The Early History of Biology at the Institute
'I owe a similar debt of gratitude to Professor Edward W. Claypole, whose Latin marginalia inscribed upon our notebooks in biology and the like, would now, I suppose, seem to be relics of an order of things that is no more.'—from the preface to The Birds of the Latin Poets by Ernest W. Martin, Associate Professor of Greek, Stanford University, 1914.
The California Institute of Technology has a prehistory, a period from 1891 to 1920 when it was a local vocational school called Throop University, then Throop Polytechnic Institute, and then, as aspirations for it rose, Throop College of Technology. It was initially at a small campus north of downtown Pasadena (on Chestnut Street between Fair Oaks and Raymond), but since 1910 has been at the present location south and east of downtown. The first listed teacher of biology at Throop (which included a grammar school, high school, and college) was Alfred J. McClatchie, A.B., listed as the only biologist on the faculty in the catalogs from 1893 through the 1895-96 school year. In 1895 Ernest Bryant Hoag, B.S. (from Northwestern in 1892), A.B. (from Stanford in 1895) became Instructor in Biology. Hoag (1868-1924) was author of, among other works, Health Studies: Applied Physiology and Hygiene, published by D.C. Heath and Company in 1909. The courses taught in this era were plant and animal physiology and classification, with a class in embryology and bacteriology started in 1895-96. Academic year 1897-88 marked the first (and for decades, the only) year a course in neurobiology, "Special Course in the Nervous System," was taught.
In 1898 the Instructor in Geology and Biology was Edward Waller Claypole. From the 1899-1900 school year to 1901-1902 he was Professor of Geology and Biology, the first biologist listed as professor. Claypole (1835-1901) was an Englishman educated at the University of London. He moved to the United States in 1872, teaching at a variety of colleges in a variety of subjects, from classics to biology to geology. He was, among other contributions to American science, a founder of the journal American Geologist, and was well-known for his studies of Devonian placoderms (armored fish). A detailed biography, both personal and scientific, can be found in American Geologist (1902, v 29).
Claypole had twin daughters, Agnes Mary Claypole (1870-1954) and Edith Jane Claypole (1870-1915). Both attended Buchtel College in Akron, Ohio, where Claypole was teaching at the time, and graduated in 1892. Both entered graduate school at Cornell. Edith earned an M.S. in 1893 with a thesis on the blood cells of amphibians; Agnes received an M.S. in 1894 with a thesis on the digestive tract of eels. Edith then went off to teach at Wellesley, while Agnes went to the University of Chicago, where she earned her Ph.D. in 1896. After two years at Wellesley, Edith entered the Cornell Medical School to earn a medical degree. Their father and stepmother moved to Pasadena in 1898, for reasons having to do with Mrs. Claypole's health, and both daughters joined them. When Edward Claypole died in August 1901 (his wife following only a few weeks later), the daughters were appointed to the Throop faculty in his place. In 1902-1903 the catalog lists Agnes Mary Claypole as Instructor in Zoology and Edith Jane Claypole as Instructor in Biology and Bacteriology. Edith left the faculty to complete her medical education at what is now UCLA (later dying, heroically, of a typhoid fever infection incurred while working toward a vaccine for European troops during WWI). Agnes remained at Throop and was promoted. In 1903-04 Agnes M. Claypole was Professor of Natural Science and Curator, the first female biology professor in the Institute. She left after holding the position for only one year, however – she married Robert O. Moody, a professor of anatomy at UC Berkeley in 1903, and afterward moved to northern California. She was listed with a star in the first seven editions of American Men of Science (the star meant she was considered one of the top 1,000 scientists in the United States in 1903), and was one of the founding members of the Pacific Division of the AAAS; she was a member of the sociology faculty of Mills College in Oakland from 1918 to 1923.
In the following school year at Throop, 1904-05, the sole biologist on the faculty list was Joseph Grinnell, Instructor in Natural Science and Curator. Grinnell was one of the great biologists of the 20th century. Born in 1877, he was a student in the college portion of Throop, receiving his A.B. in 1897. He was the third college graduate of Throop, the first graduating class having been two students in 1896, with Grinnell's class, the class of 1897, a class of one. In the 1897-8 school year Grinnell was Assistant Instructor at Throop, he then went to Stanford to receive his A.M. in 1901, and to remain a graduate student until 1903. On his return from Stanford to Pasadena, he was appointed (1904-5) Instructor in Natural Science and Curator at Throop. In 1906 and 1907 he was Professor of Biology and Curator. He began his professional work as an ornithologist and herpetologist, in 1898 publishing the noted Birds of the Pacific Slope of Los Angeles County, published by the Pasadena Academy of Sciences. In 1907 he was recruited to become the founding director of the Museum of Comparative Zoology at the University of California at Berkeley, and from 1908 to his death in 1939 was the director as well as a renowned professor at Berkeley. He is considered the first important west coast mammologist. Among his notable contributions is the concept of the ecological niche, which he first published in 1924, and the invention of the now-standard method of taking field notes on wildlife, in which journal entries, species accounts, and specimen catalogs are combined. He was an important advocate for the National Park system; his field studies at Yosemite from 1914 to 1924 (leading to the publication of Grinnell and Storer, Animal Life in the Yosemite, 1924) are still well known. Several of his books on California birds and mammals are still in print (available as reprints), nearly 100 years after they were written, and Grinnell Mountain in the San Bernardino range is named after him.
At Throop in 1906 and 1907 there was along with Grinnell another biologist, Lecturer Ernest Hoag (the same Hoag who had been Instructor from 1895-1898). In 1908-9 and 1909-1910 the Professor of Biology is listed as Carl Spencer Millikan, B.S. (from MIT in 1899). Millikan (born in 1876) had been a professor at Ripon College prior to coming to Throop, and worked on the development of the chick retina. Hoag remains listed as Lecturer through 1909-1910. Millikan is listed as professor for the last time in the Throop catalog of April, 1910; that year the only biology course offered was zoology. Despite a promise that Botany would be offered in alternate years, the Throop catalogs from 1910-11 through 1915 list the biology courses as "not offered." Biology courses are not listed thereafter, until the present division was started in 1928. Nonetheless there was at least one lecture in biology after the 1910 cessation of the teaching of the subject, the March, 1911 lecture entitled "A Zoölogical Trip through Africa" by Mr. Theodore Roosevelt.
1920s and 1930s: Foundation of the Current Division
"At the California Institute of Technology, Morgan wanted to give a concrete form to his philosophy of science in general and of biology in particular. He wanted to emphasize the new direction in which he thought biological research ought to move. It was his aim to bring together the best possible people representing the most modern lines of biological research…and allow them virtually unrestricted possibilities to interact." Allen, G.E. (1978) Thomas Hunt Morgan, the Man and His Science, Princeton, 447 pp., p. 334.
The Caltech Division of Biology, the predecessor of the present Division of Biology and Biological Engineering, was founded in 1928. Thomas Hunt Morgan, the world's pre-eminent geneticist, Professor of Biology at Columbia University, and President of the National Academy of Sciences, was hired by the Institute to start a Division of Biology. The first wing of the Kerckhoff building was constructed (1928) and Morgan set out to recruit a young but distinguished faculty in the five areas of genetics and evolution, experimental embryology, biophysics, physiology, and biochemistry. He intended later to add faculty in experimental psychology – what we call today integrative neurobiology.
Morgan recruited first in genetics, and the starting faculty in 1928 was primarily from his own laboratory and from that of R.A. Emerson at Cornell, the leading plant geneticist of the time. By the 1929-1930 school year the faculty consisted of full professors T.H. Morgan; Alfred H. Sturtevant, a Drosophila geneticist from Morgan's lab who discovered genetic mapping; and Karl J. Belar. Belar was a distinguished young cytogeneticist who had been hired away from the Kaiser Wilhelm Institute for Biology in Berlin-Dahlem. He died in an automobile accident during a trip to the desert in 1931. Ernest G. Anderson, a maize geneticist from the Emerson lab and a former postdoc in Morgan's lab, was associate professor. The assistant professors were Theodosius Dobzhansky (evolutionary genetics, a recent postdoc of Morgan's from Russia), Sterling Emerson (plant genetics, the son of R.A. Emerson), Henry Borsook, a biochemist from Toronto, and Herman E. Dolk, a plant physiologist from the Netherlands. Dolk, like Belar, died in an automobile accident, in 1932. Calvin Bridges, with Sturtevant the core of Morgan's Drosophila group, came as a Research Fellow of the Carnegie Institution, a position he held at Caltech until his death in 1938; and Albert Tyler, an embryologist, graduate student of Morgan's and Caltech's first Ph.D. in biology (1929), was appointed instructor, eventually becoming a professor (in 1938), and remaining at Caltech for the rest of his life (to 1968).
Not everyone who was asked to join the new division accepted. Among those considered but not landed were Curt Stern (a geneticist from the Kaiser Wilhelm Institute, who later visited Caltech in 1932, while in flight from Germany), Leonor Michaelis (of the Michaelis-Menten equation) and Selig Hecht, the Columbia biophysicist – each to become important scientists in their respective fields, but not at Caltech.
Some of the events of the earliest days of the new division, housed in what is now the west wing of the Kerckhoff building, were recounted in the December 1952 edition of the division's newsletter "Bio-peeps." Among them were the first class held in Kerckhoff (fall of 1928, the only course then offered was Bi 1), the first research fellows (Theodosius Dobzhansky and Yoshitaka Imai, later a distinguished plant geneticist at Tokyo University), the first graduate students (Albert Tyler, Russell L. Biddle, Carl C. Lindegren, Norwood K. Schaeffer and William A. Hetherton; Tyler was the first to graduate), the first child born to a faculty member (Harriet Sturtevant), the first female research assistant on the Caltech campus (Elizabeth L.D. Griffiths, working with Sterling Emerson), and the first female postdoc (Eileen Erlanson, who gave the first seminar in the new division).
Others hired as professors, mostly assistant professors, in the first half of the 1930s included Robert Emerson (biophysics, 1930), Hugh Huffman (biochemistry, 1931), Frits Went (plant physiology, 1932, to replace the late Dolk), George MacGinitie (a marine biologist hired in 1932 to direct the new Kerckhoff Marine Lab at Corona del Mar, and one of only two faculty members in divisional history not to have a doctoral degree—the other was Dobzhansky), and Cornelis Wiersma in physiology (1934). Later in the decade the professorial faculty was augmented by Arie Haagen-Smit (a bio-organic chemist, 1937), James F. Bonner, a plant physiologist and Caltech Ph.D. in biology (Ph.D. 1934, hired in 1938), Anthonie van Harreveld (neurophysiology, 1939) and Johannes van Overbeek (plant physiology, 1939). Kenneth Thimann, the plant physiologist, was an instructor beginning in 1930.
During the division's first decade a number of postdoctoral fellows, graduate students and visiting professors were also at work. Many of these rose later to great prominence in biology. Among the postdoctoral fellows were George Beadle, later to chair the division; Max Delbrück, later to return as Professor of Biology; Barbara McClintock; Charles Burnham; and Georgii Karpechenko (a student of Vavilov's who was to return to the Soviet Union and to die, like Vavilov, in one of Stalin's prison camps as a victim of Lysenkoism). Other international postdoctoral visitors were Cyril Darlington, Curt Stern, Boris Ephrussi, and D.G. Catcheside. Among the graduate students were Chia-Chen Tan (Jia-Zhen Tan, former vice president of Fudan University, where he started China's first department of genetics—he is considered "the founder of Chinese genetics"); Norman Horowitz, later to join the faculty of the division, and to serve as chair in the 1970s; David M. Bonner (after whom Bonner Hall at U.C. San Diego is named, and brother of James); Edward Novitski; and Edward B. Lewis – who later joined the divisional faculty, and remained a member of it until his death in 2004. Novitski wrote a memoir on aspects of the division, including his life as a graduate student in the late 1930s, called Sturtevant and Dobzhansky: Two Scientists at Odds With a Student's Recollections.
A program in undergraduate education was also established when the division was founded. The original courses taught in the 1929-30 school year were Bi 1, 2, 3, 4, and 5bc, introductory courses in biology including physiology, botany and histological technique, and a series of advanced courses such as Bi 100, Genetics, taught by Sturtevant, Anderson, Dobzhansky and Emerson, and Bi 110, Biochemistry, taught by Borsook. Bi 110 is still the biochemistry course number, more than 85 years later. After 20 years (through 1950) the total number of men who received the B.S. degree in biology was around 80 (and they were all men, as Caltech did not accept women as undergraduates until 1970). Thus the program was small by current standards. Around 80% of the graduates went on for doctoral degrees, half to earn Ph.D. degrees and half to work toward the M.D. degree.
One highlight of the 1930s was the award of the Nobel Prize for Medicine or Physiology to T.H. Morgan, in 1933 – the first of the 6 Nobel Prize winners to serve as members of the Division of Biology faculty. Another was the completion in 1938 of the second wing of the William G. Kerckhoff Laboratories – the eastern part of the present building.
"Now, his kindness, his undoubtedly sensitive nature, he hid under a protective cover of eccentricity and what I can describe no better than to say impishness. He was an imp. He liked to shock people, liked to say something unexpected, to behave in a somewhat unorthodox manner. He was dressing himself so poorly that there was at least one occasion when the laboratory janitor was taken to be Professor Morgan, and Professor Morgan was taken to be the janitor. In summer time in California, very frequently he was walking with his trousers kept in place not by a belt or by suspenders, but by a string. A piece of string." Theodosius Dobzhansky, Reminiscences, Columbia University Oral History Collection, 1962.
1940s: Changing Times
Among the major influences in the 1940s were World War II, Morgan's retirement, and the return of Beadle. The division faculty and divisional programs were well in place, and in full-scale operation by the 1940-1941 school year. The United States entered World War II in December 1941, bringing normal operations to a halt. The Division of Biology and its staff served in many ways during the war. The plant physiologists, including Bonner, spent the war looking for new plants that could be sources of rubber, to replace the Hevea brasiliensis plantations of Malaysia, which had been seized by the Japanese. Others joined the armed forces, Ed Lewis, for example, completing his Ph.D. in biology under Sturtevant, then earning an M.S. in meteorology before shipping out to the Pacific as an army meteorologist.
During the war years, no new members were added to the professorial faculty. Van Overbeek left (eventually becoming chair of the Biology Department at Texas A&M University), Huffman left (to establish a thermodynamics research laboratory of the Bureau of Mines in Bartlesville, Oklahoma), and Morgan retired (in 1942). The division was without a chairman from 1942 until 1946, with Sturtevant serving in lieu of chair as Chairman of the Biology Council, a committee of full professors (Borsook, Haagen-Smit, Sturtevant and Went). In 1946 George Beadle left his faculty position at Stanford to return to Caltech as the new division chair. He immediately began hiring new faculty, starting with Max Delbrück, who had spent the years between his departure from Caltech as a research fellow and return to Caltech as a Professor on the faculty at Vanderbilt. Shortly afterward Norman Horowitz (former graduate student of Tyler at Caltech, then postdoc with Beadle at Stanford) and Ray Owen (from the University of Wisconsin) were hired as Associate Professors, and Ed Lewis as Instructor (1947) – he was soon Assistant (1948) and Associate (1949) Professor.
By the 1949-1950 school year Herschel Mitchell (from the Tatum lab at Stanford, then a Senior Research Fellow with Beadle at Caltech from 1946) joined as Associate Professor. Among the postdoctoral fellows and visiting researchers were Renato Dulbecco, Arthur Galston, Sam Wildman, Barry Commoner, Robert Bandurski, Seymour Benzer, William Drell, Melvin Green, Urs Leupold, S.E. Luria, Clement Markert, Roger Stanier, and others, who would become renowned biologists in the 1950s and 1960s.
The year 1949 also marked the opening of two new buildings for the Division of Biology – the Biology Annex (the facility now under the Alles Patio) and the Earhart Plant Research Laboratory, torn down in the 1970s to make way for the Beckman Behavioral Biology building.
"The Division of Biology of the Institute has now been in operation twenty-one years. Its steady growth and development during this period is a tribute to the imagination and foresight of those who planned and established it. Its first Chairman, the late Professor Thomas Hunt Morgan, built its foundations well and guided its growth wisely for thirteen years…During the year the work of the Division was carried on by eleven full professors, eight research associates, seven associate professors, eight senior fellows in research, twenty-two research fellows, thirty-two graduate students and seventy-six research technical and laboratory assistants…" G.W. Beadle, Caltech Biology Annual Report, 1949.
1950s: Fungi, Flies and Phage
"Following World War II many outstanding advances in biology have been made in laboratories all over the world. To mention only a few of these: Bacteriology has been revolutionized by a group of young investigators using the methods of genetics, cytology, and biochemistry. Our knowledge of viruses has been greatly increased, particularly from a biological point of view. Through a growing interest by physicists and physical chemists in biological problems, biophysics has grown rapidly. It has made extensive use of the electron microscope, the preparative and analytical centrifuges, the Tiselius electrophoresis apparatus and other techniques completely unknown to the biology of a few years ago." Caltech Biology Annual Report, 1950.
In terms of faculty appointments, the 1950s were quantitatively static. In the course of the decade all of the associate professors hired in the late 1940s became full professors, and four new faculty members joined the division, all at the level of associate or full professors (Arthur Galston, 1951, Renato Dulbecco, 1952, Roger Sperry, 1954 and Robert Sinsheimer, 1957). By the 1957-58 school year, and until the end of the decade, there were no associate or assistant professors. Four faculty members departed in the '50s – MacGinitie retired in 1957, E.G. Anderson in 1959, Went departed (1959) to become head of the Missouri Botanical Garden, and Arthur Galston left to join the faculty at Yale in 1955.
The three new senior appointees of the 1950s who were still at Caltech by the end of the decade were Renato Dulbecco, Roger Sperry, and Robert Sinsheimer. Two were to win Nobel Prizes, Dulbecco for work in animal viruses (by the time of his award he had left to join the new Salk Institute), and Sperry for discovering that different brain hemispheres serve different functions in humans. Sinsheimer was later Chancellor of the University of California at Santa Cruz, which named its Robert L. Sinsheimer Laboratories after him.
Remarkable scientific progress was made in the laboratories of those appointed in the '40s, three of whom (Beadle, Delbrück, and Lewis) were also to win Nobel Prizes. A roster of some of the students, postdocs and long-term visitors to the Delbrück lab in 1951-60 gives an idea of the nature and significance of the work there - Jean Weigle, Renato Dulbecco, Marguerite Vogt, Seymour Benzer, Giuseppe Bertani, Margaret Lieb, Gunther Stent, Elie Wollman, Dale Kaiser, Gordon Sato, Ole Maaloe, Niccolo Visconti, Robert Sinsheimer, James Watson, Harry Rubin, George Streisinger, Naomi Franklin, Andre Lwoff, Charles Steinberg, Frank Stahl, Howard Temin, Matthew Meselson, Harriet Ephrussi-Taylor, Francois Jacob, Sydney Brenner and Millard Sussman. Similar lists could be drawn for other divisional laboratories.
One other notable piece of progress was the admission of women as graduate students in what had before been an all-male student body. Dr. L. Elizabeth Bertani was the first of many women to be granted a Ph.D. in biology at the Institute, for her 1957 thesis "Studies on the Establishment of Lysogeny by Bacteriophage P2." In fact, Division of Biology faculty had decided to admit women a full ten years earlier, having recommended to the Institute the admission of women to graduate standing after a unanimous faculty vote in 1947. It took some time for the Institute to catch up.
"The next question considered was "Does the Division want to recommend that women be admitted to graduate study in Biology?" After much discussion and a statement from each member present, Dr. Sturtevant moved that the Division go on record as favoring the admission of women graduate students at the Institute. Dr. Bonner seconded the motion and it was unanimously carried. Dr. Sturtevant recommended that Dr. Beadle inform the proper authorities of the Division's recommendation." Minutes of Staff Meeting May 20, 1947, p.2.
Beadle's Nobel Prize in Medicine or Physiology was awarded in 1958.
In the '50s the division was active in building – the Norman Church Laboratories of Chemical Biology opened in 1955, largely a chemistry building, but with a small western wing for Division of Biology labs. Construction was started for the Campbell Plant research Laboratory, a greenhouse completed in 1960 (and demolished in the 1980s); and the Gordon A. Alles Laboratory for Molecular Biology – also completed in 1960. E. G. Anderson's retirement in 1959 marked the end of research at the division's 10-acre farm in Arcadia, which had been used for maize studies since the founding of the division; the "Arcadia Farm" is now part of the campus of Temple City High School.
"I discovered a little book called Viruses, which was from a symposium held at Cal Tech in 1950. It's a remarkable book. I read avidly about phages in it and got very excited. This was the beginning of the phage ideas – stuff that was going to become clear in the next twenty years! So I started to work on bacteriophages..." Sydney Brenner, My Life in Science, BioMed Central, 2001, p.21.
"When the Pasadena meeting on protein structure finished at the end of September, the full horror of being in Pasadena hit me. Not knowing how to drive a car, much less owning one, I was effectively confined to the girlless Caltech campus and had to continue living at the faculty club... I became part of Max Delbrück's ground-floor phage group in the 1930s-style Kerckhoff Biology Building." James D. Watson, Genes, Girls and Gamow, Alfred A. Knopf, New York, 2002, p. 45.
The 1960s opened with a burst of new faculty, all appointed for the 1960-61 school year. At the level of full professor Alan Hodge, an electron microscopist, and Anton Lang, the renowned plant physiologist, were appointed, Lang as a replacement for Frits Went. Robert Edgar and Charles Brokaw (who received his B.S. in biology from Caltech in 1955) were appointed assistant professors. In the following years of the decade 10 additional new faculty appointments were made: Derek Fender as associate professor and Giuseppe Attardi as assistant professor for the 1962-3 academic year; William Dreyer as full professor for the 1963-4 year; Felix Strumwasser as associate and William Wood as assistant professor for 1964-5; Jerome Vinograd as full professor for 1965-6; Seymour Benzer as full professor for 1967-8; Daniel McMahon as assistant professor in 1968-9; and James Olds as full professor and James Strauss as assistant professor in 1969-70. The 14 new faculty hired in the '60s were only partly balanced by retirements and departures – Sturtevant and Borsook retired, Beadle, Dulbecco and Lang departed, and Tyler died. The division thus experienced substantial net growth.
Beadle's departure deserves separate mention. After 14 years as division chair he was invited to become the President of the University of Chicago, and departed Caltech in January of 1961. He was replaced as division chair by Ray Owen, among whose contributions to immunology had been the discovery of immune tolerance, leading to our present abilities in organ transplantation. Owen at first accepted appointment only as acting chair until a new outside chairman could be found, but became division chair for the 1962-3 academic year. In April, 1968 he was succeeded by another internal candidate, Robert Sinsheimer.
There were many research highlights in the 1960s, including the Bonner lab's explorations of chromatin, Sinsheimer's work on the single stranded DNA virus ϕX174, the beginning of the classic work of Edgar and Wood on the genetic control of phage assembly, Olds's explorations of the results of direct brain stimulation, Sperry's discovery of hemispheric specialization in human brains, Lewis's initial studies of the genes of the Bithorax complex, and the beginning of Benzer's work on the neurobiology and genetics of behavior in fruit flies.
The 1960s were also the decade when the biology undergraduate program began to grow. From the beginning of the division until 1962 the typical biology graduating class was 3 or 4 students, rarely more. In 1962 11 graduated, with classes of 8, 12, 10, 4, 11, 8 and 10 following through 1969. There were a total of 83 B.S. degrees in biology given in the decade, with fewer than 120 total degrees given from the 1930s to 1960. Biology was beginning to become popular with Caltech undergraduates. A few of the students who earned a B.S. in biology in the '60s are Thomas Jovin, Leroy Hood, Leland Hartwell, Ira Herskowitz and Douglas Brutlag – familiar names now, 30 years later, and another set of examples of the saying that Caltech is a good place to go to, and a good one to come from.
The decade ended with the award of the 1969 Nobel Prize to Max Delbrück for his leadership in establishing phage genetics and the field of molecular biology.
"'We planned the experiment that day,' Jacob said. 'That's when we decided, Sydney and myself, to go to Cal Tech. I had been invited by Delbrück to come and spend a month there, and Sydney had been invited by Meselson'…they put three of the six [tubes] into Meselson's usual centrifuge, the other three into a second machine downstairs in Dulbecco's lab. They started them up. Then they found that Weigle's water bath was contaminated by the spilled P32, so they rinsed it out and hid it behind the Coca-Cola machine in the basement to cool off. In Brenner's remembrance, the next day was the day they went to the beach…" Description of the experiments in which the existence of messenger RNA was proven, in June, 1960; The Eighth Day of Creation, H. F. Judson, Simon and Schuster, 1979, pp. 433-439.
1970s: Brain Research
The 1970s, similarly to the 1960s, began with a series of new appointments. The first was Leroy Hood, a former Caltech undergraduate and graduate student (B.S. 1960, Ph.D. 1968), as assistant professor in 1970, followed by appointment of the electron microscopist and cell biologist Jean-Paul Revel as full professor. In 1971 Eric Davidson was appointed as associate professor and Richard Russell as assistant professor. In preparation for and following the opening of the newly-built Beckman Laboratory for Behavioral Biology (January, 1974) several young neuroscientists and biophysicists were appointed: Henry Lester and Jack Pettigrew (1973), John Allman and Ronald Konopka (Caltech Ph.D. 1972), for the '74-5 academic year, Mark Konishi and James Hudspeth ('75-6), David van Essen (Caltech B.S. 1967, appointed for '76-77), and Jeremy Brockes (1978). Howard Berg (Caltech B.S. 1956) was appointed as Professor of Biology in 1979. In addition a new start in cell and molecular biology was begun by the hiring of Elias Lazarides as assistant professor in 1976, and Tom Maniatis as associate professor in 1977. The 15 new appointments in the decade were nearly balanced, however, by the retirements of Emerson, Haagen-Smit, van Harreveld, Wiersma, and Delbrück; by the deaths of Vinograd and Olds; and by the departures of Hodge, McMahon, Russell, Sinsheimer and Wood (to chair the Molecular, Cellular and Developmental Biology Department at the University of Colorado).
Sinsheimer's departure necessitated the appointment of a new division chair, and this position was filled by Norman Horowitz, Caltech Ph.D. 1939 (with Tyler), collaborator of Morgan's while a student, postdoctoral fellow of Beadle's at Stanford in the 1940s, and Caltech faculty member from 1946. Horowitz assumed the chairmanship September 1, 1977.
Among the research topics gaining new prominence in the '70s were, in molecular biology, electron microscopy of DNA, led by Professor of Chemistry (and later Professor Emeritus of Biology) Norman Davidson; development of automated protein sequencers by Dreyer and Hood; and the introduction of recombinant DNA technology by Maniatis. The numerous appointments of neurobiologists opened a number of new fields for the division, from Hudspeth's work on hair cells, to Konopka's on circadian rhythms, van Essen's on the visual cortex, and Konishi's on sensory integration in owl behavior.
It was also in this decade that former faculty member Renato Dulbecco, who had been at Caltech from 1949 to 1962, was awarded the 1975 Nobel Prize, along with his former Caltech graduate student Howard Temin, and a young MIT professor (and former Dulbecco postdoctoral fellow), David Baltimore, who would later come to Caltech as its President and Professor of Biology in 1997.
Undergraduate education gained more prominence in the division, as the number of biology B.S. graduates took another sharp upturn. There were 210 graduates from 1970 through 1979, with one class, that of 1974, having 37 graduating biologists – still a record for the division. The first year women were admitted as undergraduates was 1970. This action was a result of the recommendation of an Institute committee chaired by biologist Ray Owen. The first wave of female graduates was in 1974, as the women admitted as freshman in 1970 completed their work – and 10 of the 37 biology graduates that year were women. At least one female biologist graduated even earlier, though not as a biology major – Sharon Long entered as a sophomore transfer student in 1970, graduating in 1973 as an Independent Studies major. She became (and remains) a noted biologist, professor and dean at Stanford. A random selection of other undergraduate biologists of the '70s who are now well-known are James Gould, William Chia, Brian Seed, James Posakony, Thomas St. John, Ed Hedgecock, and Joe Kirschvink – Kirschvink is now a Professor in Caltech's Division of Geological and Planetary Sciences.
1980s: Gene Cloning
The first new faculty members to start in the 1980s were John Hopfield, appointed as full professor of Biology and Chemistry, and Elliot Meyerowitz, appointed Assistant Professor of Biology. In 1981 Mary Kennedy and Barbara Wold became assistant professors, followed by Ellen Rothenberg in 1982, and in the same year John Abelson and Mel Simon came as full professors. In 1983 the assistant professors' ranks were incremented by Mark Tanouye and Scott Emr, and Paul Patterson joined as a full professor; in 1984 James Bower and in 1985 David Anderson became assistant professors. Judith Campbell also joined the Division of Biology in 1985 as Associate Professor of Chemistry and Biology; prior to this (from 1977) she had been a faculty member solely in the Division of Chemistry. In 1986 assistant professors Howard Lipshitz and Christof Koch joined, Koch as Assistant Professor of Computation and Neural Systems, a new interdisciplinary area started by, among others, Hopfield. Paul Sternberg arrived in 1987 as Assistant Professor of Biology, as did Pamela Bjorkman, William Dunphy and Kai Zinn in 1989. All told 19 new appointments were made to the biology professorial faculty in the decade.
Of course the new additions were partly balanced by departures – there were the retirements of Bonner (1981), Horowitz (1982), Owen (1983), Mitchell and Sperry (1984), Fender (1986) and Lewis (1988); and in addition Maniatis, Strumwasser, Berg, Hudspeth, Pettigrew, Brockes and Konopka departed. This still left room for an increase in the faculty by 5, to a total of 33.
The chairmanship changed hands twice in the decade; in 1980, with the appointment of Lee Hood as chair, and again in 1989, when Hood stepped down and John Abelson became division chair. Among the highlights of the decade was the award of the Nobel Prize to Roger Sperry in 1981. The building program also continued in the '80s, with the construction of the Braun Laboratories (1982) and of the Beckman Institute (1989), each of which contain a mix of labs from different divisions – Braun being shared between the Divisions of Chemistry and Biology, and the Beckman Institute housing not only the semiautonomous Institute but also laboratories from Biology, Chemistry, and elsewhere.
One of the research themes of the 1980s was the advent and wide utilization of gene cloning, and gene sequencing. Maniatis came as one of the pioneers of these new technologies, as did Simon and Abelson, and Meyerowitz, Wold, Lipshitz and Zinn as new faculty members trained in them. Hood in particular among the faculty switched his laboratory to exploit the new possibilities of genomics, and one result of this was the invention in the Hood lab of the automated DNA sequencer, which played a central role in the genome projects of the 1990s and beyond.
1990s: Genomics, Proteomics, and Macromolecular Machines
The 1990s continued the hiring of new faculty that characterized the previous decade. In 1990 Gilles Laurent was hired as Assistant Professor of Biology and of Computation and Neural Systems, followed by appointment of Scott Fraser and Alexander Varshavsky as full professors (1991 and 1992, respectively), Stephen Mayo as assistant professor in 1992, Raymond Deshaies and Erin Schuman as assistant professors and Richard Andersen as full professor in 1993, Marianne Bronner as full professor and Bruce Hay as assistant professor in 1996, and Shin Shimojo as associate professor in 1997. The year 1997 also saw the arrival of David Baltimore as President and Professor of Biology, the first time a biologist had been president of the Institute. Assistant Professor Jose Alberola-Ila joined in 1998.
The decade also saw the departures of Mark Tanouye, Elias Lazarides, Lee Hood, Scott Emr, David Van Essen, and Howard Lipshitz, the retirement and departure of John Hopfield, and the formal (but not actual!) retirement of Seymour Benzer. John Abelson completed his term as chair in 1995, to be replaced by Mel Simon.
The long tradition of using Drosophila genetics to explore fundamental biological questions was recognized by two prestigious prizes awarded that decade. Seymour Benzer received the Crafoord Prize in 1993 for his work examining the genetics of behavior. And Ed Lewis won the 1995 Nobel Prize for Physiology or Medicine for his work in developmental genetics – the sixth Nobel Prize awarded to a member of the divisional faculty.
One of the new research directions that became evident in the '90s was structural biology, with Pamela Bjorkman of the Division of Biology and Douglas Rees of the Division of Chemistry solving numerous important protein structures by modern methods of x-ray crystallography, and Stephen Mayo producing new theoretical methods for understanding protein folding. Genomics also became a core part of the division, with Caltech contributions to the model organism genome projects such as Paul Sternberg's work with Caenorhabditis elegans and Elliot Meyerowitz's work with Arabidopsis thaliana. Mel Simon played a key role in the human genome project – it was his laboratory that invented the BAC clones that made it possible, and that produced the libraries that were sequenced.
The undergraduate program remained highly vigorous in the 1990s. While the large classes of the 1970s were not seen again in the 1980s (the largest class of B.S. graduates in that decade was the class of 1989, with 18), the 1990s surpassed all earlier decades, with total of 223 undergraduate degrees from 1990 through 1999.
2000 and Beyond: The Past is Prologue
The division achieved its 75th anniversary in 2003 with approximately 700 faculty, students, postdoctoral fellows, and staff.
The new decade started with a change in leadership; Elliot Meyerowitz was appointed the chair of the division in 2000, after Mel Simon completed his term. In that decade there were 12 new appointments to the faculty, David Chan as assistant professor in 2000, Thanos Siapas and Grant Jensen as assistant professors and Michael Dickinson as full professor in 2002, Michael Elowitz as assistant professor in 2003, Ralph Adolphs as professor and Angelike Stathopoulos as assistant professor in 2005, Dianne Newman as professor and Sarkis Mazmanian as assistant professor in 2006, and from 2008 to 2009 David Prober, Doris Tsao, and Alexei Aravin were appointed as assistant professors (Aravin arrived in 2010). Siapas and Dickinson were joint appointments (Biology and Engineering and Applied Science) and represented the growing influence of a new bioengineering option. There were departures, as well; Charles Brokaw retired in 2000, James Bower left in 2001, John Abelson retired in 2002, Mel Simon retired in 2005, Jean-Paul Revel in 2006, James Strauss in 2007, and Erin Schuman and Gilles Laurent left to start a new Max Planck Institute for Brain Research in Frankfurt in 2008. The decade also saw the deaths of Bill Dreyer and Ed Lewis in 2004, Norm Horowitz in 2005, Seymour Benzer in 2007, and Giuseppe Attardi in 2008. All but Horowitz and Dreyer were active in research at the time of their death.
A new building, the Broad Center for the Biological Sciences, was completed and occupied in September 2002. Seven laboratories from the division (including Chan and Jensen, who were hired to help occupy the new building, and Michael Elowitz, a computational and synthetic biologist who arrived in mid-2003) moved there, as well as a lab from the Division of Chemistry and one from Engineering. Since the opening of this modern building designed for interaction between groups and research areas, the basement (initially shell space) has been fully developed, bringing a total number of 13 biology laboratories to the space.
The new decade of the teens started as the new century had, with a change in leadership; Stephen Mayo replaced Elliot Meyerowitz as chair after Meyerowitz completed his second 5-year term in mid-2010—the first Chair since George W. Beadle (who was chair for 14 years) to serve the maximum of 10 years. New arrivals so far in this decade have included Lea Goentoro as assistant professor and Rob Phillips as professor in 2011, Viviana Gradinaru as assistant professor and Markus Meister as professor in 2012, Mitch Guttman as assistant professor in 2013, and Yuki Oka and Elizabeth Hong as assistant professors in 2015. Ray Owen died in 2014 at the age of 98, severing one of the last direct links to the Beadle years, and Paul Patterson died at the untimely age of 70 while still the leader of an active research laboratory.
A reorganization of the division also took place, which resulted in a considerable increase in faculty size: in 2013 the Division of Biology voted to incorporate bioengineering (previously organized under the Division of Engineering and Applied Science), and to change its name to the Division of Biology and Biological Engineering—the first change of a division name at Caltech since 1970. Along with this change came an enthusiastic vote to provide joint appointments to members of the bioengineering group who were faculty in other divisions, leading to the addition of 10 new professorial faculty members. The new additions were Frances Arnold and Rustem Ismagilov from Chemistry and Chemical Engineering, and John Dabiri, Morteza Gharib, Richard Murray, Niles Pierce, Lulu Qian, Michael Roukes, Erik Winfree, and Changhuei Yang from Engineering and Applied Science. The size of the new BBE division, therefore, will be 50 assistant and full professors—an all-time high and a precursor of exciting times to come. As the BBE division enters its 87th year, the traditional areas of intellectual concentration—genetics, biochemistry, developmental biology, immunobiology, microbiology, molecular biology and neurobiology—are joined with the new areas of genomics, synthetic biology, and systems biology. But as in the beginning, the BBE division of 2015 is dedicated to educating students and postdocs, and to solving four of the great intellectual and practical problems of humankind: the origin and evolution of life, the mechanisms by which a cell functions, the development of organisms from fertilized eggs, and the mechanisms by which the brain works.
George W. Beadle Professor of Biology