A Short, Informal History of the Biological Sciences at Yale University

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A Short, informal History of the Biological Sciences at Yale university

Peter B. Moore

Department of Chemistry, Yale University, New Haven, Connecticut in May 2012, Yale University’s Graduate school of Arts and sciences held a reunion for all who have received doctorates from Yale in the biological sciences. The proceedings began with two presentations on the history of biological research at Yale: one focused on the Med - ical school, and the other centered on the rest of the University. This essay is a lightly ed - ited version of my account of the history of the biological sciences outside the Medical school.

introduction corporated into the academic structure of As everyone knows, biology, broadly Yale University. Indeed, the history of bi - construed, is today a vast, lavishly sup - ology at Yale has been full of twists and ported, scientific enterprise that attracts the turns that still affect the way biology is interest of a larger and more heterogeneous done in New Haven. group of scientists than any other branch of In the first place, unlike the other science. Given its prominence in today’s major branches of science (e.g., physics, scientific universe, it may come as a sur - chemistry, astronomy, geology, etc.), biol - prise that biology was the last of the major ogy has never had a single departmental scientific disciplines to become fully in - home at Yale, nor has there ever been a de -

To whom all correspondence should be addressed: Peter B. Moore, Department of Chem - istry, Yale University, PO Box 208107, New Haven, CT 06520-8107; Tele: 203-432-3995; Fax: 203-432-5781; Email: [email protected].

†Abbreviations: DPA, Department of Philosophy and the Arts; EEB, Ecology and Evolu - tionary Biology; MB&B, Molecular Biophysics and Biochemistry; MCDB, Molecular, Cellu - lar and Developmental Biology. 551 552 Moore: The biological sciences at Yale

partment focused on biology that could tHe 19tH centurY: tHe SHeffield Scientific ScHool claim to be more than first among equals. Today, there are a large number of depart - It is impossible to understand what hap - ments outside of the Medical School in pened to biology at Yale outside the Medical which biology is done. Those include the School in the century that followed Silli - obvious suspects ― Molecular, Cellular and man’s appointment if you do not know Developmental Biology (MCDB †), Ecology something about the history of the division and Evolutionary Biology (EEB), and Mo - of the University that began in 1846 as the lecular Biophysics and Biochemistry Yale School of Applied Chemistry. For the (MB&B) ― but they also include Chem - record, a year later, Yale established a sub - istry, Geology, and Biomedical Engineering. division called the Department of Philoso - In order to understand how biology came to phy and the Arts (DPA), which was charged be scattered all over the University, you need with supervising post-bachelor’s degree ed - to go back to the beginning. ucation in all fields other than Law, Theol - ogy, and Medicine. Today we refer to it as the Graduate School of Arts and Sciences. BeginningS (Similar ventures were launched at other in - In its first century, the 18th century, stitutions of higher education at about the Yale College offered its students a modest same time.) level of instruction in the natural sciences The first two faculty appointments in and mathematics, but systematic instruction Applied Chemistry were John Pitkin Norton, in the sciences did not begin until 1804, Professor of Agricultural Chemistry, and when Benjamin Silliman took up his duties Benjamin Silliman, Jr., Professor of Practi - on the faculty of Yale College. His title was cal Chemistry. Their appointments had some “Professor Chemistry and Natural History,” important fine print attached. The Corpora - the latter being a catchall phrase applicable tion stipulated that neither man was to in - to all branches of science including geology, struct students enrolled in Yale College, nor which interested Silliman a lot, and what - was either to be supported by Yale College ever aspects of biology Silliman found funds. It may seem odd to the reader that the amusing. Corporation would have been willing to es - In 1810, only 6 years after Silliman’s ap - tablish two new positions, but unwilling to pointment, the Yale Corporation took the first support them. However, this paradox has a of the many steps it has taken since that en - simple explanation: The Corporation had no sured that the biological sciences would money. Nevertheless, the Corporation’s de - never have a single home at Yale. It agreed to mand that Applied Chemistry be self-fi - the establishment of the “Medical Institute of nancing led ultimately to its downfall, as we Yale College,” which was to be a joint ven - shall see. ture of Yale College and the Connecticut Since the education of students who al - Medical Society. It is now called the Yale ready had college degrees was a major com - Medical School. In its original manifestation, ponent of its mission, the School of Applied the Medical Institute was no more than a Chemistry was initially part of DPA. In trade school, but by the second half of the 1852, Yale established a School of Engi - 20th century, there was more biological re - neering, which was also incorporated into search being done at the Medical School than DPA, and two years later, Applied Chem - in the entire rest of the University. istry merged with the School of Engineering Since its founding, the Medical School to create the Yale Scientific School. For has been viewed by Yale as the proper home many years, DPA was little more than a for most, if not all, human biology, and little wrapper organization for the Scientific more will be said about it here because the School, which was the branch of the Uni - focus of this essay is the history of the bio - versity that educated the overwhelming ma - logical sciences outside the Medical School. jority of the students on whom DPA Moore: The biological sciences at Yale 553

bestowed advanced degrees. This should not necticut.) There was a stark contrast between come as a surprise; the Ph.D. degree was de - the forward-looking curriculum of the vised by German universities in the mid- Sheffield School and that of the 4-year bach - 19th century for the training of professional elor’s program offered at Yale College, scientists. which still emphasized Greek, Latin, and Norton, who was Yale’s first bio - Philosophy, just as it had a century before. chemist, died only a few years after the Sci - Why should any of this ancient history entific School was founded, and he was be of concern to recent recipients of ad - replaced by John Addison Porter, a recent vanced degrees in the biological sciences Yale College graduate who had been teach - from Yale? The reason is that from 1860 to ing at Brown. In 1856, Porter single-hand - about 1920, the Sheffield School was the edly solved the daunting financial problems home of most of the biologists at Yale out - that the Scientific School confronted in its side the Medical School. Some of them were early years by marrying Josephine Sheffield. pretty good. For example, around the turn of She was the daughter of Joseph Sheffield, a the last century, Lafayette Mendel collabo - railroad tycoon who lived in New Haven, rated with Thomas Osborne at the Con - and through his son-in-law, Joseph became necticut Agricultural Experiment Station on sufficiently interested in the Yale Scientific a series of nutritional experiments that, School to give it several gifts so large that among other things, demonstrated that ly - in 1861, in gratitude, the Corporation re - sine and tryptophan are essential nutrients named it the Sheffield Scientific School. for mammals. Their work also contributed Thereafter, the Scientific School was a mightily to the discovery of the first vitamin: financially independent subdivision of Yale vitamin A. There were other respectable bi - that ran its own degree programs in science ologists on the Sheffield faculty, e.g., Chit - and engineering, and at Joseph Sheffield’s tenden in Physiological Chemistry, Beecher request, it was provided with its own Board in Paleontology, Brewer in Botany, and Sin - of Trustees. It appears that the Corporation nott in Plant Genetics. (Sinnott’s textbook acquiesced to this peculiar arrangement in on genetics was still in use in the late 1950s, anticipation of future gifts from Sheffield, as the author can attest from personal expe - which, gratifyingly, did ultimately material - rience.) ize. In addition to the income the School re - For many years, the President of Yale ceived from tuition and from Sheffield’s attended Sheffield commencement cere - benefactions, it received government funds monies, which in those days included a re - due to the fact that from 1863 to 1893, it was search talk given by one of its advanced Connecticut’s land grant institution. degree recipients. Thus, Noah Porter was on The Sheffield School offered its stu - hand in 1879 to hear a talk entitled “The dents several bachelor’s degree programs in Anatomy of the Common Gray Mussel.” Af - engineering, as well as a variety of graduate terward, to the amusement of all within programs in science and engineering. In ad - earshot, Porter was heard to remark: “Won - dition, it also ran a non-specialized, 3-year derful. That young man got more out of that bachelor’s degree program that included not clam than most people would get out of an only the math, science, and engineering you entire cow.” would have anticipated, but also courses in Although Sheffield was certainly the the humanities, including English, which home of much of the science being done at was not taught at Yale College in that era. Yale in the second half of the 19th century, (Wilbur Cross, the man after whom the it was not the home of all of it. The Yale Col - Graduate School medal bestowed on its lege faculty at that time included some sub - most distinguished alumni is named, taught stantial scientists, e.g., Dana in Geology and English in the Sheffield School before going the formidable J.W. Gibbs in Theoretical on to a remarkable career in politics that in - Chemistry. Nor did the two divisions of the cluded the governorship of the State of Con - University act entirely independently. There 554 Moore: The biological sciences at Yale

was some cross-listing of courses so Yale For example, in 1918, Yale was supporting College students could take courses in several separate German departments, which Sheffield, and vice versa, but nevertheless, must have seemed as lunatic then as it would an observer of the scene in New Haven today. That committee’s recommendations, around 1900 would have been astonished to which were announced in 1919, led to revo - discover a single institution running two un - lutionary changes in the way Yale is organ - dergraduate programs so totally different in ized and governed that included the character. Furthermore, it is likely that this establishment of the office of Provost. Noth - observer would have concluded that in many ing as cataclysmic has taken place at Yale ways, the Sheffield School was a much more since. lively and forward-looking institution than Among the many recommendations of Yale College. the Committee were several that taken to - gether amounted to nothing less than a call for the dissolution of the Sheffield Scientific on tHe revolution of 1919 And School. Not surprisingly, these recommen - itS AftermAtH dations met with passionate opposition for Sadly, by the first decade of the 20th reasons that in hindsight appear compelling. century, the Sheffield School’s relationship In fact, the Sheffield trustees were advised with Yale College, which had never been all by their attorney that they would be in that amicable, turned sour. About 1907, the breach of their fiduciary duties if they were University acquired Pierson Sage Square, to accede to the liquidation of the School. the area often referred to today as Science The response of the Corporation, which was Hill, and at about the same time, Yale Col - dominated by unsympathetic Yale College lege added Ross Harrison to its faculty. Har - graduates, was to reduce the Sheffield rison was a distinguished biologist who School’s sentence from instant execution to made major contributions to the develop - a slow death by a thousand cuts that contin - ment of tissue culture techniques. He was ued until about 1956. The demise of the lured to New Haven from Johns Hopkins Sheffield School as an independent compo - with an offer that included a promise that he nent of the University left behind myriad would be housed in a new building on Sci - problems having to do with the education of ence Hill ― Osborn Memorial Laboratory engineers that Yale has still not satisfactorily ― and that a new department would be set solved. The fact of the matter is that up for him: Zoology. The Sheffield School Sheffield was destroyed by the University was pressured into moving its biological that had created it, not because it had failed programs into Osborn Memorial Laboratory, financially or in any other way, but because which it had largely done by World War I. it had become too successful and too inde - Given its long and successful involvement pendent. in the biological sciences, the Sheffield In the years that followed, many of the Board might well have felt it deserved to be Departments that had been components of treated as the senior partner in this new, joint the Sheffield School were dispersed venture with Yale College, but the Corpora - throughout the University. For example, tion had other ideas. Physiological Chemistry moved to the Med - Toward the end of World War I, the ical School in 1923, where it eventually be - Corporation appointed a committee of came the Department of Biochemistry. alumni to consider ways in which the Uni - Sheffield’s botanists were integrated into a versity might be reorganized so as to im - Zoology-Botany operation in Osborn Me - prove both its governance and its efficiency. morial Laboratory. The non-physiological The Corporation had substantial reasons for parts of Sheffield Chemistry merged with concern on both counts, not the least of them the Yale College Chemistry program that being many duplications of academic effort had been educating undergraduates for about in the different divisions of the University. 30 years, and in the early 1920s, the newly Moore: The biological sciences at Yale 555

merged department moved into Sterling graduate (class of 1860) who persuaded his Chemistry Laboratory. wealthy uncle, George Peabody, to fund the establishment of a museum of natural his - tory at Yale. In 1870, Marsh, who, like some cHemiStrY And BiologY other members of the Yale faculty of his This is not the place to provide an ac - generation, was unencumbered by a Yale count of the subsequent history of the De - salary, headed off to Wyoming to collect the partment of Chemistry, but its long fossils that would ultimately fill his uncle’s involvement with the biological sciences museum. Marsh eventually became Profes - needs to be acknowledged. In the first place, sor of Paleontology, a discipline that is a it is important to point out that for many fundamental to evolutionary biology. Pale - years the phrase “organic chemistry” meant ontology remains an important part of what the chemistry of the molecules found in liv - is today the Department of Geology and ing organisms, rather than the chemistry of Geophysics, as well as being a component all compounds that contain carbon. Sec - of the Department of Ecology and Evolu - ondly, many of the Department’s faculty tionary Biology, and its association with the contributed to our understanding of bio - Peabody Museum is as strong as ever. chemistry. For example, one of the people It is also appropriate to note the contri - Chemistry obtained from the wreckage of bution made to both biology and geology by the Sheffield School was Treat B. Johnson, Bertram Boltwood in the first decades of the an organic chemist who did a lot of impor - 20th century. Boltwood, who was a tant work on the chemistry of pyrimidines, Sheffield graduate, was one of the pioneers nucleic acids, and proteins. Rudolph Ander - in radiochemistry in the United States. He son, who joined Chemistry in the mid- began his faculty career in Physics, but later 1920s, was Editor of the Journal of moved to Chemistry. Boltwood was the first Biological Chemistry for 21 years. John to date minerals by measuring the amounts Kirkwood, who was entrusted with the reor - of lead and uranium they contain. Isotopic ganization of the Department of Chemistry dating has been a critically important tool in the 1950s, was a theorist interested in, for paleontologists ever since because it en - among other things, the physical chemistry ables them to fix the fossils they collect in of biological macromolecules, and he estab - time. lished a tradition of research in biophysical chemistry within the Department of Chem - tHe BiologicAl ScienceS Since istry that continues to this day. 1920 Compared to the upheavals that oc - geologY And BiologY curred in the first two decades of the 20th When thinking about Biology at Yale, century, the history of the biological sci - one should never forget Geology. Geology ences since 1920 has been placid. By the late was the second of the sciences to be fully in - 1950s, there were three biology-related de - stitutionalized at Yale. By 1812, Yale Col - partments outside the Medical School: Bio - lege was offering its students a course in physics, Botany, and Zoology. Botany and geology that was distinct from Silliman’s Zoology shared Osborn Memorial Labora - chemistry course, and in 1850, Dana, who tory and jointly taught an introductory biol - was Silliman’s successor, became the first ogy course that was half zoology and half Silliman Professor of Geology and Mineral - botany. From the point of view of the un - ogy. From a biological perspective, the sin - dergraduate curriculum of the day, bacteria gle most important development in Geology were largely ignored because no one was in the second half of the 19th century was sure what to think about them. It had only the appointment of Othniel Marsh to the been a few years since Edward Tatum, who Sheffield faculty. Marsh was a Yale College was a member of the Botany Department, 556 Moore: The biological sciences at Yale

and his graduate student Joshua Lederberg years later, Pollard left Yale for Pennsylva - had discovered conjugation in E. coli , a find - nia State University, precipitating a crisis ing that one could argue is the single most both in leadership and scientific direction. momentous discovery ever made by Yale bi - In 1962, under the guidance of Frederick ologists. Molecular biology would not have Richards, Biophysics became the Depart - taken off the way it did in the 1950s and ment of Molecular Biology and Biophysics 1960s if there had been no bacterial genet - (MB&B), a title that better reflected its sci - ics. entific direction, but a dispute soon broke In 1962, Zoology and Botany merged to out with the Department of Biology over the form the Department of Biology, an entity propriety of MB&B’s use of the word “biol - that turned out to be short-lived. In the late ogy” in its name, and MB&B was forced to 1990s, the Provost convened a committee become MB, i.e., Molecular Biophysics. that was charged with finding a solution to MB got its second B back a few years the problems that had arisen because of the later as a result of governance problems that inability of Biology’s physiologically ori - had arisen in the Medical School’s Depart - ented faculty to agree on appointments with ment of Biochemistry. These problems were their evolutionary/ecological colleagues. solved by President Brewster by merging The upshot was that although the members Molecular Biophysics with Biochemistry to of the Provost’s committee unanimously produce a new entity that would have a pres - agreed that intellectually it made no sense ence on both sides of the campus. (This whatever to separate evolution from the rest move returned Physiological Chemistry to of Biology, from a purely practical point of the part of campus from which it had been view, it might be best if its two components so rudely ripped four decades earlier.) Since went their separate ways. So it was that the 1969, this arrangement has served Yale well, Department of Ecology and Evolutionary even though I am sure that, if asked, all of Biology and the Department of Molecular, the faculty associated with MB&B, both liv - Cellular and Developmental Biology were ing and dead, would tell you that physically born. No doubt that in another 25 years or divided departments are to be avoided so, the biological sciences departments at whenever possible. However, the notion that Yale will undergo yet another such reorgan - it might be a Good Thing to strengthen the ization. Fission and fusion events are every - ties between biologists who live in the Med - day occurrences in biology. ical School and those who reside on the cen - tral campus has obvious merit. In biology, bigger is usually better, especially for grad - BiopHYSicS uate students, provided institutions are able Biophysics is a much younger entity to manage size effectively. than any we have talked about so far. The person most responsible for bringing it into concluding remArkS existence was Ernest Pollard, a nuclear physicist who joined the Physics faculty in It seems to me that the history of the the late 1930s. After the war, he and several Sheffield Scientific School may offer an ex - of his younger colleagues became interested planation for an aspect of the culture of this in using ionizing radiation as a probe for University that had long puzzled me because studying the organization of viruses and or - it is so irrational. Even when I was a Yale ganisms. For a number of years, Biophysics undergraduate in the late 1950s, I was aware was supported by the Hartford Foundation that the non-scientific parts of Yale viewed and occupied space owned by that Founda - its scientific components unsympathetically. tion in Valhalla, New York. In 1959, Bio - The fact that unlike all of the other universi - physics returned to New Haven and moved ties Yale considers its peers, Yale has never into the sixth floor of Gibbs Laboratory, had a president who began his/her career as which had been specially built for it. Only 2 a scientist is but one manifestation of this Moore: The biological sciences at Yale 557

persistent aspect of the Yale zeitgeist. Per - biggest return on its investment? The suc - haps it is a product of the competition that cess of the biological sciences at Yale in the developed in the 19th century between Yale 21st century will be determined by the wis - College and the Sheffield School, which dom with which such decisions are made, must have looked very threatening to the and Yale is going to need all the help it can faculty of Yale College in, say, 1900. get to make them right. I invite its graduate In addition, one cannot help but specu - alumni to make their opinions known. late that had the Yale Corporation treated the Sheffield School with more respect in 1919, Acknowledgments: i gratefully acknowledge the debt i owe to Professor William sum - Yale might still be a leading school for en - mers, who happens to be the author of the gineering, as it was then. Might it be wise chapter on MB&B that appears in reference for Yale to make the investments today that (2), for his advice on where to begin my search for information about the history of bi - would be required to revitalize engineering? ology at Yale. i would have been lost without Remarkably, despite all of the trauma it has him. suffered since 1919, Yale’s engineering fac - ulty is still big enough and strong enough to referenceS nucleate such a venture. I can think of noth - ing Yale could do today that would have a Detailed references have not been in - more positive effect on its standing in the ac - serted into this essay sentence by sentence, ademic world. lest the reader be misled into thinking that Turning now to more specifically bio - all the statements made in an essay of this logical issues, it is interesting to reflect that sort can be or should be supported as they despite being forced to give up the phrase would be in a scientific article. That said, “molecular biology” in its name, at the time three works were consulted extensively in it was founded, the regular faculty of the writing of this essay. MB&B and its joint appointees did in fact 1. Chittenden RH. History of the include a large fraction of all of the mem - Sheffield Scientific School of Yale Univer - bers of the Yale faculty on both sides of the sity, 1846-1922. New Haven, CT: Yale Uni - campus who were doing molecular biology. versity Press; 1928. Today it would make no sense whatsoever The author of this marvelously inform - to organize a department around the theme ative, well-written, two-volume work was of molecular biology. The reason is that the last head of the Sheffield Scientific today, the mindset and experimental ap - School. It is a rich source of information proaches of the molecular biologist pervade about the history of the Sheffield School, its all of biology; one way or another, we are faculty, and its often troubled relationship all molecular biologists now. This obvious with the rest of Yale. Everything said about fact may trigger the next big reorganization the Sheffield School in this essay derives of the biological sciences at Yale. from this single source. Finally, as the 21st century unfolds, 2. Altman S. editor. Science at Yale. Yale will confront many issues that will bear New Haven, CT: Yale University, New directly on the professional lives of its bio - Haven; 2002. logical faculty and students. How big should Only 300 copies of this small volume the biological enterprise be allowed to grow were printed. It was published at the time of at Yale, given that outside funding is likely the University’s tercentenary to commemo - to be harder to obtain in the future than it rate the contributions made to Yale by its was between 1950 and 2000? How should science departments. It contains short his - biological research done on the West Cam - tories of the nine science departments that pus be integrated with that done elsewhere at existed in 2000 outside of the Medical Yale? Indeed, might it be better to make it School and Engineering and Applied Sci - free-standing? What parts of the biological ence. They vary a lot in breadth and style, as enterprise are the ones that will give Yale the is typical of multi-authored works. Never - 558 Moore: The biological sciences at Yale

theless, short of spending many months that the Geological Sciences at Yale. Geological I did not have to spare burrowing through Society of America, Centennial Special Vol - the University’s archives, it would have ume 1; 1985. pp. 355-370. been impossible for me to write an essay This paper explores the early history of anywhere near as complete as this one with - the earth sciences at Yale in far more detail out it. than the terse account provided in reference 3. Skinner BJ, Narendra BL. Rummag - (2). Like references (2) and (3), it was a de - ing Through the Attic; or a Brief History of light to read.