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Why did Caftan officials pursue a so near retirement by Judith R. Goodstein to establish the schoots division of biology?

This story 0/ the genesis 0/ biology at Caltech is The biochemist Henry Borsook liked to tell excerpted/rom Millikan's School: A History of the about a conversation Thomas Hunt Morgan had California Institute of Technology (© judith R. with the physicist , a campus Goodrtein) and is reprinted here with pemziSJiol1 of the visitor in the early thirties. At a point in the publisher. W. W. Norton & C01llpcmy. Probably no conversation, Einstein supposedly asked, "What one wm in c/ better positioll to u'rite Calterh's history in hell are you doing in a place like thisl " "The thcm judith Goodstein. who, since becoming the future of biology rests in the application of the Imtittite'sjirJt archillist ilz 1968, hm had an inside methods and ideas of , , and trark on learning 11'hat went on when. In the yettrs mathematics," replied Morgan. The physicist sinre. she has built the arrhilles, nou' housed in persisted. "Do you think you will ever be able to expanded ncu,/acilities in the Beckman Imtitute. into a explain in terms of chemistry or physics so notable in bistory-cmd ctlso knOUT important a biological phenomenon as first love)" where all the bodies are bllried Research for the book. "What did you say to that one)" Borsook asked u'hirh apjJropriately apjJearJ in time j()r Calteeh's Morgan afterward. "I tried to explain something Ce7Ztennial, cl'as supported by the HClples Foundcttion. about the connection between sense organs and In addition to her post as arciJillirt. Goodstein has cdw the and ." "You didn't believe Thomas Hunt Morgan, with , at Columbia been a faculty ass(){iate since 1982 and registrar since that yourself, did you)" Borsook asked. "No," University in 1917. J 989. She ectrned her BA frail! Brooklyn College in said Morgan, "but I had to say something to J 960 and PhD from the Unillersity o/VvmhillgtoJl in him." 1969. Millikan's School u'ill be cte'ailable in What he "had to say" says a lot about his bookstores in October or lIlay be ordered from the plans, however. Thomas Hunt Morgan was 62 publisher lcith the coupon Oil the back cot'er. when he came to Caltech in 1928. By then, he had earned a worldwide reputation as a remark­ The establishment of a Department of able teacher, a clear writer, and an impressive Biology, rather than the traditional depart­ researcher. In 1933, he would win the Nobel ments of and , calls for a word Prize for his discovery of the chromosomal of explanation. It is with a desire to lay mechanism by which character traits are passed emphasis on the fundamental principles on from parent to offspring through the interac­ underlying the processes in and tion of . that an effort will be made to bring together, in a single group, men whose All of that work had been accomplished in one common interests are in the discovery of the room at that held a bunch unity of the phenomena of living of bananas hanging in the corner and eight desks rather than in the investigation of their crammed into a space measuring 16 feet by 23 manifold diversities. feet. In the room, as it was known, Morgan -THOMAS HUNT MORGAN, 1927 had elevated the lowly fruit fly, the

Engineering & Science/Summer 1991 13 Contributing to the squalor was Morgans 0/ squashing his flies (after he'd finished counting them) on his 17lelanogaster, into the most famous experimental mold it attracted. Along one wall stood a in the world. kitchen table used by the student hired to wash counting plate, Why did Cal tech officials pursue a biologist so bottles and prepare fly food. This area of the which he left near retirement to establish the school's division room was Morgan's only concession to hygiene. unwashed on his of biology? The answer to the question begins Bridges, an unremitting tinkerer, sat at a desk with the ways and means of Morgan's Drosophila covered with odd-looking pieces of apparatus he desk. group at Columbia. had made from items at hand. Capable of long According to the Russian Theodo­ periods of routine work and intense "fits and sius Dobzhansky, Morgan ran the fly room by his SpurtS" of ingenuity, Bridges gradually over­ own rules. Traveling on a Rockefeller-financed hauled Morgan's primitive laboratory techniques: International Education Board fellowship, he designed a binocular microscope (most Dobzhansky in 1927 arrived in from workers used a hand lens to examine flies; Leningrad thinking that Morgan was "just next Morgan used a jeweler's loupe), invented new to God" and his laboratory "close to Heaven." To ways to etherize flies, developed new incubators, his dismay, he found what he called "a very small, improved culture bottles, and whipped up poorly equipped, and positively filthy" labora­ alternative foods for flies. tory, run by a man obsessed by "pathological Bridges had his faults, but jealousy, according stinginess." The Morgan operation made to Morgan, was not one of them: "In fact, one of Dobzhansky's laboratory facilities back home in his most admirable traits was his freedom from Russia look very good in comparison. Morgan's priority claims of any kind." Morgan first met longtime co-workers Calvin B. Bridges and Bridges in 1909 when he took Morgan's courses Alfred H. Sturtevant sat and worked in the same in general biology and . Hoping to room, along with graduate students, postdoctoral find research work, Bridges put himself through fellows, like Dobzhansky, and assorted visitors, Columbia on scholarships and odd jobs. Morgan ranging from Yoshitaka Imai, a Japanese geneti­ hired him in 1910 to wash glassware, but gave cist, to Alexander Weinstein, a recent fly-room him a desk and promoted him to the job of PhD. Often, all the desks were occupied, in­ breeding flies and looking for after cluding the two reserved for guests. Bridges spotted a fly with bright eyes in a dirty Cleanliness was unknown here. The workers bottle. Bridges excelled at finding new mutants, competed for space with cockroaches that which he "immediately announced." This skill reproduced in awesome numbers. Contributing (Sturtevant insisted he "had the best 'eye' for new to the squalor was Morgan's habit of squashing types") paid off in 1916, when he published, in his flies (after he'd finished counting them) on his the first issue of , a detailed paper dealing counting plate, which he left unwashed on his with flies that had extra and missing desk. As the pile oflifeless flies grew, so did the . Not only could Bridges explain

14 & Science/Summer 1991 great many advantages, particularly for a foreign guest. You can ask anyone a question you wish to enlighten yourself on any problem which Left: Morgan's fly­ arises. You also listen to the conversation room crew didn't flinch from eating between the people. As far as training is con­ among their speci­ cerned, nothing better can really be imagined." mens_ Here a lunch Jammed with people and paraphernalia, Morgan's celebrates the return of Sturtevant (front laboratory, in short, was an ideal training ground left, holding beer for budding experimental , good for bottle and cigar, from everything from selecting projects on which to the Army in 1919_ Clockwise around the base PhD research to testing new techniques and table from Pithecan­ analyzing experimental data. thropos wearing Sturtevant tells a similar story. "Everybody Sturtevant's old uni­ form are Muller, did his own experiments with little or no super­ Morgan, Lutz, Mohr, vision," he wrote on one occasion, "but each new Huettner, Schrader, result was freely discussed by the group." Anderson, Weinstein, Dellinger, and Bridges Morgan's Drosophila group did not go in for (with book,. organized coffee breaks, nor did it set aside a Right: , certain time of the day for laboratory discussion. who began his career with Morgan as a bot­ "Instead," recalled Sturtevant, "we discussed, tle washer, was pro­ planned, and argued-all day every day." He moted to the job of added, ''I've sometimes wondered how any work breeding flies, and given a desk, which these exceptions; he provided convincing proof got done, with the amount of talk that went on." he covered with hand­ of the theory of . Bridges But Morgan did have one cardinal rule: you had made pieces of appa­ delighted in building up and studying the to pick your own research topic. ratus as well as bottles of flies. Drosophila stocks and mapping the position of To do otherwise could be academically fatal, as genes in each chromosome. He "was so one aspiring Drosophila geneticist, Edgar Alten­ good at this that he contributed many more burg, discovered. Having been given desk space mutants than did the rest of us," Sturtevant in the fly room, Altenburg asked Morgan to once admitted. suggest a fruit fly problem for graduate work. Sturtevant owed his desk in Morgan's labora­ Close by Morgan's office was the aquarium room. tory to a childhood passion for recording the He took Altenburg there, dipped his finger in a pedigrees of horses. A book on Mendelism that tank of stagnant water, and held it up to the he read as a college sophomore opened new light. 'There are a lot of Daphnia in here," he worlds, he later wrote, "for I could see that the said to Altenburg. "Why don't you work on principles could be applied to the inheritance of them?" Humiliated by the experience, Alten­ colors iQ the horses whose pedigrees I knew so burg quickly switched to genetics. well." He wrote up an account of his findings Dobzhansky nearly made the same mistake. and submitted it to Morgan, his biology teacher. Once unpacked, he wasted no time in asking Much impressed, Morgan urged the young man Morgan "to suggest a topic." "After all I was to publish the account, which he did in due coming from afar, and although I knew what Jammedwith course. Sturtevant always believed this was the they had published earlier, I didn't know what people and para­ reason why in the fall of 1910 Morgan invited they were doing at the time, less still ... what phernalia) Mor­ him into the laboratory and gave him a desk and they were planning to do" in the future, he said, some DroJ()philcl to work on. By that time, adding, "I did not know how foolish that was." gan}s laboratory} Sturtevant knew for sure that he wanted to do At first, Morgan brushed him off with a joke. in short} was an genetics. When Dobzhansky asked him again for some­ Sturtevant was the bookish one. Piles of books thing to read, "the Boss" reached into his desk, ideal training and reprints, stacked high, covered his desk. In took out a reprint dealing with the effects of ground for bud­ the course of cleaning the room one summer, so temperature on the development of Drosophila ding experimental the story goes, a workman found it necessary to (a subject of no interest to the Russian biologist), rearrange some of Sturtevant's papers, uncovering handed it over to the newcomer, and turned biologists. a shriveled mouse. away. A zoologist by training and a geneticist It didn't take Dobzhansky long to discover with an expert's knowledge of the natural varia­ what made the fly room tick. He later told an tions in lady beetles and with a keen desire to interviewer, "So this one room had six people study problems of , Dobzhansky quickly working in it, a situation which doubtless had a made his peace with Morgan's managerial style.

Engineering & Science/Summer 1991 15 Morgan "thought that everybody should work on was coined only in 19(9) were parts of chromo­ the problem which he sees fit," and Dobzhansky somes, unless they had evidence rooted not in knew he was "perfectly capable of choosing" what statistical studies of monastery peas but rather in he wanted to do in Morgan's laboratory. It was a observable laboratory phenomena. Thus, the perfect Ii t. question facing Morgan and like-minded col­ Morgan was a southerner, born in Lexington, leagues was twofold: First, how far could , in 1866. His maternal great-grandfa­ Mendel's work be taken as an authentic descrip­ ther was , author of "The Star­ tion of heredity in organisms? Second, how Spangled Banner," and his father's brother, John correct was the theory that chromosomes were Hunt Morgan, had been a notorious Confederate indeed the physical basis of inheritance? The general. Preferring to politics, validity of the chromosome theory took Calvin Thomas Hunt Morgan in his youth combed the Bridges-first Morgan's student and later his backwoods and byways of rural Kentucky and collaborator-only a few years (1914-16) to western , collecting fauna and fossils. establish. The evidence needed to convince One summer, he earned his keep working for the Morgan that Mendel's "genes" were indeed U.S. Geological Survey, tracing coal seams. After carried on chromosomes took longer to graduating in 1886 with a BS in zoology from accumulate. the , Morgan spent the Of this period (1910-11) in Morgan's scientific summer months working in a marine biological life, the biologist John Moore has remarked, laboratory at Annisquam, Massachusetts, and "Whereas it was difficult for Morgan to accept then enrolled as a graduate student at Johns the data of others in suggesting that genes are Hopkins, earning his PhD-his dissertation parts of chromosomes, it was not nearly so involved studying different species of sea spi­ difficult whea his own data showed the same ders-in 1890. By the time Morgan joined the thing." In the end, Morgan's studies with Columbia University faculty, in 1904, he was Drosophila convinced him of the necessity of known far and wide for his work in experimental associating specific hereditary characteristics with embryology and . specific chromosomes. He equated Mendel's But the studies that brought lasting fame to elements of heredity with invisible genes at The studies that Morgan were those connected with Drosophila. known locations in visible chromosomes and in brought lasting He had begun breeding fruit flies in 1908 in an the process created a new science of genetics. effort to determine what role-if any--chromo­ Fruit flies have been called the geneticist's best fame to Morgan somes played in the transmission of physical friend. They reach maturity quickly, reproduce were those con­ traits from one generation to the next. themselves frequently, and are inexpensive to Morgan began his research with Drosophila just rear. Shortly after he had begun research on nected with as biologists were beginning to appreciate for the Drosophila, in the autumn of 1910, Morgan Drosophila. He first time the long-neglected findings of the recruited Bridges and Srurtevant, both then still 19th-century Austrian monk . undergraduates, to help with the fly work. Two had begun breed­ Mendel's genetic experiments on plant hybrids, years later, he brought a graduate student, the ing fruit flies in published as a short report in 1866, led him to physiologist Hermann]. Muller, into the fold, 1908 in an effort conclude that traits in garden peas such as seed an association that was to have less happy shape, pod color, and plant-stem length were consequences. Muller was a good scientific to determine what determined by fundamental units of inheritance, choice-his work was clearly outstanding-but role. . . chromo­ which he called "elements." Alternative forms he and Morgan made a poor match in tempera­ existed for every hereditary trait as well: round ment. From the first, Muller's relations with the somes played in seeds and wrinkled; green pods and yellow; short group, and with Morgan in particular, were the transmission stems and long-one of which always stood out strained. Part of the problem was the pecking ofphysical traits decisively in the pea plant. Today every school­ order. Scientific decorum mattered a great deal child knows how Mendel, toiling alone in his to Muller, who came to feel that others in the fly from one genera­ monastery's vegetable patch, theorized the room got credit for his ideas and experimental tion to the next. existence of dominant and recessive traits, work. In conversations with the psychologist through repeated crossings of innumerable pea Anne Roe in the 1940s, Sturtevant testified that plants. Yet Mendel's work was ignored and "Muller was a very essential part of that group," forgotten until 1900, when it was rediscovered adding, "We didn't see eye to eye but I got a lot independently by three botanists. out of him." By the time Morgan moved his Nevertheless, many researchers, Morgan laboratory, Drosophila stocks, and research group included, were initially reluctant to accept the to Pasadena, in 1928, Muller had long since left notion that Mendel's "elements" (the term Columbia and launched his own research team at

16 Engineering & Science/Summer 1991 {(Our program, when we get it going} should speak/or ztse. If.... n

On October 21, 1933, a luncheon at the Athenaeum hosted by Robert Millikan in honor of distinguished visitors Guglielmo Marconi, inventor of the wireless, and his wife (third and fourth from right) also turned into a celebration of Morgan's , which had been the University of Texas in Austin, where he be­ make other significant discoveries both at announced the pre­ came the first geneticist to demonstrate that x Columbia and at Caltech, but none came close vious day_ A pleased­ rays cause . Like Morgan, Muller to matching the thrill of his "first job on looking Morgan stands second from eventually won the Nobel Prize, but the personal Drosophila. " right, next to Mrs. rift between the two was never entirely repaired. Two years later, Morgan, Sturtevant, Muller, Millikan. Millikan, The discovery that genes are arranged in a and Bridges joined forces to produce the first (second from left) is flanked by Mr. and single line in chromosomes like beads strung textbook on Drosophila genetics, which they Mrs. Allen Balch, who, together on a loose necklace was made by entitled The Mechanism of Mendelian Heredity. A in addition to financ­ Sturtevant in 1911. At the time, he and Morgan landmark in the history of 20th-century biology, ing the building of the Athenaeum, also gave had been talking about the meaning behind some the volume quickly became the bible of the new $1 million toward the diagrams by H. E. Castle of coat colors in rabbits. science of genetics. In the hands of Morgan and biology laboratory The diagrams, they decided, were meant to be a his co-workers, the genetics of Drosophila in­ that helped bring Morgan to Pasadena representation of the spatial relationships of the volved a rigorous and experimental search for the (although Kerckhoff genes on a given chromosome. How nice it secrets oflife that lay sprinkled within the got his name on it). would be~to figure out the geometrical relation­ chromosomes of a tiny fly. ship between genes and chromosomes! "I think I By this time, Morgan had largely turned his can do it," Sturtevant told Morgan all of a attention elsewhere. He left the day-to-day sudden. "I suddenly realized," he recalled 50 operations of the fly room in the care of his years later, students, who were technical virtuosi of the first that the variations in strength of linkage, order. A great synthesizer, Morgan distilled their already attributed by Morgan to differences in findings, popularized their work, and shouldered the spatial separation of the genes, offered the the responsibility for bringing their results to a possibility of determining sequences in the wide, frequently nonscientific audience. In fact, linear dimension of a chromosome. I went by the time he left Columbia in 1928, Morgan home and spent most of the (to the could no longer follow in detail what his younger neglect of my undergraduate homework) in colleagues, Bridges and Sturtevant in particular, producing the first chromosome map, which were doing. included the sex-linked genes y, w, m, and r, This happens to , even to the best of in the order and approximately the relative them. And it's often sad to observe. In Morgan's spacing that they still appear on the standard maps. case, however, distance worked to his advantage. As one geneticist familiar with Morgan's working He published his results in 1913. Sturtevant habits pur it, "For Morgan himself the Drosophila later told an interviewer that the discovery of the work was only one aspect of a biologist's search­ linear arrangement was the most exciting thing ing." Professor of experimental zoology at he had ever done scientifically. He went on to Columbia for 24 years, Morgan in 1928 was still

Engineering & Science/Summer 1991 17 In 1931 Caltech's Division of Biology included (front row, from left): H. Borsook, H. Dolk, H. Sims, A. H. Sturtevant, S. Emer· son, H. Huffman, T. H. Morgan; (back row): H. Schott, C. Burn· ham, W. Lammerts, K. Lindstrom·Lang, E. Ellis, G. Keighley, J. Bonner, A. Tyler, searching, as is plain from the following lines he work in biolegy in Pasadena. "The best chance" G. W. Beadle, and J. wrote to , shortly after for success, he indicated, would be "to put some Schultz. Missing from accepting the Caltech job: physicists in the biological laboratory, and some photo are E. Ander· biologists in the physical laboratory." son, K. Belar, ... I am writing to you something of the ideas Morgan's prophetic remarks set the tone of T. Dobzhansky, R. that are shaping themselves in my about Emerson, and K. biology at Caltech for the next half century. It Thimann. the organization of our biological work. Would it not be a good plan to think in terms was, for example, Caltech's physicist-turned­ of "The Biological Laboratories," rather than biologist, Max Delbruck, himself winner of the of a "Biological Department." This would Nobel Prize in 1969, who helped lay the founda­ allow greater freedom in giving each group an tions of modern and the brave independent footing and allow greater flexi­ new world that we've only begun to glimpse. bility in the future. As I have intimated to Morgan kept his word to Hale "to get together you, I think, I have no ambition to "boss the the best men available," starting with three he job,"' but rather to get together the best men knew-Sturtevant, Bridges, and Dobzhansky. available, to settle down to my own work, and He also recruited as teaching fellows three then do all I can to coordinate and help matters forward along constructive lines. graduate students ftom Columbia, including Our program, when we get it going, should Albert Titlebaum. From the University of speak for itself. ... And, while I am anxious to Michigan, he plucked Ernest Anderson and emphasize the dynamic or physiological Sterling Emerson, both PhDs in plant genetics, character of the work, I shall try to avoid the but well versed in the genetics of animals as well. criticism that we are leaving the older and less Anderson, 37, came as an associate professor of important sides of biology in the background. genetics; Emerson, 29, as an assistant professor. This can best be done, perhaps, if we point out Another geneticist from Columbia, Alexander that we are not so much attempting to dupli­ Weinstein, did not get to Caltech-in a way that cate work that is being done well elsewhere, as says much about the school's early ways. in furnishing opportunities for the more ad­ Weinstein, Morgan told Millikan in the vanced and less well developed lines of modern research. spring of 1928, had been working in the fly room and had just successfully repeated Muller's use of Morgan's days of "the fly room" mentality x rays to induce mutations. If appointed to Cal­ were behind him. "Only through an exact tech, he would continue the work in Pasadena knowledge of the chemical and physical changes and teach the introductory course in biology. taking place in development can we hope to raise Morgan was proposing to make Weinstein an the study of development to an exact science," assistant, perhaps even an associate, professor, at Morgan told Caltech's elders in 1927, shortly an annual starting salary of$3,500. Emerson's after they had approached him about organizing starting salary was $3,800 and Anderson's

18 Engineering & Science/Summer 1991 $4,000. As professor of genetics, Sturtevant received $6,500, placing him near the top end of the Cal tech pay scale; Morgan was hoping to raise UT he best chance Sturtevant's salary to $7,500. (As head of the new department of biology, Morgan himself {for success made $10,000, the same as Millikan and Noyes.) would be} to put A "with distinct literary ability," broadly trained in biology, fluent in mathematics some physicists in and physical chemistry, Weinstein ("a fine type, the biological not aggressive") struck Morgan as the right man laboratory and for the Cal tech job. "I have hesitated a long time before bringing his name forward," Morgan ad­ some biologists in mitted in his letter to the school's head, "but I the physical think for the position proposed he is the most suitable man at present available." laboratory. JJ No one on the faculty, save for members of the National Academy of , replied Millikan, made more than $7,000 a year. In Sturtevant's ca~e, Millikan agreed, Morgan might have to pay that much or more to get him, but he might first try less expensive inducements-paying traveling costs to meetings back east or moving expenses. Weinstein's case was scarcely different. Millikan had at least "three brilliant young men" in the assistant professor ranks, all making between $3,000 and $3,300. Offer him $3,500, Millikan counseled, but then "give him a chance to match his pace to an associate professorship with these other men of about his age." In any case, he left all decisions about rank and salary in Morgan's hands. Morgan did not change Sturtevant's starting salary. He offered Weinstein $3,500 as an assistant professor, which the seasoned fly-room veteran refused, pointing out that Emerson, barely out of graduate school, was making more and that Anderson and Sturtevant, who were about his age, were getting higher faculty positions. Morgan bristled. "I ... consider the matter finished, as I do not think we want to have a man who makes points like that," he informed Millikan by letter that May. Too cheeky perhaps for Morgan's taste, Weinstein went on to teach genetics at Minnesota, then branched out into zoology and the history of sci- ence at Johns Hopkins. He later taught physics at City College Morgan and Arie of New York and eventually wound up at Haagen·Smit, who Harvard. later pioneered studies in the Another possible appointment in biology was chemical of , a prominent biophysicist at smog and its Johns Hopkins. Michaelis, however, had several sources, in 1938. Morgan had per­ strikes against him, according to Caltech's new suaded Haagen. biology head. One was his apparently pro­ Smit to come to nounced ethnicity. In the same letter of 1928 to Caltech as associ· ate professor of bio· Millikan recounting his dealings with Weinstein, organic chemistry Morgan lamented that Michaelis already had the previous year. "collected about himself a few young Jews." "He

Engineering & Science/Summer 1991 19 himself is markedly Semitic," added Morgan. "I the Rockefeller Foundation has given him money have my doubts whether we should want to start to secure the services of a physiologist. He is under all these conditions, and shall make no combing England and the Scandinavian countries moves." Morgan recommended against hiring to find one who is not Jewish, if possible." The Michaelis. Fifty years later, Leonor Michaelis's informant added, "From the English reception of daughter read the discussion about her father in this announcement, I am inclined to believe that the Caltech archives and said "it was shocking" to he will have difficulty in finding a first-rate learn that the call never came because, as she put Englishman who will be willing to go to Pasade­ it, "he was aJew." People, scientists included, na." Indeed, Morgan was unable to recruit rarely take the time to write shocking letters any anyone in England or Scotland. Just before more; they simply talk on the telephone instead. sailing home, he hired a Dutchman, C. A. G. Dobzhansky, who worked side by side with Wiersma, who evidently had the right pedigree. Morgan tor many years, described him as a biol­ According to a Rockefeller official in Amsterdam, ogist with a razor-sharp mind, "a man of wide Morgan "gave somewhat the impression of being I n the genetics education which should have made him very 'desperate. '" laboratory) the broad, but curiously enough, did not." "In many In the fall of 1928, the founding members of ways," his Caltech colleague once recalled, "he Caltech's biology division assembled for the first atmosphere of the was a very contradictory person." He'd had a time in Pasadena. Traveling by the Santa Fe original fly room number of Jewish co-workers at Columbia­ Railway, Morgan and his wife, Lilian, also a was soon re­ Weinstein, Muller (he claimed one Jewish biologist, left for Pasadena on September 6, ancestor), Tyler (born Titlebaum, he changed it 1928, after spending the summer, as always, at created after moving to Pasadena)-and he brought the Marine Biological Laboratory in Woods Hole, many others to Caltech, including Henry Massachusetts, Dobzhansky, his wife, and Miss Borsook, Jack Schultz, and Norman Horowitz, Wallace, Morgan's illustrator and secretary, left who many years later said, Woods Hole and joined him in Pasadena several weeks later, carrying among them "the sacred The question of Morgan's alleged anti­ flame," the Drosophila stocks. Morgan met them Semitism bothers me. I was closer to him at the train station. Bridges rurned up soon than most graduate students during 1936-39, afterward. Emerson had gone fishing with his because he and Tyler and I spent every father-in-law in Canada meanwhile and arrived at weekend at the marine station. I never noticed any anti-Semitism whatsoever on his the end of October, a month after the academic part. On the contrary, he was always nice to term had begun. Srurtevant's wife was expecting me, and I have always believed that it was he a baby, and Sturtevant "thought she had a right who got me a National Research Council to be born in the East." They arrived two months Fellowship when I finished my PhD in 1939. late. Anderson was completing a research trip to Berlin and arrived even later, close to Christmas. "But time and again he would make, especial­ Finding the new biology building unfinished, ly when irritated, anti-Semitic remarks of the Morgan and his group set up a makeshift labora­ most crude sort," remembered Dobzhansky. tory in the chemistry building, in Arthur Noyes's Morgan had a reputation for making outra­ office. By the time classes began, two rooms in geous remarks, for teasing those with different Caltech's new KerckhoffLaboratories of the beliefs. "But he was never mean," insisted Biological Sciences were ready for occupancy. Sturtevant. Robert Millikan was often the butt Tucked away by itself in the northwest corner of of Morgan's gibes, for, unlike the atheist Morgan, the campus, the building was a brisk five-minute Millikan was a pious Protestant. But Morgan's walk to the physics, chemistry, and engineering penchant for saying the wrong thing eventually laboratories across the quadrangle. A student caused an international flap in scientific circles. who took courses there in 1930 recalls that the In 1934, Morgan went abroad, partly to pick up building "was connected by a boardwalk to the his Nobel Prize in Stockholm, partly to recruit rest of the campus. In the winter, the territory new staff members. As Morgan had told a between Gates and Kerckhoff became a sea of Rockefeller Foundation official beforehand, he mud, known generally on the campus as Lake wanted "to look over the ground at first hand and Kerckhoff. " make sure that the men we have in mind are the Caltech officials had promised the biologists kind we are looking for." While in London, that they would not have to teach any courses Morgan attended an elegant reception hosted by that first year. But under pressure from the the Royal Society. "He has announced to all who undergraduates, they offered one-in beginning will listen," an eyewitness later reported, "that biology-in the spring term. Morgan and

20 Engineering & Science/Summer 1991 Sturtevant divided up the lectures, while Ander­ meant that a piece of the third chromosome had Above: Russian son, Emerson, and Sturtevant ran the laboratory become attached to the tiny fourth." The ancient geneticist Theodo· associated with it. Partial to Darwin, Morgan in of -direct evidence of the serial sius Dobzhansky, his homework assignments often asked students order of genes on chromosomes-stared Dob­ who joined Morgan at Columbia in 1927 and to read a portion of his masterpiece, The Origin of zhansky in the face. "I don't remember whether I came with him to SpecieJ, and to write a report on it. emitted a loud yell," he later said. Caltech, enjoyed In the genetics laboratory, the atmosphere of By spring 1929, Dobzhansky had produced camping and often used the pursuit of the original fly room was soon re-created. A long the first cytological map of the fly's long, rodlike biological specimens bench stood in front of the two windows. Dob­ third chromosome. To his joy, when he com­ as an excuse to zhansky and Sturtevant sat at opposite ends of it pared the linkage map, which summarized pursue his favorite pastime. looking at their flies-Dobzhansky on the left, statistical data based on many genetic experi­ Above right: Kerck· Sturtevant on the right. "The students sat in ments, to the cytological map, he discovered that hoff Laboratory was between and listened to the wise conversation and the two maps agreed with each other. The ability still under construe· to tion when Morgan, contributed it when they could," one former to predict the inheritance of certain characteris­ his researchers, and student remembers. tics, Morgan once said, justified the construction his flies arrived in Intrigued by Muller's x-ray work, Dobzhansky of genetic-linkage maps, "even if there were no 1928. The sea of mud in front of it had used his time at Woods Hole during the other facts concerning the location of the genes." became known as summer to irradiate flies. He spent fall and Dobzhansky's work in Kerckhoff Laboratory Lake Kerckhoff. winter in Kerckhoff studying the chromosomal offered irrefutable, direct evidence of the correct­ aberrations caused by the x rays and arranging ness of Morgan's classical theory. them on a chart, using genes as markers. "Just Meanwhile, time was running out on Dob­ what I expected to see in chromosomes, I don't zhansky's postgraduate fellowship. Morgan had remember," Dobzhansky later told an interview­ succeeded in getting him a six-month extension, er, but he decided one day to look under a mi­ which meant he had to wind up his research and croscope at the rearrangements he had projected return home to the Soviet Union at the end of would be observed between the fly's third and June 1929. One day, Morgan walked into the fourth chromosomes. Practiced in dissecting genetics laboratory and, according to Dobzhan­ beetles, Dobzhansky removed the ovaries of a sky, "asked the question, 'Dobzhansky'-or young female fly, embedded them in paraffin, rather, he called me to the end of his days, he and sectioned and stained them. It was a long, could not pronounce this name, which of course I tedious process. He looked through the eyepiece. don't blame him, it's a devil of a name-he called "Suddenly I saw an incredible thing," he later me 'Dobershansky' 'Would you like to join our recalled, "namely, I saw a chromosomal plate staff as assistant professor?'" It took him no time which had just one little dot ... and a chromo­ at all to answer yes. some never seen before, a long rod, which clearly In educational matters, Morgan did not be-

Engineering & Science/Summer 1991 21 Right: Morgan and Albert Tyler, then a grad student and later a member of the faculty, examine fertilized eggs of marine organisms at the Corona del Mar marine laboratory in 1931. Left: Also in the early thirties, Calvin Bridges points out a mutant on his "totem pole," a map classify­ ing mutations in a particular chromo­ some as "best, good, poor, dead" for the purpose of doing genetic experiments.

lieve in graduate courses; he believed in reading. ing the number of visible bands of these chromo­ But even in Morgan's day, graduate students took somes. He went on to produce a series of courses, whether required or not. Seminars drawings that are still consulted by fly geneti­ abounded, including the general biology seminar cists. "Bridges's map," the Drosophila whiz each Tuesday night, which Morgan always Edward Lewis remarked more than 50 years later, attended (and at which he introduced the "is still a masterpiece." speaker). Keen competition existed between the Caltech In one of these seminars, in 1933, a visiting geneticists and Painter's research group at the biologist reviewed a paper by two German University of Texas. In 1934, Painter wrote an researchers on the salivary-gland chromosomes of indignant letter to the editor of the journal flies known as Bibio, or March flies, so named Geneticf. According to Painter, the two groups because they are commonly abundant in spring. were "in a sense competing," and the Texas group The Germans had observed in the nuclei of had already hit two "home runs": Muller's dis­ the larval salivary glands rope-like , covery of x-ray induced mutations, and Painter's which they correctly interpreted as "giant own work on the salivary glands. But now he chromosomes." A number of geneticists were in was upset that Bridges had reviewed his salivary­ the audience that day, Morgan and Bridges gland manuscript. (Did Morgan have a hand in included, but they did not get excited by the this? he asked.) Worse still, Bridges's salivary­ report. Their attitude changed overnight when gland work was now "making a splurge in the Theophilus Painter, a geneticist at Texas, drew newspapers"-the New York Times had sent a and published the first map of these chromo­ reporter to Spring Harbor to cover Bridges's somes for Drosophila and pointed out how the talk on his salivary-gland research-while his, banding pattern could be used to study the break Painter's, contributions had "been belittled." points of any chromosomal rearrangements. Painter's public howl strikes a familiar chord. It As Dobzhansky tells the story, Bridges showed is not uncommon for scientists to believe that up in his laboratory one morning just thereafter their work is underappreciated. and said, "Dobzhansky, show me the salivary Painter's most pressing complaint, however, glands." Although he probably knew more about concerned a Science News Service press release Drosophila than anyone else, Bridges had no about Bridges's work, which the magazine's experience in dissecting larval fruit flies. Indeed, editor had sent ro Painter. From reading the he had never even seen the salivary glands. marked-up copy, Painter could see that Bridges Dobzhansky dissected a larva and showed the had corrected his estimate of the size of the results to his visitor. And Bridges jumped into salivary-gland chromosomes and that the new the study of these giant chromosomes with a figures now agreed with Painter's-which had vengeance. He set about identifying and extend- not yet been published. He felt that Bridges had

22 Engineering & Science/Summer 1991 taken advantage of the situation. He wrote in his in the same journal. Muller prefaced his techni­ letter, "I do not intend to reflect in any way on cal remarks by calling "the attention of American Dr. Bridges. On the other hand, the men in ... readers ... ro the fact that essentially the same [our} laboratory are unwilling to allow competi­ findings and interpretation" (presented in tors in our field to enjoy the privilege of examin­ Bridges's paper) had already appeared in print in ing our work a year prior to publication when we the Soviet Union under Muller's name and that of have no opportunity to see theirs." his two Soviet co-workers. In truth, rivalry is the "Weare not interested in home runs," Morgan handmaiden of science, and the quest to be first is replied, after reading a copy of Painter's letter. a motivating force and a powerful to As far as he, the dean of American biologists, was creative work. Good scientists are nearly always concerned, the two research groups were "cooper­ keen competitors. ating," not competing. Far from admitting any Closer to home, Morgan made a magnanimous wrongdoing, Morgan defended his group's honor, gesture when in October 1933 he received the but held out a peace offering: articles submitted Nobel Prize in and . From by Cal tech would now be sent to Austin before the proceeds of the prize, some $40,000, he de­ publication. In a separate note to the editor, D. ducted his traveling expenses to Stockholm and F. Jones, Morgan blamed Painter's "ourburst" on back, and divided the rest of the cash prize three Muller, noting, "[His} attitude has always been ways: one-third of the money went to his own antagonistic to us ... although he has generally children, one-third went to Bridges's children, managed to keep this under cover and we have and one-third to Sturtevant's. consistently ignored it, treating him in the most Biology at Caltech in the thirties was special friendly way, because we regarded his attitude as because of its emphasis on genetics, the essential wrong and inexcusable." It is reported that science for the future of biology. Caltech had Painter later mellowed in his view. staked out its claim, and that was Morgan's Painter's story bears telling because of Muller's doing. At other first-class universities, including experience several years later. At Cal tech , Mor­ Harvard and Princeton, genetics took a backseat There was, Mor­ gan had laid down the rule that getting papers to other branches of biology, such as comparative gan maintained, published was an individual's responsibility, not , embryology, and physiology. Cal tech the department's. Bur as Sterling Emerson once was unconventional not only in its choice of a rational, physi­ freely admitted, Morgan's group had no trouble discipline but also in its methods of discovery. co-chemical expla­ getting him to submit their papers to Science and For one thing, Morgan's approach was completely the American Naturalist, edited by J. McKeen experimental; no courses in descriptive biology nation behind Cattell, a personal friend. When Morgan sub­ existed. According to people in the department all biological mitted a colleague's paper, recalled Emerson, "it then, Morgan "said that as long as he had any say would come out in the next issue. It might be in this matter, there would never be a class in phenomena. any time if you sent it in." Being friends of or in ." friends paid off, as Calvin Bridges discovered. In setting the intellectual tone for the new In 1936, Bridges was preparing a paper for division, Morgan was guided by his instincts, and Drosophila on the Bar gene, a spontaneously by an outlook much broader than even his best mutating gene that reduces the size of the fly's pupils could muster. If he had an ideology, it eye. As Bridges drew near the end, word reached "was that genetics was the root to finding out the Caltech group that Muller was also working how life works," as Dobzhansky put it. Stur­ on this gene. Donald Poulson, a graduate stu­ tevant said it in a slightly different way: "Mor­ dent in the lab at the time, told an interviewer in gan's objective in biology was the development of 1978 what he remembered: "I don't know mechanistic interpretations. Anything teleologi­ whether I should say anything about this, but I cal was sure to arouse his antagonism." There think it's. current now-Dobzhansky had had a was, Morgan maintained, a rational, physico­ letter from Russia, from one of his friends, which chemical explanation behind all biological said, 'Muller has solved the Bar story.''' phenomena. Morgan took matters in hand. Bridges's paper The Caltech plant physiologist and biochemist was submitted on February 21, to Science! and was James Bonner remembers that when he was a published a week later. The habitually frugal graduate student in biology, in the early thirties, Morgan, it seems, had wired the entire paper to a fellow graduate student in physics, "Willy the journal's editor. So much for Morgan's Fowler, who will of course deny this," said to "cooperation" among Drosophila groups. Three him, '''Biology? How are you ever going to make months later, a special article by Muller on his a science out of that?'" Morgan came to Caltech own cytological analysis of the Bar gene appeared to answer that question.

Engineering & Science/Summer 1991 23