORAL DEGREES 1957 Royal Agricultural College of Sweden 1957 Faustus Liebig University, Giessen, Germany 1958 Royal Veterinary and Agricultural College of Denmark 1964 Michigan State University 1969 University of Illinois 1970 Kansas State University 1970 University of Wisconsin 1971 Swiss Federal Institute of Technology, Zurich 1975 Agricultural University of Norway AWARDS 293 1946 Morrison Award of the American Society of Animal Science 1956 Honored Guest, American Society of Animal Science 1957 1958 Charles F. Curtiss Distinguished Professor in Agriculture, Iowa State University Borden Award for research in dairy production, American Dairy Science Association 1960 Herman von Nathusius Medal of the German Society for Animal Breeding 1965 Armour Award for animal breeding and genetics, Ameri- can Society of Animal Science 1965 Medal of the Mendel Centennial Association, Czechoslova- kia 1966 Order of Merit in Science, Italy 1968 National Medal of Science MEMBERSHIPS 1967 Member, National Academy of Sciences 1972 Member, Royal Society of Edinburgh Foreign member of the Academies of Science or Agricul- ture of Sweden, Norway, and Italy
294 BIOGRAPHICAL MEMOIRS B I B LI OGRAPH Y 1921 Inheritance in swine. i. Hered., 12:57-71. 1922 An hereditary notch in the ears of {ersey cattle. {. Hered., 13:8- 13. The influence of age and individuality upon the yield of wool. Proc. Am. Soc. Anim. Prod., 1922:105-9. 1923 With E. N. Wentworth. Inheritance in swine.J. Agric. Res.,23:557- 82. With i. M. Jones and i. H. tones. I. Fattening steers on cottonseed meal and hulls with and without corn. II. The influence of age on fattening steers. Tex. Stn. Bull. 309. With I. M. Jones. The influence of individuality, age and season upon the weight of fleeces produced by range sheep. Tex. Stn. Bull. 311. 1924 Twinning in Brahma cattle. I. Hered., 15:25 -27. "Double ears" in Brahma cattle. I. Hered., 15:93-96. With I. M. tones. The influence of individuality, age and season upon the weights of fleeces produced by Angora goats under range conditions. Tex. Stn. Bull. 320. 1925 The possibility of sex control by artificial insemination with centri- fuged spermatozoa. J. Agric. Res., 30:893-913. With J. M. Jones. Methods of selecting wool samples in shrinkage studies. Proc. Am. Soc. Anim. Prod., 1925: 115 -17. 1926 Practical methods of estimating the proportions of fat and bone in cattle slaughtered in commercial packing plants. J. Agric. Res., 32:727-55.
JAY LAURENCE LUSH 295 Inheritance of horns, wattles, and color in grade Toggenburg goats. I. Hered., 17:72-91. With W. H. Black. How much accuracy is gained by weighing cattle three days instead of one at the beginning and end of feeding experiments. Proc. Am. Soc. Anim. Prod., 1926:206-10. 1927 "Percentage of blood" and Mendelism. i. Hered., 1 8 :35 1- 67. Practices and problems involved in crossbreeding cattle in the Coastal Plain of Texas. Proc. Am. Soc. Anim. Prod., 1927:58- 61. With I. M. tones. A statistical interpretation of some Texas lamb feeding data. Proc. Am. Soc. Anim. Prod., 1927: 167-70. 1928 Changes in body measurements of steers during intensive fatten- ing. Tex. Stn. Bull. 385. With F. W. Christensen, C. V. Wilson, and W. H. Black. The accu- racy of cattle weights. I. Agric. Res., 36:551-80. 1929 With W. H. Black and A. T. Semple. The use of dressed beef ap- praisals in measuring the market desirability of beef cattle. I. Agric. Res., 39: 147 - 62. Atavism in Jersey cattle. I. Hered., 20:381-83. Twins in Jersey cattle. i. Hered., 20:510 -13. With I. M. tones. The inheritance of cryptorchidism. Proc. Am. Soc. Anim. Prod., 1929:57-61. 1930 "Duck-legged" cattle on Texas ranches. J. Hered., 21:84-90. With I. M. tones and W. H. Dameron. The inheritance of cryptor- chidism in goats. Tex. Stn. Bull. 407. With I. M. tones, W. H. Dameron, and O. L. Carpenter. Normal growth of range cattle. Tex. Stn. Bull. 409. With O. C. Copeland. A study of the accuracy of measurements of dairy cattle. I. Agric. Res., 41:37-49. Earlessness in Karakul sheep. J. Hered., 21:107-12. "Nervous" goats. J. Hered., 21 :242-47.
296 BIOGRAPHICAL MEMOIRS How farm animals inherit. (Review of Christian Wriedt's Heredity in Live Stock. ~ J.. Hered., 21 :306 -8. 1931 Interpreting the results of group feeding experiments. Proc. Am. Soc. Anim. Prod., 1930:44-55. The number of daughters necessary to prove a sire. }. Dairy Sci., 14:209-20. Predicting gains in feeder cattle and pigs. I. Agric. Res., 42:853- 81. 1932 Genetic aspects of the record of performance work with swine. Proc. Am. Soc. Anim. Prod., 1931:51-62. With M. D. Lacy. The ages of breeding cattle and the possibilities of using proven sires. Iowa Stn. Bull. 290. With M. D. Lacy. How old are your bulls and cows and what dif- ference does it make? (Abridged ed. Iowa Stn. Bull. 290.) Iowa Stn. Bull. 290A. An empirical test of the approximate method of calculating coef- ficients of inbreeding and relationship from livestock pedi- grees. J. Agric. Res., 456:565-69. With A. B. Chapman. Twinning, sex-ratios, and genetic variability in birth weight in sheep. I. Hered., 23:473-78. The relation of body shape of feeder steers to rate of gain, to dressing percent, and to value of dressed carcass. Tex. Stn. Bull. 471. The amount and kind of inbreeding which has occurred in the development of breeds of livestock. In: Proceedings of the Sixth International Congress of Genetics, vol. 2, pp. 123-26. Menasha, Wisc.: Brooklyn Botanic Garden. Mutton, and how it gets that way. (Review of John Hammond's Growth and the Development of Mutton Qualities in the Sheep.) J. Hered., 23:312 - 14. 1933 With W. F. Dickson. Inbreeding and the genetic history of the Ram- bouillet sheep in America. i. Hered., 24:19-33.
JAY LAURENCE LUSH 297 The use of statistical methods in animal husbandry. Proc. Am. Soc. Anim. Prod., 1932, 15-29. Linebreeding. Iowa Agric. Exp. Stn. Bull. 301. With P. B. Pearson. A linebreeding program for horse breeding. I. Hered., 24: 185 -91. The bull index problem in the light of modern genetics. I. Dairy Sci., 16:501-22. 1934 With A. L. Anderson, C. C. Culbertson, and W. E. Hammond. The reliability of some measures of productiveness in brood sows. Proc. Am. Soc. Anim. Prod., 1933:282-87. With Mogens Plum. Freshening ages of purebred cows in Iowa cow testing associations. I. Dairy Sci., 17:625-38. With G. M. Harris and E. N. Shultz. Progress report on comparison of lactation and yearly records. J. Dairy Sci., 17:737-42. With W. H. Black and A. T. Semple. Beef production and quality as influenced by crossing Brahman with Hereford and Short- horn cattle. U.S. Dep. Agric. Tech. Bull. 417. A herd of cattle bred for twenty years without new blood. I. Hered., 25:209-16. With H. O. Hetzer and C. C. Culbertson. Factors affecting birth weights of swine. Genetics, 19: 329-43. 1935 Progeny test and individual performance as indicators of an an mar's breeding value. J. Dairy Sci., 18: 1-19. The inheritance of productivity in farm live stock. V. Discussion of preceding contributions. Emp. l. Exp. Agric., 3:25-30. 1936 With I. C. Holbert and O. S. Willham. Genetic history of the Holstein-Friesian cattle in the United States. I. Hered., 27:61- 72. Genetics and animal breeding. (Review of C. Kronacher's Genetik and Tierzuchtung.~. Hered., 27:201-3. Genetic aspects of the Danish system of progeny-testing swine. Iowa Agric. Exp. Stn. Res. Bull. 204. With B. H. Thomas, C. C. Culbertson, and F. I. Beard. Variations
298 BIOGRAPHICAL MEMOIRS in the softness of lard produced in the record performance testing. Proc. Am. Soc. Anim. Prod., 1936:258-59. 1937 With Dorsa M. Yoder. A genetic history of the Brown Swiss cattle in the United States. I. Hered., 28:154-60. Identical twins in cattle. (Review of C. Kronacher's Neue Ergebnisse der Zwillingsforschung beim Rind.) At Hered., 28:415-18. Animal Breeding Plans. Ames, Iowa: The Collegiate Press. With A. E. Molln. The degree to which litter size is a constant characteristic of sows. Proc. Am. Soc. Anim. Prod., 1937:133- 37. 1938 With Earl N. Shultz. Pedigree promise and progeny test among sires proved in Iowa Cow Testing Associations. l. Dairy Sci., 21 :421-32. Teaching animal breeding. Proc. Am. Soc. Anim. Prod., 1938: 175- 80. 1939 With P. S. Shearer and C. C. Culbertson. Crossbreeding hogs for pork production. Iowa Agric. Exp. Stn. Bull. 380. With }. C. Berry. High records contrasted with unselected records and with average records as a basis for selecting cows. I. Dairy Sci., 22:607-17. With A. L. Anderson. A genetic history of Poland-China swine. I. Hered., 30: 149-56,219-24. President's address. Proc. Am. Soc. Anim. Prod., 1939:11-18. 1940 intra-sire correlations or regressions of offspring on dam as a method of estimating heritability of characteristics. Proc. Am. Soc. Anim. Prod., 1940:293-301. With D. M. Seath. "Nicking" in dairy cattle. J. Dairy Sci., 23:103- 13.
JAY LAURENCE LUSH 299 1941 With H. W. Norton III and Floyd Arnold. Effects which selection of dams may have on sire indexes. i. Dairy Sci., 24:695-721. With P. S. Shearer and C. C. Culbertson. Crossbreeding hogs? Farm Sci. Rep., January:8-11. Applications of genetics to animal breeding. Proc. Iowa Acad. Sci., 48:65 - 72. (Also in Portuguese in: Ceres EBrazil], 6~19441:44- 51.) 1942 With L. N. Hazel. The efficiency of three methods of selection. I. Hered., 33:393-99. With F. S. Straus. The heritability of butterfat production in dairy cattle. I. Dairy Sci., 25:975-82. With A. E. Molln. Litter size and weight as permanent character- istics of sows. U.S. Dep. Agric. Tech. Bull. 836. With H. H. Stonaker. Heritability of conformation in Poland-China swine as evaluated by scoring. I. Anim. Sci., 1:99-105. With Leslie E. Johnson. Repeatability of type ratings in dairy cattle. J. Dairy Sci., 25:45-56. 1943 Animal Breeding Plans, 2d ed. Ames, Iowa: The Collegiate Press. 1944 The optimum emphasis on dam's records when proving dairy sires. J. Dairy Sci., 27:937 - 51. Are better hogs coming? Farm Sci. Rep., April:3-6. 1945 Animal Breeding Plans, 3d ed. Ames, Iowa: The Collegiate Press. 1946 Chance as a cause of changes in gene frequency within pure breeds of livestock. Am. Nat., 80:318-42. With G. E. Dickerson and C. C. Culbertson. Hybrid vigor in single crosses between inbred lines of Poland-China swine. I. Anim. Sci., 5:16-24. With Roberto M. Miranda and C. C. Culbertson. Factors affecting
300 BIOGRAPHICAL MEMOIRS rate of gain and their relation to allotment of pigs for feeding trials. J. Anim. Sci., 5:243-50. Out on first record? Iowa Farm Sci., January: 6-7. With R. C. Cook. Genetics for the millions. (Review of Dunn and Dobzhansky's Heredity, Race and Society.) ]. Hered., 38:299-305. 1947 With Robert R. Shrode. The genetics of cattle. Adv. Genet.,1 :209- 61. With Raul Brequet, Jr. Heritability of amount of spotting in Hol- stein-Friesian cattle. J. Hered., 38:98-105. Family merit and individual merit as bases for selection. Am. Nat., 81 :241-61, 362-79. 1948 With G. G. Carneiro. Variations in yield of milk under the pen keeping system in Brazil. J. Dairy Sci., 31 :203-11. With W. F. Lamoreux and L. N. Hazel. The heritability of resistance to death in the fowl. Poult. Sci., 27~41:375-88. The genetics of populations. Mimeo. 381 pp. 1949 Heritability of quantitative characters in farm animals. In: Proceed- ings of the Eighth International Genetics Congress, pp. 356-75. Lund: Berlingska Boktryckeriet. Ernest W. Lindstrom, 1891-1948. J. Hered., 40~2) :44-46. The algebra of genetics. (Review of C. C. Li's An Introduction to Population Genetics.) J. Hered., 40: 156. 1950 With R. W. Touchberry. The accuracy of linear body measurements of dairy cattle. J. Dairy Sci., 33~1~:72-80. Inheritance of susceptibility to mastitis. I. Dairy Sci.,33~2~:121-25. With R. H. Nelson. The effects of mild inbreeding on a herd of Holstein-Friesian cattle. I. Dairy Sci., 33:186-93. With R. R. Shrode. Changes in milk production with age and milk- ing frequency. J. Dairy Sci., 33~5~:338-57. Review of R. A. Fisher's "The Theory of Inbreeding." Am. J. Hum. Genet., 2~1~:97-100.
JAY LAURENCE LUSH 301 With Martin Polhemus and Walter C. Rothenbuhler. Mating sys- tems in honey bees. J. Hered., 4 1 (6~: 1 5 1-55. With L. N. Hazel. Computing inbreeding and relationship coeffi- cients from punched cards. I. Hered., 41:301-6. 1951 Summary. (Symposium on selection, Chicago, November 1949.) J. Anim. Sci., 10:18 - 21. The impact of genetics on animal breeding. A general invitation review and forecast. I. Anim. Sci., 10~2~:311-21. Inbreeding and outbreeding as practiced with poultry. In: Proceed- ings of the Fourth Pacific Northwest Chicken and Turkey Breeders Roundtable, pp. 11-18. Mimeographed. Numbers of sires, dams and progeny required for reliable progeny testing. In: Proceedings of the Fourth Pacific Northwest Chicken and Turkey Breeders Roundtable, pp. 68-74. Mimeographed. 1952 With Walter R. Harvey. Genetic correlation between type and pro- duction in Jersey cattle. J. Dairy Sci., 35~3~:199-213. How dominance and gene interaction modify the effectiveness of breeding plans. In: Proceedings of the First Poultry Breeders Roundtable, pp. 15-25. Mimeographed. 1954 Breeding structure of populations. I. General considerations. In: Statistics and Mathematics in Biology, pp. 537-42. Ames: Iowa State College Press. With J. E. Legates. A selection index for fat production in dairy cattle: Utilizing the fat yields of the cow and her close relatives. I. Dairy Sci., 37~6~:744 - 53. With G. G. Carneiro. Reproductive rates and growth of purebred Brown Swiss cattle in Brazil. J. Dairy Sci., 37:1145-57. Rates of genetic changes in populations of farm animals. In: Pro- ceedings of the Ninth International Congress of Genetics, vol. 6, Suppl., pp. 589-99. Florence: Ex. Officina Typographica Flo- rent~na.
302 BIOGRAPHICAL MEMOIRS 1955 Estimates of heritability in breeding problems. In: Breeding Beef Cattle Adapted to Unfavorable Environments, King Ranch Centen- nial, pp. 113-26. Austin: University of Texas Press. With Lon D. McGilliard. Proving sires and dams. {. Dairy Sci., 38: 163-80. With D. E. Madden and L. D. McGilliard. Relations between parts of lactations and producing ability of Holstein cows. l. Dairy Sci., 38~11~: 1264-71. Gene action as related to physiological characteristics. In: Proceed- ings of the Fourth Poultry Breeders Roundtable, pp. 7-23. Mimeo- graphed. Statistics in investigations in animal production. I. Ind. Soc. Agric. Stat., 7:7-22. 1956 Dairy cattle genetics. i. Dairy Sci., 39~6~:693-94. With L. D. McGilliard. Changes in type classifications of dairy cattle. J. Dairy Sci., 39~7~: 1015-26. Answer to query about repeatability of number at a birth. Biomet- rics, 12~1~:84-88. Theoretical consequences of breeding for the heterozygote. In: Proceedings of the Fifth Poultry Breeders Roundlable, pp. 3-26. Mi- meographed. Recent advances in animal breeding. In: Proceedings of the Western Section, American Society of Animal Production, pp.20-21. Mimeo- graphed. 1957 With David E. Anderson and Doyle Chambers. Studies on bovine ocular squamous carcinoma. II. Relationship between eyelid pigmentation and occurrence of cancer eye lesions. }. Anim. Sci., 16:739-46. With David E. Anderson and Doyle Chambers. Studies on bovine ocular squamous carcinoma. III. Inheritance of eyelid pigmen- tation. I. Anim. Sci., 16: 1007-16. With C. E. Meadows. Twinning in dairy cattle and its relation to production. J. Dairy Sci., 40: 1430 - 36.
JAY LAURENCE LUSH 303 1958 With C. M. van Krosigk. Effect of inbreeding on production in Holsteins. J. Dairy Sci., 41: 105-13. Practical applications of performance testing. The Shorthorn World, 43~111:44, 318-21. Genetics in plant and animal breeding. (Translated title.) Tolvmandsbladet, 30(10) :403 -9. With D. W. Blackmore and L. D. McGilliard. Genetic relations be- tween body measurements at three ages in Holsteins. I. Dairy Sci., 41: 1045-49. With D. W. Blackmore and L. D. McGilliard. Relationships between body measurements, meat conformation, and milk production. .DairySci.,41:1050 - 56. 1959 With F. Pirchner. Genetic and environmental portions of the vari- ation among herds in butterfat production. I. Dairy Sci., 42:115-22. Making use of new knowledge about basic principles. In: Proceed- ings of the Eighth Poultry Breeders Roundlable, pp. 141-55. Mimeo- graphed. With Ivar Johansson. Zucht- und selektionsmethoden. In: Hand- buch der Tiersuchtung, vol. 2, pp. 383-473. 1960 . Improving dairy cattle by breeding. I. Current status and outlook. I. Dairy Sci., 43:702-6. 1961 With John D. Wheat. Accuracy of partial trapnest records. Poult. Sci., 40(2):399-406. Mejoramiento Animal. Publicacion Tecnica no. 6. Buenos Aires: CAFADE. Selection indexes for dairy cattle. Z. Tierz., 75~3~:249-261. Large farm animals. Germ Plasm Resour., 66:127 - 36. Der Sinn und die Bedeutung des Erblichkeitsanteiles. In: Vortrage des II Internationalen Ferienburses u.s.w., Mariensee, Germany., pp. 171-99.
304 BIOGRAPHICAL MEMOIRS 1962 With T. M. Sutherland. Effects of inbreeding on size and type in Holstein-Friesian cattle. I. Dairy Sci., 45~3~:390-95. Obituary: Edward N. Wentworth. Rec. Genet. Soc. Am.,31:20-21. 1964 With Hector A. Molinuevo. Reliability of first, second and third records for estimating the breeding value of dairy cows. J. Dairy Sci., 47~8~:890-93. 1965 With Ben Bereskin. Genetic and environmental factors in dairy sire evaluation. III. Influence of environmental and other extra- neous correlations among the daughters. J. Dairy Sci., 48:356- 60. 1967 With D. I. Kelleher and A. E. Freeman. Importance of bull x herd- year-season interaction in milk production. J. Dairy Sci., 50: 1703-7. 1968 Importance of family structure in the dairy cattle population. l. Dairy Sci., 51 :296-306. With R. M. Acharya. Genetic progress through selection in a closed herd of Indian cattle. I. Dairy Sci., 5:1059-64. 1969 Genetic unknowns in animal breeding a century after Mendel. Trans. Kans. Acad. Sci., 71:309. Pushing back the frontier of animal breeding. In: Proceedings of the Eighteenth Annual Session, National Poultry Breeders Roundtable, pp. 93-111. Mimeographed. 1971 Research in animal production: Its accomplishments and present prospects. In: Bulletin of the Swiss Association of Agricultural Grad- uates from the Eidgenossischen Technischen Hochschule. (ETH), Zu- rich, November, pp. 45-62.
JAY LAURENCE LUSH 1972 305 Early statistics at Iowa State University. In: Statistical Papers in Honor of George ~ Snedecor, pp. 211-26. Ames: Iowa State University Press. 1973 Teaching animal breeding and training graduate students. In: Pro- ceedings of the Animal Breeding and Genetics Symposium in Honor of Dr. pay L. Lush, American Society of Animal Science, American Dairy Science Association, and Poultry Science Association, Blacksburg, Virginia, July 29, 1972, pp. 78-88. 1974 Dedication of the Jay L. Lush auditorium response by Jay L. Lush. Iowa State J. Res., 48~4~:281-84.