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Archival Resources for the History of Genetics & Allied Sciences

ISSUED BY THE LIBRARY & MUSEUM OF THE AMERICAN PHILOSOPHICAL SOCIETY

New Series, No. 22 October 2020

IN THIS ISSUE

From the Librarian — Patrick Spero 3

Celebrating Mendel’s 200th — M. Susan Lindee and John J. Mulvihill 4

The Allan Wilson Papers — Marina DiMarco 6

The Johannes Holtfreter Papers — Michael Dietrich 12

The Richard Lewontin Papers — Charles Greifenstein 17

THE AMERICAN PHILOSOPHICAL SOCIETY LIBRARY & MUSEUM

The Mendel Newsletter October 2020

The Mendel Newsletter American Philosophical Society Library & Museum 105 South Fifth Street Philadelphia, PA 19106-3386 U.S.A.

www.amphilsoc.org/library

Editor Managing Editor Michael Dietrich Adrianna Link Department of History and Philosophy of Science American Philosophical Society 1017 Cathedral of Learning [email protected] University of Pittsburgh Pittsburgh, PA 15260 [email protected]

Editorial Board

Garland E. Allen, Washington University Susan Lindee, University of Pennsylvania Ana Barahona, National University of Mexico Jane Maienschein, Arizona State University John Beatty, University of Minnesota Staffan Müller-Wille, Cambridge University Michael R. Dietrich, University of Pittsburgh Diane B. Paul, University of Massachusetts, Boston Kaori Iida, Graduate University for Advanced Studies, Jan Sapp, York University, Toronto Sokendai Vassiliki Betty Smocovitis, University of Florida Barbara Kimmelman, Philadelphia University

The Mendel Newsletter, New Series, No. 22 (October 2020) Issued occasionally by the Library & Museum of The American Philosophical Society. Available only in digital format, readers may print and redistribute copies. The current issue and many back issues may be downloaded from The Mendel Newsletter webpage at www.amphilsoc.org/library/mendel.

Inquiries relating to article contributions to The Mendel Newsletter may be sent to the Editor.

Photograph of Gregor Mendel, ca. 1860, from the Curt Stern Papers, American Philosophical Society. APS graphics:2210

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From the Librarian

Patrick Spero

HILE much of the world has had to close its doors due to the pandemic, I am happy to report that the Library W & Museum, while certainly disrupt- ed, has maintained an active acquisitions program. Since our last newsletter, the Society has acquired several papers that may be of interest to readers, including the papers of Francisco Ayala, Nina Ja- blonski, David Hungerford, and rare first editions of several of Darwin’s works, some of which came as recently as this summer. Longer-term, the Library & Museum just completed a two-year long review of its collection development policy. The Society has undertaken this review about every ten years. The revised pol- icy highlights the Library’s strength in genetics and identified the field as a key priority for col- lecting in the future. Its statement reads, in part, “Within the life sciences, the Library & Museum should continue focused collecting in those areas already strong – including genetics, evolution, and science, including most recently a conference en- eugenics.” The committee in charge of the history titled “Evidence: The Use and Misuse of Data.” of science collecting area also highlighted evolu- Next year, we hope to open our exhibition, Dr. tionary biology as an area of particular interest in Franklin, Citizen Scientist, delayed due to the the future. coronavirus pandemic. Our future programs in- clude an exhibition on the history of climate sci- In addition to its collection, the Library & ence in 2022 and on women in science in 2023. Museum continues to build programming around the history of science. Such programming raises I hope you all will check out our website to awareness of our collections, shares what is being see the range of virtual programs we currently of- discovered in them, and serves the Society’s mis- fer and tune into some of them. In fact, if you sion of promoting knowledge. In the past few have research from the APS’s Library & Museum years, we have added a long-term predoctoral fel- you’d like to share, you might consider presenting lowship to support research in the history of sci- at our weekly brown bag lunches. And, by all ence, and we continue to host a large slate of means, please let us know about important papers short-term resident research fellows. We have also that might be a good fit at the APS. organized a number of events on the history of

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Celebrating Mendel’s 200th Birthday July 22, 2022

M. Susan Lindee and John J. Mulvihill [email protected] [email protected]

ENDEL is a fascinating figure in the he accepted the Children’s Hospital Foundation-- history of genetics, needless to say. Kimberly V. Talley Chair of Genetics, Professor His pea experiments are known to of Pediatrics, University of Oklahoma. In 2014, the average 6th grader. His status as he became part-time consultant to the National M a monk has been used to propose Human Genome Research Institute. that religion and science are not in conflict. His Mulvihill’s research has focused on the genet- quantitative results have been critiqued as too ics of human cancer, with an emphasis on late ge- good to be true. And he is iconic, like Darwin: netic and reproductive effects in cancer survivors This is the Mendel Newsletter because Mendel’s and on germ cell mutagenesis. Elected a Director name is a legible shorthand for the entire history of the American College of Medical Genetics, he of modern genetics. is also a member of the Committee on Ethics, Johann Mendel was born July 22, 1822, and Law, and Society of the International Human Ge- died as Abbot Gregor Mendel on January 6, 1884. nome Organisation (HUGO) and a past Scientific As his 200th birthday looms, we have been think- Advisor of the Radiation Effects Research Foun- ing through options for how to recognize this im- dation, Hiroshima and Nagasaki, Japan. While he portant figure, and make sense of his historical is employed by Oklahoma, he is currently “on and modern roles in genetics, meteorology, and loan” to the National Human Genome Research agriculture. Institute of the NIH, and he lives and works in While Lindee is presumably known to many Philadelphia. readers of the Mendel Newsletter, Mulvihill prob- The commemorative year is intended to pull ably is not. His commitments are those of a clini- together an international program of education cal geneticist in tune with history who wants the and awareness for scientists and the public. Ge- broader public and the scientific community to netics as a scientific field has a history of huge appreciate Mendel’s contributions more accurate- public interest and stunning success, as well as ly and completely. historic abuses manifest in racist and eugenicist A pediatrician and medical geneticist with 20 ideas. The program for 2022 will be, in Mulvi- years' experience at the National Cancer Institute, hill’s words, “conducted responsibly, with integri- he was chief of the Clinical Genetics Section of ty, rigor, inclusion, innovation, and the Clinical Epidemiology Branch and Director of collaboration.” Attention to the historical roles of the Interinstitute Medical Genetics Program of the Mendel can become a critical resource for think- National Institutes of Health (NIH). In 1990, he ing clearly about knowledge and religious com- became founder, chair, and professor of Human munities, the origins of modern genetics and Genetics at the University of Pittsburgh. In 1998,

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agriculture, and the ethical questions raised by As Mulvihill suggests, Mendel can be held as information about heredity today. an exemplar of mentoring (in both directions, giv- ing and taking). He was a nineteenth-century sci- entist who was able to thrive at the interdisciplinary interactions of botany, evolution, mathematics, physics, and agriculture. He con- tributed to improved fruit trees, ornamental flow- ers, horticulture, meteorology, and beekeeping. He also worked as a bank director, and was popu- lar in his community of priests.

General knowledge of his work—inculcated first often in elementary school—might inspire public interest in this commemoration in ways that could productively call attention to enduring

issues in the history of genetics. A broad discus- sion of Mendel’s legacies could illuminate tech- nical, social, ethical and moral concerns in the 200 years since his birth. Historians of science have a special contribution to make to this effort and are hereby invited to suggest options and ide- as. Anyone interested should contact Susan Lindee, [email protected] and John J. Mulvihill, [email protected]. “Mendel’s Research Garden,” n.d from the Curt Stern Pa- pers, American Philosophical Society, APSimg:507. Working geneticists are an important audience for all events relating to this bicentennial of Men- del’s birth. Mulvihill has even proposed that it might be useful, for publicity reasons, to discuss sequencing Mendel’s DNA—with the resulting ethical debate itself of interest as a way of inform- ing the public and the scientific community. He has established partnerships for this program with the two Mendel Museums in Brno, CZ, with sci- entific societies including the American Society of Human Genetics and the American Association for the Advancement of Science, with Jackson Laboratory and with several universities. Roger Turner at the Science History Institute is working to develop meteorological perspectives on Mendel’s life (Mendel collected significant meteorological data seen as relevant to agricul- ture); I am working with Section L of the AAAS to propose an event at the 2022 meeting that will explore Mendel and modern genetics.

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The Allan C. Wilson Papers at the Bancroft Library, University of California, Berkeley

Marina DiMarco [email protected]

LLAN C. Wilson was a University of to pull one of his particularly long-winded California, Berkeley biochemist postdocs off a conference presentation stage whose work in molecular evolution did not help matters.)1 Although some ex- ranged from the timing of human- pected young Wilson to become a farmer or A primate divergence and the produc- a veterinarian, he became interested in more tion of Mitochondrial Eve to the isolation of general biochemical processes, and a local ancient DNA from a Siberian wooly mam- animal scientist convinced him to study zo- moth. The Wilson papers are available to ology and chemistry at the University of researchers as part of the History of Science Otago. Otago was far away and expensive, and Technology collection at the Bancroft and Wilson took his BSc. there despite a Library of the University of California, great deal of financial hardship. Berkeley. These papers are an important After graduation, Wilson traveled to the source of material regarding the twentieth US to work with David S. Farner of Wash- century development of molecular systemat- ington State University and to complete a ics, human genetics, the Human Genome MS in zoology working on hormonal regula- Diversity project, and the history of the Uni- tion of bird behavior. From there, he went versity of California. on to his doctoral work at the University of California, Berkeley, where he studied under Early life and training Arthur Pardee. Pardee was a former student Wilson was born in Ngaruawahia, New of Linus Pauling, perhaps best known for the Zealand, on October 18, 1934 and grew up famous 1959 PaJaMo (Pardee, Jacob, and on the family dairy farm in Helvetia. Early Monod) paper on gene regulation in bacte- on, he honed his talents for biological in- ria. After a postdoc with Nathan Kaplan at quiry by careful observation of local birds Brandeis, Wilson returned to Berkeley to set (including a nearby ostrich farm) and by up his own lab in the biochemistry depart- working to improve the family’s cattle herd. ment, where he would spend the rest of his Despite these bovine origins, his proud sta- academic career. tus as a New Zealander provoked a lifetime of sheep jokes and pranks from students and colleagues. (Wilson’s use of a shepherd’s crook 1 Cann (2014), p. 469.

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Proteins, primates, and molecular clocks alone.” Similarly, Wilson and Sarich (1969) Wilson was frustrated by what he per- argued that their molecular clock conferred ceived as the failure to synthesize molecular greater certainty and objectivity than the biochemical and population genetic ap- comparatively fragmentary fossil record, proaches to the study of evolution. This frus- and that a quantitative, molecular method tration persisted throughout his career and should resolve controversy among anthro- motivated much of his work in the emerging pologists, paleontologists, and geneticists with regard to the timing of human-primate field of Molecular Evolution. In his first 3 year as a professor of biochemistry, Wilson divergence. met Vincent Sarich, an anthropology PhD Such consensus was not forthcoming. student with whom he began work on an Instead, this work provoked the first of immunological clock for primate evolution. many controversies throughout Wilson’s Sarich and Wilson’s clock built on Emile career, as it contradicted other established Zuckerkandl and Linus Pauling’s 1965 paleoanthropological theories (for instance, comparison of hemoglobin alpha chain se- the status of Rampithecus ape jaw fossils as quences, which assumed a constant, univer- a hominid).4 Paleoanthropologists perceived sal rate of substitution in the amino acid Wilson and Sarich’s work as an arrogant sequence in order to estimate the time dismissal of their tradition, and they were elapsed between evolutionary events from not alone: creationists were also unhappy observed sequence differences.2 Using se- about the biochemical demonstration of sim- rum albumin immunological reactivity to ilarity between humans and apes.5 quantify differences between homologous Despite (or, more likely, because of) this proteins in human and primate populations, controversy, Wilson continued to work on Wilson and Sarich (1967) put the divergence primate evolution, and together with student between humans and primates at about 5 Mary-Claire King, he quantified the se- million years prior. Wilson saw this as a quence similarity of human and chimpanzee methodological unification of molecular and proteins and DNA. In another landmark pa- population genetic approaches. per in 1975, King and Wilson proposed that This use of molecular clocks had more gene regulatory differences, rather than se- quence differences, explained the biological in common with Zuckerkandl and Pauling’s 6 early work on molecular clocks than mere differences between the two. method. In “Molecules as Documents of Evolutionary History,” (1965) Zuckerkandl Mitochondrial Eve and Pauling argued that molecules contained Wilson wanted to extend the molecular the greatest amount of a living organism’s clock to provide a temporal framework for evolutionary history in an information theo- retic sense, and that molecular semantides (DNA, RNA, and polypeptides) were the 3 For more on the status of “informational molecules” most certain or least “cryptic” sources of and the development of molecular evolution as a dis- information about evolutionary history. cipline, see Dietrich (1998), Suarez-Diaz (2007; 2009) and Sommer (2008; 2016). They argued that the “most rational, univer- 4 Klein and Takahata (2002), p. 238-239. sal, and informative molecular phylogeny 5 Allan Wilson to Susan Smith, December 16, 1969. will be built on semantophoretic molecules Allan Wilson Papers, Bancroft Library, University of California Berkeley (BANC MSS 95/22 c), microfilm 2 For more on Zuckerkandl and Pauling and the de- edition (hereafter AWP), Series 3, Carton 5, Reel 13. velopment of molecular clocks, see Morgan (1998). 6 King and Wilson (1975).

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evolution, and this required new clock in- tion from scientific criticisms, where advo- puts. Because mitochondrial DNA (mtDNA) cates of the competing multiregional hy- was a small molecule with a relatively rapid pothesis used it to dismiss the result (which rate of mutation, mtDNA clocks made it arguably contradicted their theory), and oth- possible to view evolutionary time in com- ers argued that the population genetic as- paratively high resolution. Furthermore, sumptions needed to build and root the tree mtDNA was thought to be exclusively inher- were suspect. In popular press, African Eve ited through the maternal line, and not sub- became a global celebrity, and much of the ject to recombination; Wilson and discussion centered around the notion of Eve colleagues thought this relatively simpler as a first woman who was not just African, mode of inheritance made it a more reliable but Black. This reflected at least one com- basis than nuclear DNA for evolutionary mon misconception about Mitochondrial inference.7 When Wes Brown joined the Eve, who was by no means the first woman, Wilson lab in the late 1970s, he brought but merely the last one to have her mito- with him from Caltech new techniques for chondrial DNA survive in the population. It isolating and amplifying mtDNA.8 Building was also symptomatic of the reproduction of on Brown’s (1980) work and new computa- race in molecular evolution, where (despite tional technologies like David Swofford’s Cann’s protests) such trees were taken by Phylogenetic Analysis Using Parsimony some to show evidence of biological race (PAUP) algorithm, Wilson’s student Rebec- differences, and where competing theories ca Cann built a restriction fragment length of human evolution routinely charged their polymorphism (RFLP) mtDNA tree that adversaries with racism.10 identified an African single most recent 9 Cann, Stoneking, and Wilson worked to common mitochondrial ancestor. Mark address these criticisms and misconceptions, Stoneking, another Wilson student, analyzed notably trying to rebrand Mitochondrial Eve samples from Papua New Guinean popula- as a “lucky mother,” with little success. Alt- tions to further calibrate the tree. In 1987, hough Mitochondrial Eve and the “Out of Cann, Stoneking, and Wilson published Africa” hypothesis are often touted as high what became known as the “Out of Africa” points of Wilson’s research career, neither tree, featuring an ancestor who quickly be- Cann nor Wilson saw Mitochondrial Eve as came known first as “African,” and, later, as entirely successful. Later work by Ston- “Mitochondrial Eve.” eking, Linda Vigilant, and other students in Much like the human-primate diver- the Wilson lab lent additional support to the gence paper two decades before, this work Out of Africa model using new sequence sparked immediate scientific and popular data and increasingly sophisticated computa- controversy. At first, Wilson embraced the tional methods, and after several years it was “African Eve” branding, hoping it would more widely accepted.11 Meanwhile, Wilson encourage others to take seriously the possi- worked to promote mtDNA techniques in bility that all modern humans originated in his own lab and in his role as an associate Africa. Later, however, it became a distrac- 10 Notably, these included Wilson’s former student

7 and collaborator Vince Sarich, whose teachings and For more on early nuclear DNA research in molecu- writings on race sparked outrage and protests on the lar evolution, see Marianne Sommer’s account of the Berkeley campus (“Campus LIfe”). Sarich’s book, Cavalli-Sforza lab in History Within (2016). Race: The Reality of Human Difference is dedicated 8 Brown and Vinograd (1974). to Wilson (Sarich and Miele 2004). 9 Brown (1980), Cann et al. (1987). 11 Vigilant et al (1991).

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editor of the Journal of Molecular Evolu- important starting point for molecular paleo- tion. priests. anthropology, as well as for many forensic and medical applications. modern genetics. Ancient DNA and PCR Of immediate relevance was the im- In addition to his work to establish portance of PCR for improving HIV testing mtDNA as a basis for molecular clocks, and treatment in the late 1980s. At the Wilson was also at the forefront of devel- height of the Reagan administration’s esca- opment of techniques for isolating and am- lation of the Cold War and its indifferent plifying so-called “ancient DNA,” such as approach to the AIDS crisis, Wilson worked the mtDNA sequences from an extinct on PCR technologies that would facilitate quagga and samples from a wooly mam- HIV research, corresponded about collabo- rating with Cubans to identify human re- moth.12 Because Wilson had close connec- tions to Cetus, he and his students not only mains that were thought to belong to helped to develop and calibrate polymerase Christopher Columbus, and routinely pro- tested nuclear weapons research policy at chain reaction (PCR) protocols, but they 16 were also among the first to bring PCR to Berkeley. Though Wilson is often de- bear on problems in molecular evolution. scribed as a reserved character, he had no Together with Russell Higuchi and Svante reservations about advocating for the inclu- Pääbo, Wilson worked to isolate DNA from sion of women as invited conference speak- museum collection specimens, including not ers (in a scathing letter to program only the quagga but also many birds, extinct organizers, he points out “Your program has zebras, and a 7,000 year-old human brain.13 no women”). He also had no trouble refus- (Pääbo recalls an early prototype of a PCR ing to give a talk where he found the invita- tion of other speakers (South African machine made with washing machine valves 17 routinely flooding the Wilson lab.)14 Though scientists) politically distasteful. Wilson is widely celebrated for the new mo- lecular technologies he brought to bear on The Human Genome Diversity Project the study of evolution, in this sense, Wil- Ancient DNA did not distract Wilson son’s approach was more continuous with from his work on the relationships among the natural historical tradition in biology, contemporary human populations, where his and many of his students went on to ap- interests expanded to include the evolution pointments in natural history museums as of cognition and the possible relationships well as biology research (although Wilson’s between evolution, language, and the brain. opportunistic approach to sample acquisi- In a 1989 presentation to the Merry Evolu- tion, which included filling freezers with tionists informal discussion group, provoca- roadkill and instructing his young daughter Ruth in the art of getting New Zealand frogs tational traditions in biology, see Strasser (2019). For through international customs on his behalf, more on the frogs, see Allan Wilson to Ruth Wilson, might have appalled some natural histori- August 21, 1971, and Allan Wilson to Customs Offi- ans).15 This work on ancient DNA was an cials, Auckland Airport, New Zealand, August 20, 1971. AWP Series 3, Carton 6, Reel 13, Frame 289. 16 Schmeck (1985), Jonathan Ericson to Allan Wil- 12 Higuchi et al (1984). 13 son, April 21, 1987, AWP Series 3, Carton 8, Reel Zagorski (2006). 21; Allan Wilson to Jonathan Ericson, October 19, 14 Andrews (2008). 1987, AWP Series 3, Carton 8, Reel 21. 15 See Cann (2014), p. 467. For more on the continui- 17 Allan Wilson to Dr. Wilfried W. deJong, Novem- ty of the natural historical, experimental, and compu- ber 5, 1987. AWP Series 3, Carton 8, Reel 21.

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tively titled, ``Eve and the Chomsky Muta- part of his scientific legacy. His influence tion,” Wilson drew on Luigi Cavalli- extends via his dozens of graduate students Sforza’s work to hypothesize that the devel- and postdocs, many of whom have made opment of language could explain the migra- significant scientific contributions of their tion out of Africa suggested by his mtDNA own, and also by way of the California bio- work with Cann, Stoneking, and others.18 technology industry, of which he was a ma- Though they had many intellectual disa- jor advocate.21 greements, Wilson and Cavalli-Sforza joined forces to advocate for the Human The Wilson Papers Genome Diversity Project, an effort to col- Dr. Ellen Prager organized Wilson’s sci- lect and preserve DNA samples from isolat- entific papers for donation to the archive by ed populations around the world. While Leona Wilson in 1994. Many papers and Wilson, Cavalli-Sforza, and others saw this folders contain Prager’s notes, which are as a liberatory project which would celebrate often very informative. The Wilson papers human diversity, promote scientific inquiry, contain 20 linear feet of material: 15 cartons, and move away from a Eurocentric concep- a box, one volume, and an oversized folder. tion of the human genome, many indigenous Most of this material has been transferred to people and other advocates were skeptical of 50 reels of microfilm, which is the primary its anti-racist advertisement and critical of 19 means of accessing the collection. Materials its implementation. are arranged chronologically within each Wilson did not live to participate directly series, with notes from Dr. Prager estimating in this final controversy. In late 1990, Wil- dates when they are absent. Papers Dr. Prag- son was diagnosed with leukemia, and he er deemed confidential, such as letters of died July 21, 1991 while undergoing treat- recommendation, have been removed. ment in Seattle. His public call for the The collection is arranged chronological- HGDP appeared posthumously in October 20 ly within several series, which are organized 1991. Wilson’s former student, prolific as follow. Series 1 contains publications research collaborator, and longtime lab 1957-1998. Series 2 contains materials relat- manager, Dr. Ellen Prager, wrapped up op- ed to manuscript preparation 1962-1992 (in- erations in his lab. Wilson’s many important cluding rejected manuscripts with reviewer methodological and theoretical contributions comments that may never have been pub- to molecular evolution and numerous pres- lished). Series 3 includes Wilson’s profes- tigious fellowships and awards are only one sional correspondence 1959-1991; in addition to his students, Wilson’s corre- 18 Allan Wilson, lecture notes, March 31, 1989. AWP spondents included Mayr, Howell, Series 8, Carton 14, Reel 41, Frame 435. This lecture Lewontin, Dayhoff, Zuckerkandl, McCar- immediately followed Cavali-Sforza’s address to the Merry Evolutionist society, a series which Wilson co- thy, Fitch, Margoliash, Nei, Ohta, and many organized with Hy Harman. other major figures of twentieth century bi- 19 See M’Charek (2005 ab), Sommer (2016), and ology. Series 4 includes sabbaticals and spe- Reardon (2017). As Reardon (2017) notes, the HGDP cial trips 1967-1989 and related became known as the Vampire Project because it was correspondence (perhaps most notably, a perceived as being more interested in the blood of reassuring note from home that his lab had indigenous people as research subjects than it was in the survival of indigenous communities. The contro- not burned down in his absence during versy sparked by the HGDP has had lasting effects on both genome research and indigenous communities. 21 Allan Wilson to Senator John Garamendi, June 29, 20 Cavalli Sforza et al (1991). 1988. AWP Series 3, Carton 8, Reel 21.

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Berkeley campus riots of 1969). Series 5 Accessing the Collection contains notes from lectures 1962-1990, More information on the Wilson Papers is both at Berkeley and elsewhere. Series 6 available online: contains teaching materials 1965-1991. Se- https://oac.cdlib.org/findaid/ark:/13030/kt95 ries 7 contains documents from Wilson’s 8035w4/ education 1953-1962. Series 8, labeled “Pro- fessional 1956-1991,” contains information on various career awards and honors, but The Wilson papers are located at the Ban- also any publicity that Wilson kept about the croft Library of the University of California lab, including several international news Berkeley in Berkeley, CA. Because some clippings. Series 9 contains material from material may be stored offsite, it is im- Wilson’s early career (1959-1971), and Se- portant to contact the archives before visit- ries 10 contains research materials from ing. Bring a USB drive to store microfilm 1965-1990. The final Series 11 contains scans on pain of being forced to buy one notes, articles and letters concerned with with a Berkeley logo on it. Wilson’s death and the fate of the lab, 1990- 1994.

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The Johannes Holtfreter Papers at the Marine Biological Laboratory

Michael R. Dietrich [email protected]

HE archives of the Marine Biological pore lip had been placed. Spemann’s group de- Laboratory in Woods Hole, MA have termined that transplanted tissue had influenced made the papers of embryologist Johan- the surrounding tissue to differentiate and form nes Holtfreter available to researchers. the second body axis. As a result, Spemann T While the correspondence series in this dubbed the dorsal blastopore lip region the organ- collection has been processed, other parts are not izer. yet fully processed. Nevertheless, this collection Holtfreter would eventually engage in organ- represents a very significant source of material izer research, but he did not share Spemann’s regarding the rise of experimental embryology in temperament or work habits. Upon graduation the twentieth century. organizer. Spemann urged Holtfreter to continue his research Holtfreter’s Life and Work at the Naples Zoological Station. Holtfreter went Born in 1901 in northern Germany, Hans to Italy, but instead of pursuing embryological Holtfreter would become one of the most influen- research, he wandered away from to the island of tial experimental embryologists of the twentieth Ischia in the Bay of Naples, where he painted century. The son of a whiskey manufacturer, Holt- frescoes in the church in St. Angelo. Although freter studied at the Universities of Rostock and Holtfreter was a very talented artist, he could not Leipzig before arriving at the University of Frei- support himself with his art. burg where he studied under the renowned em- When he returned to Germany, having ex- bryologist, Hans Spemann. hausted his personal funds, Holtfreter discovered In 1924, while Holtfreter finished his doctor- how difficult it would be to find a position in sci- ate on organogenesis in frog embryos, Hilde Proe- ence without Spemann’s full endorsement. Teach- scholdt was performing a series of transplantation ing high school or becoming a fishery biologist on experiments that would lead Spemann to articu- the remote island of Helgoland seemed like his late his concept of the organizer. Proescholdt had only options. Fortunately, in 1928, Otto Mangold, taken cells from an amphibian embryo at the blas- who had been a student of Spemann’s and Hilde tula stage just before gastrulation occurred and Proescholdt’s husband before her untimely death, created different tissue layers and a body axis and had an opening at the Kaiser Wilhelm Institute for transplanted them into another location on a sec- Biology in Berlin-Dahlem. He offered the job to ond embryo. The second embryo then developed Holtfreter, before consulting Spemann. Holtfreter two separate body axes: one at the usual site and accepted immediately and began one of the most the second where the transplanted dorsal blasto- scientifically productive periods of his life. In

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Mangold’s group, Holtfreter returned to the or- tory at Yale University. Harrison and Spemann ganizer. Unlike Spemann, who preferred to think were collaborators and friends. Holtfreter knew of the organism as a whole, Holtfreter became in- Harrison and had agreed to study tissue culture in terested in the specific developmental potential of his lab when he accepted the fellowship. Howev- different regions in isolation from the rest of the er, when he got to New Haven, he was much more embryo. Using the glass needle technique perfect- interested in exploring New York city. Eventual- ed in the Spemann group, Holtfreter divided blas- ly, he left Harrison’s lab and traveled across the tulas into many different parts and then followed United States and then on to Hawaii and Bali. the development of each region as it grew in isola- Holtfreter lingered on Bali, which had a profound tion. These thousands of tissue explants had to impact on him personally. When his funds ran survive long enough for any potential differentia- low, he reluctantly completed his circumnaviga- tion to be detected. Up to this point, embryos had tion of the globe to return to Germany. been cultured in filtered water where they grew While Holtfreter had been immersed in his poorly and tended to become infected and die. In journey, Germany had undergoing its own trans- order to create better growing conditions, Holt- formation as the Nazi Party consolidated political freter experimented with different saline solutions power throughout the 1930s. Back in Germany, until he found a mixture that allowed embryonic Holtfreter felt alienated, but he would soon find tissue to survive and thrive. This solution became himself in more serious trouble. In 1938 accepted known as Holtfreter solution and was a crucial an invitation to speak at the Congress of Physics, innovation that allowed Holtfreter and other em- Chemistry, and Biology at the International Expo- bryologists to perform successful transplantation sition in Paris. As a German participant, the Ger- experiments. man government expected him to make hi As he explored the fate different regions, presentation in German. Instead Holtfreter gave Holtfreter also began to question what it was his paper in French. To make matters worse, he about regions, such as the organizer, that allowed did not return immediately to Germany. With the them to have such influence on surrounding tis- honorarium from the Congress, he explored Alge- sues. Spemann thought the organizer’s properties ria. The German government took notice and up- rested in features of its structure. Holtfreter on his return, he was denounced and imprisoned. thought that the organizer’s effects may have Joseph Needham and many other friends arranged nothing to do with living tissue and performed a for his emigration. In 1939, he joined Needham’s series of transplantation experiments with killed group at Cambridge University. Because he left organizer tissue. When this killed tissue induced for political reasons, Holtfreter was considered to the same effects as living tissue, Holtfreter con- be an enemy alien and was interred in a Canadian cluded that the organizer’s influence must be the prison camp until 1942. result of an inducing substance—a chemical that Holtfreter laws unable to do research as a diffused through surrounding tissue and deter- prisoner and quickly became embittered. Using mined its fate. This discovery started embryolo- wooden shingles from the sides of the building in gists on a search for inducing substances and in which he lived, he painted landscapes of his “Ba- effect began research on biochemical embryology. linese dreamland.” Realizing the great talent im- Holtfreter’s success at the Kaiser Wilhelm In- prisoned nearby, officials at McGill University in stitute was widely recognized. In 1934, he became Montreal began to arrange for the release of aca- an associate professor in the Zoology Department demic from the prison camps, John Berrill cam- at the University of Munich. A year later, a Rock- paigned for Holtfreter’s release and in 1944 efeller Foundation Fellowship and a private travel Holtfreter joined the faculty at McGill. grant allowed him to join Ross Harrison’s labora-

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Free to pursue embryology again, Holtfreter 1946, Holtfreter took a position in Rochester, returned to the biology of gastrulation. The beau- where he would stay until his death in 1992. tifully coordinated movements characteristic of In Rochester, Holtfreter continued his work on gastrulation had been demonstrated by Walter the experimental embryology of amphibians. Vogt ‘s technique of vital staining. Following Most notably, he returned to an observation he Vogt, Spemann had seen gastrulation as the result had made in Germany about the affinity of differ- of organism level organization and control. Holt- ent tissue types for each other. Ectoderm and en- freter rejected this approach in favor of a more doderm did not have a strong affinity for each mechanistic perspective that emphasized physical other, but mesoderm had a strong attraction to forces and the actions of specific cells. By ex- both ectoderm and endoderm. As cells became planting different groups of cells, Holtfreter de- more differentiated this affinity grew stronger. In veloped a new explanation of the mechanics of the 1950s Holtfreter and his student Philip gastrulation based on the elongation of bottle cells Townes systematically studied this phenomenon to create an invagination and the surface tension of affinity and selective adhesion of cells. Townes produced by the coating of the blastula, which and Holtfreter created mixtures of cells from dif- then drew in surrounding cells. The apparent co- ferent tissue layers and noted how they reassem- ordination of cell movement was thus explained in bled. They found that cells formed distinct tissue physical-mechanical terms. layers based on their affinities. This preferential At McGill, Holtfreter continued to explore the movement and association led Holtfreter and physical and chemical influences on embryogene- Townes to conclude that cellular processes creat- sis, especially the processes of neural develop- ed the organization evident in the tissue layers of ment in early embryos. In 1941, Lester Barth an embryo. demonstrated altering the concentration of salt in While Holtfreter’s research had a tremendous a solution could induce ectoderm to develop into impact, in the United States, he was also partly neural tissue. Holtfreter extended Barth’s results responsible for the spread of the techniques of ex- by systematically adjusting the salt concentration perimental embryology developed in the Spemann and pH of solutions with presumptive neural ecto- group. In the early 1950s, Holtfreter worked derm. Not only did Holtfreter induce this tissue to closely with his closest friend, Viktor Hamburger, form neural tissue, they also formed rudimentary to write the definitive overview of research on sensory organs. Because the tissue and solution amphibian development for The Analysis of De- used in these experiments did not include any or- velopment. Holtfreter and Hamburger had been ganizer tissue, Holtfreter claimed that they friends since their student days in Spemann’s lab. demonstrated that neuralization was independent Their joint chapter summarized the state of the art of the organizer and that, in fact, the salt solution in experimental embryology as it reviewed the facilitated autoneuralization by inhibiting the sup- results that earned Spemann the Nobel prize, that pression of the capacity for neuralization inherent launched biochemical embryology, and that made in the ectoderm. transplantation experiments the hallmark of exper- While McGill freed him from an internment imental embryology in the twentieth century. camp and allowed him to return to research, Holt- In 1968, Holtfreter retired from the University freter was never very happy there. One of Holt- of Rochester. He spent his remaining days travel- freter’s close friends from Berlin, Curt Stern, had ing with his spouse, Hiroka Ban Holtfreter, as immigrated to the University of Rochester in New well as painting and sketching until he lost his York. Eager to move to the United States, in sight near the end of life.

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Johannes Holtfreter’s sketches of Amblystoma development. March 23 and 24, 1928. Holtfreter Papers, Marine Biological Laboratory Archives, Woods Hole,

Holtfreter’s Papers from family members from the 1930s, as well as In 2008, the Holtfreter collection was deliv- letters regarding Holtfreter’s immigration to Eng- ered to the Marine Biological Laboratory in 19 land and the United States. boxes. Historian Garland Allen worked with Holt- freter’s widow, Hiroko, to donate selected profes- Notes sional correspondence, extensive laboratory notes, This collection contains an impressive set of some manuscript drafts, as well as several 16 mm laboratory notes. Holtfreter was forced to leave films and a projector for viewing those films. Cambridge, England very quickly when he was Holtfreter’s art was distributed to collections in sent to a Canadian internment camp and entrusted Rochester, NY and Richtenberg, Germany. his personal belongings to Hal Waddington. As This collection is subdivided into correspond- Holtfreter notes in his letters, Waddington was ence, experiment notes, experiment illustrations, less than diligent in keeping track of Holtfreter’s photographs, and films. The correspondence se- things. Some of Holtfreter’s personal belongings ries has been curated and organized chronologi- were lost, but not his laboratory records from cally. Some notable correspondents have their Germany. own folders. Other parts of the collection will be Beginning with five binders of notes from processed at a later date. The collection has 16 1928 to 1933, Holtfreter’s notes detail his experi- linear feet of materials. mental work on amphibian embryos, often hour by hour. These notes include his very important Correspondence work on transplantation and explants. With some Most of the correspondence in this collection breaks, these notes continue from 1940 through is from the post-war period. Notable correspond- the 1960s. Richly illustrated, these handwritten ents include Holtfreter’s close friend, Viktor notes include descriptions in German and English, Hamburger, Otto Mangold, Robert Briggs, Tuneo often with abbreviations. Yet, one can still pick Yamada, Katsuma Dan, C. H. Waddington, Curt out Holtfreter’s experiments on gastrulation, sur- Stern, Richard Goldschmidt, and Jane Oppenhei- face tension, neurulation, cell inclusion, cell ag- mer. There are some significant letters to and gregation and cell movement.

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oirs of the National Academy of Science 73 Films (1998): 3–22. The Holtfreter collection includes 44 films and Gilbert, Scott, ed. A Conceptual History of Mod a film projector. I have not reviewed all of these ern Developmental Biology. New York: Ple- films. Many seem to be from Holtfreter’s trip to num Press, 1991. Japan in the 1960s when he was researching the Hamburger, Viktor. The Heritage of Experimental movements and aggregation of slime molds. Oth- Embryology: Hans Spemann and the Organiz- ers may be of his experiments on Amoebic er. New York: Oxford University Press, 1988. movements in the 1970s. Holtfreter, Johannes with H. Bautzmann, Hans Spemann, and Otto Mangold. “Versuche der Subject Files Analyse der Induktionsmittel in der Embryo- nalentwicklung.” Naturwissenschaften 20 When the collection was donated, it contained (1932): 971–974. a few folders organized by subject. These includ------“Veränderung der Reaktionsweise im altern ed folders on Animals, Appointments, Habilita- den isolierten Gastrulaektoderm.” Roux’ Ar- tion, Immigration, Internment, and Travel. chive für Entwicklungsmechanik 138 (1938): Holtfreter also collected a file of newspaper clip- 163–196. pings in German, Japanese, and English. In his ------“Differenzungspotenzen isolierter Teile der later years, he also engaged in some biographical Anurengastrula.” Roux’ Archive für Entwick- reflections and was the subject of some biograph- lungsmechanik 138 (1938): 657–738. ical profiles. Drafts of this biographical material ----- “A Study of the Mechanics of Gastrulation.” are also included in this collection. Parts 1 and 2. Journal of Experimental Zoolo- gy 94 (1943): 261–318; 95 (1944): 171–212. Accessing the Collection ----- “Neural Differentiation of Ectoderm through More information on the Holtfreter Collection Exposure to Saline Solution.” Journal of Ex- is available online at: perimental Zoology 95 (1945): 307–340. http://archives.mblwhoilibrary.org/repositories/3/r ----- “Neuralization and Epidermization of Gastru esources/234. la Ectoderm.” Journal of Experimental Zoolo- gy 98 (1945): 169–209. The Marine Biological Laboratory Archives is Holtfreter, Johannes and Viktor Hamburger. located in the Lillie Building at the Marine Bio- “Amphibians.” In Analysis of Development, logical Laboratory at Woods Hole, MA. Because edited by B. H. Willier, P. A. Weiss, and V. some material may be stored offsite, it is im- Hamburger, 230–296. New York: W. B. portant to contact the archives before visiting. Saunders, 1955. Keller, Ray. “Holtfreter Revisited: Unsolved Hours: 8 am to 5 pm, Monday through Friday Problems in Amphibian Morphogenesis,” De- Contact: [email protected] velopmental Dynamics 205 (1996): 257–264. Phone: (508) 289-7341 Steinberg, Malcolm, and Scott Gilbert. “Townes and Holtfreter (1955): Directed Movements Further Reading and Selective Adhesion of Embryonic Am- Gerhart, John. “Johannes Holtfreter’s Contribu phibian Cells.” Journal of Experimental Zool- ogy 301A (2004): 701–706. tions to Ongoing Studies of the Organizer.” Townes, P. and Johannes Holtfreter. “Directed Developmental Dynamics 205 (1996): 245– Movements and Selective Adhesion of Em- 256. bryonic Amphibian Cells.” Journal of Exper------. “Johannes Holtfreter.” Biographical Mem imental Zoology 123 (1955): 53–120.

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The Richard Lewontin Papers at the American Philosophical Society

Charles Greifenstein [email protected]

T has been fun to write pieces for the truly revolutionized population genetics by survey- Newsletter as it lets me delve deeply into ing loci in Drosophila pseudoobsucra using gel I an important collection, recently acquired. electrophoresis and demonstrating that many loci Examples of these were the John M. Opitz pa- are polymorphic. pers (no. 20, 2015) and the Arno Motulsky pa- pers (no. 16, 2007). The coronavirus crisis having forced the closing of the APS, I have not been to the Library since March, which puts me in an unfortunate position; I can’t examine any collection. Nevertheless, I thought it worth mak- ing a stab at chronicling a recent acquisition: the Richard Lewontin papers. I remember enough about the collection, which I retrieved from Harvard in December 2018, to relate some sense of it, and there are particular circumstances about the papers that should be pointed out. Lewontin was one of Dobzhansky’s students at Columbia, where he received an MA in math- ematical statistics and a PhD in zoology. After Richard Lewontin in his laboratory at Harvard University. No appointments at several institutions, Lewontin date. From personal collection of Michael R. Dietrich. Copied with permission from R. C. Lewontin. ended up at Harvard, where he spent the rest of his career. Now 91, he has emeritus status. For a population geneticist, Lewontin is widely known, and became more so as his writing and Training in mathematics provided Lewontin speaking developed into social commentary. His with the skills to make important contributions profile rose with such work as his on race, arguing to population genetics and evolutionary biology. in a paper, for instance, that race classification is of With Ken-Ichi Kojima he developed equations little meaning given its small part in human genetic for change in haplotype frequencies and natural variability. He has also argued that organisms do selection at two loci. His paper with Jack Hubby not exist passively in their environment but also ac-

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tively shape the environment for future genera- creation. University records pertaining to individu- tions. His most visible public activity is as be- als, including student and employee records, are ing a critic of social biology, which explains closed for a minimum of 80 years.” While “Person- behavior and social arrangements as evolution- al archives of faculty, administrators, students, and arily advantageous. Lewontin questions the her- alumni/ae may be subject to restrictions established itability of human traits, such as intelligence, by the donor. University records in personal ar- faulting IQ testing as an inadequate measure of chives are subject to the same 50- and 80-year re- it. Germany. strictions on access required by University policy.”1 The papers at the APS cover Lewontin’s Folders were removed according to 28 criteria.2 long and significant career. They in fact arrived Each folder removed was marked with a form indi- at the APS in more than one accession. The first cating the folder title and the reason it was removed. (4 linear feet) was in 1979 when he deposited As far as I know, the contents of each folder were copies of a good deal of important correspond- not fully vetted before its removal. A folder, for in- ence. Familiar names abound: Crow, Delbrück, stance, could have had a name on it of an individual Dobzhansky, Haldane, Mayr, Neel, Wright. who worked in Lewontin’s lab and later was a cor- Two smaller of accessions assorted material and respondent and collaborator. That folder would be cassette recordings arrived in 2003. I drove a removed as it could contain employee or fellowship van up to Cambridge to pick up the rest in 2018. records. If a folder was marked “Oa-Op” and there was something in the folder meeting the criteria, the The papers were kept in a room Vanserg whole folder was removed without examination. Hall, pretty much by themselves. There were 11 file cabinets and assorted banker’s boxes. With The APS usually allows access to papers when the help of Steve Orzack and Diane Paul, I they arrive, as long as they have been examined and packed up 108 linear feet of material. are partially organized. Because the Lewontin pa- pers that were kept in the cabinets were boxed in One cabinet held manuscripts of Lewontin’s original order and were arranged alphabetically, ac- work. I have not examined these files specifical- cess is possible through the Curator of Manuscripts. ly, but they appear pretty comprehensive. There were many draws with copies of the printed work. About 5 cabinets held reprints of others’ 1 See https://library.harvard.edu/how-to/access-materials- work, a huge amount. I would have preferred to harvard-university-archives. 2 Noted on a document sent to Charles Greifenstein by Har- go through these discard photocopies and com- vard University Archives, August 2018. monly-available reprints, but time did not per- mit. At any rate some reprints now are growing in value, so examining them could yield surpris- es. They were removed from their hanging fold- ers and kept in alphabetical order. The banker’s boxes held some of Lewontin’s early research notebooks. Some four cabinets held correspondence, the meat of the papers. There is much interesting material in the correspondence, but folders have been removed by the Harvard University Ar- chives. University policy states that “access to University administrative records [is restricted] for a period of 50 years from the date of their

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