Ramblin’ and resonating with Eric- an unlikely but enduring bond

Ramblin’ and resonating with Eric- an unlikely but enduring bond

Author’s Accepted Manuscript Ramblin’ and resonating with Eric- an unlikely but enduring bond Thoru Pederson www.elsevier.com/locate/developmentalbio...

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Author’s Accepted Manuscript Ramblin’ and resonating with Eric- an unlikely but enduring bond Thoru Pederson

www.elsevier.com/locate/developmentalbiology

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S0012-1606(16)00020-8 http://dx.doi.org/10.1016/j.ydbio.2016.01.025 YDBIO6995

To appear in: Developmental Biology Cite this article as: Thoru Pederson, Ramblin’ and resonating with Eric- an unlikely but enduring bond, Developmental Biology, http://dx.doi.org/10.1016/j.ydbio.2016.01.025 This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting galley proof before it is published in its final citable form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

Ramblin’ and resonating with Eric- an unlikely but enduring bond

Thoru Pederson

a

a,*

Program in Cell and Developmental Dynamics, Department of Biochemistry and Molecular

Pharmacology, University of Massachusetts Medical School, Worcester, MA 01605.

* Corresponding author. E-mail address: [email protected]

“Here’s to you, my ramblin’ boy…” (Tom Paxton)

That’s a quirky title but so was our friend plus it seeds what I want to record. By “ramblin” I mean that my friend was rough-hewn in many ways (as the dropped “g” conveys) and also that he had a polymath mind (“rambling” as “searching”) that effortlessly swept across many corridors of knowledge (vide infra). By “resonating” I mean that whereas he was as profoundly different from me in almost all ways as can be imagined, yet in all our years as friends we did indeed resonate so gloriously, coming to us as a surprise at first, and then evolving from our first instant affinity to a powerful covalent bond that was one of the deepest and most cherished relationships in my life in science, though one “unlikely” from first principles. I first tuned into Eric in 1964, when I attended a symposium on Differentiation and Development sponsored by the New York Heart Association at which Alfred Mirsky, Eric’s PhD mentor at Rockefeller, introduced the session on Regulation of Gene Expression, one of the most memorable talks (Mirsky, 1964). I was amazed that a protein biochemist who previously (with Linus Pauling) had made a breakthrough about hemoglobin denaturation-renaturation, was now an erudite raconteur of an entirely different field, gene regulation in development. So after returning after the meeting I went once or twice a week to the Syracuse University Department of Zoology library (which is how a student followed the literature in those days- imagine!)

watching for papers from the Rockefeller group, and there I “met” Eric for the first time (e.g. Davidson, Allfrey and Mirsky, 1964). A few months later I took the 1965 Embryology Course, directed by Carnegie’s Jim Ebert and with Don Brown also on the faculty. With Mirksy’s 1964 talk still in my mind, one afternoon Don mentioned a name: Eric Davidson. This was in the context of work by Herman Denis which Don had emphasized in one of his course lectures; Herman’s study was an extension of work Eric had done at Rockefeller the year before (vide supra). As often happens, a certain name sticks in one’s mind when one has developed, à priori, a “prepared mind”. I rushed to the MBL Library and read Eric’s latest papers and saw that embryology was changing from a science of observation to one of causation. We have all had this experience, when we suddenly become aware of a particular name that seems to be recurring, an intrusive phenomenon and yet suffused with a growing admiration. (Case in point: while writing this article I became aware, for the first time, of the popular singer “Adele”). So I said to myself, who is this guy Davidson? A decade went by and then I met Eric at the 1972 Gordon Conference on Animal Cells and Viruses (Fig. 1) and later at a meeting of the NIH Molecular Biology study section. Eric’s term on that study section overlapped with that of Elliott Robbins, my post-doctoral mentor at Albert Einstein College of Medicine. When Elliott came back from his first meeting of the study section he walked into the lab and said “Thoru- there is a guy on the study section who knows everything. Eric Davidson.” I can still see this in my memory as if it was yesterday, and by then Eric’s name was “all in” with me. But I still had not really gotten to know this person.

Another decade went by and then, by the circumstances of several meetings we were at, our friendship took off. Very strange, as we were not in the same fields at all and yet we found great pleasure in talking. And did we talk, into the wee hours, on all things. We soon realized that we shared two things: a profound distaste for political careerists who leverage their B.S. (used here as an abbreviation for the waste product of bovine ingested food, not the undergraduate college degree) into a faux career (more on this to follow) and that we loved the history of embryology. I soon realized that I had misjudged my friend in this nascent relationship: he was not only a master of embryology, he was a master of many things. I had fallen into a friendship with the first autodidact I was to encounter. Over the ensuing years I watched with envy all the key steps in Eric’s wonderful scholarship, including his books (each sui generis), his partnership with Roy Britten which led to gene regulation being written down as principles of nucleic acid biophysics in an epochal transformation of embryology, and his cloning of the sea urchin actin gene which was the beginning of his major mid-/late career achievement, viz. the formal portrayal of gene regulatory networks based on empirical measurements (Pederson, 2006; Tu et al., 2013). He created a seminal Gene Regulatory Network course at MBL and powerfully drove it even in recent years when his somatic health was failing. Like all his friends and colleagues, I was amazed at his zeal to keep teaching this course as well as running his productive lab at Caltech while his physical condition was deteriorating. As a profound sign of his character and strength, in all our times together in the last years of his life, I never heard him complain. All I heard and saw was his quintessential, lifelong intrepid passion to advance the frontiers of

development. At the time of his death, his latest book with Isabelle Peter had just come out and was being applauded (Mann, 2015). From our years together I have the following composite of memories and perspectives. Eric could be tough but he was never mean. When people misjudged his motive, they didn’t know the person and didn’t know that his zeal for truth sometimes got in the way of his manners. This is one area in which we were, though such close friends, totally opposite. My upbringing had taught me never to raise a voice or, worse, cause a ruckus. His childhood had been in a richly intellectual home and there were no holds barred. Over the years I realized, from Eric, that my childhood had been deficient in this respect. As our friendship evolved and we increasingly shared more and more things, I began to see my friend in broader dimensions. His zeal for beholding development as the true wonder it is was probably in part something inherited from his father, a gifted artist. A strand of DNA, or at least some meiotically recombined genes, came to the son, who drew beautifully and was a passionate devotee of form, wherever encountered. (When I once asked him if he had pondered a career in art he replied “If your father is Joe Namath, you don’t go out for football”. Ironically, Eric loved football and donned full gear for an informal team of Caltech acquaintances and was known for his voracious play). I came to know two deep traits of Eric as our friendship got more intimate in the 1990s and beyond. One was his love of American folk music (not the 1960’s Bob Dylan variety but its precursors). When I learned about the recordings of the last of those musicians he had made on many treks into the mountains of Virginia starting in the

1960’s, I asked him how this had come to be, him a New York kid for whom this form of American music would have not been expected to be of interest. He said it was because this music was about people, pure and uncontaminated. Eric was along with Allan Lomax a pioneer of recording this music. Here we need to applaud Jane Rigg, Eric’s longtime lab manager and scholar of embryology literature, who undertook the production of mastered versions of the initial tapes Eric and she had recorded in their treks. And Eric was no amateur historian of American mountain music. He played the banjo brilliantly himself (Fig. 2). In his beautifully evocative piece in this issue Caleb Finch has described Eric’s forays into early American folk music in greater detail. I have long pondered this intense love Eric had of early American mountain music and think it may have been a key to his psyche. I suspect it was a meeting of his inherited artistic talent and his socialist upbringing. More than once he conveyed to me the warmth and passion he felt about the people he met in his odyssey and the one and only time I ever saw my friend reveal profound emotion, with tears, was when he described one particular man, Wade Ward (known to Eric and others as “Uncle Wade”) who had become a cherished muse to him in those back hills of southwest Virgina. He had two professional colleagues to whom he was as close as any scientist can be, Mirsky and Roy Britten, but from his avocational life he shed tears for a simple man who knew some music and lore and who had kindly befriended him. Therein is a deep clue to Eric’s soul and I was privileged to witness it, up at his house that time. My friend was a socialist liberal and in the time of Joseph McCarthy might have been branded with that toxic label “fellow traveler”. He had a deep reverence for anyone or any people who had been mistreated.

In the 1970’s he was a leader in demolishing a fraudulent ChineseAmerican embryologist that the Rockefeller Foundation had corruptly backed in Beijing, his zeal being both for the scientific misconduct as well as the damage to the next generation of Chinese developmental biologists. He wrote a devastating (dissenting) critique in a National Academy report on Chinese science. This was one of Eric’s finest, and yet not widely known moments. My memories of my friend will always be the keenness of his intellectual penetrance of any subject we happened to discuss, the amazing breadth and recall of his mind and his enduring outlook on life where he always wanted social justice to prevail. I will also always remember his knife-like sense of humor, finding comedy not in the trivial but in the true human comedy of the absurd. Most of all, I will always cherish that he found something in me that he enjoyed. Again: “unlikely”. Eric was born into a Jewish home but was not religious. But, nonetheless, I want to close with a parable that seems fitting. As the story goes, a man left two letters to his survivors. The first letter said: “Go ask the rabbi if I can be buried with my socks on.” So they went and asked the rabbi, and he said “No.” The first letter had also said: “After you see the rabbi, open the second letter.” They then did and it said “You see, I cannot even leave with my socks on.” This parable gave rise to a cherished principle which translates from the Hebrew as “the crown of a good name”. Eric Davidson, agnostic, and my dear friend, leaves us wearing that crown. Acknowledgments: I thank Jane Rigg and Tuck Finch for information Eric’s and their treks into the heartland of American mountain music.

Their recordings are in the Smithsonian Folkways Recordings archive at the Smithsonian Institution, Washington, DC. References Davidson, E.H., Allfrey, V.G. and Mirsky, A.E. (1964) On the RNA synthesized during the lampbrush chromosome phase of amphibian oogenesis. Proc. Natl. Acad. Sci. USA 52: 501-508. Mann, R.S. (2015) How genomes generate animals: a view from Pasadena. Review of “Genomic Control Processes: Development and Evolution” by I.S. Peter and E.H. Davidson. FASEB J. 29: 4764-4765. Mirsky, A.E. (1964) In: “Differentiation and Development. Proceedings of a Symposium Sponsored by the New York Heart Association”. Little, Brown and Co., Boston. pp. 45-48. Pederson.T. (2006) The sea urchin’s siren. Dev. Biol. 300: 1-9. Tu, S., Pederson, T. and Weng, Z. (2013) Networking development by Boolean logic. Nucleus 4: 85-91. Figures Fig. 1. A portion of the group photo taken at the 1972 Gordon Research Conference on Animal Cells and Viruses, Tilton School, NH). Our subject is first on the left in the front row. Photo and permission provided by the Gordon Research Conferences. Fig. 2. Time out for another passion while at MBL, 1996. Photo taken by Peggy Bierer and kindly provided by Jane Rigg. Fig. 3. At the 2009 Cold Spring Harbor Annual Symposium “Evolution: The Molecular Landscape”. At this CSHL talk Eric was at the zenith of his

career-epochal command of development and evolution and he remained so all thereafter until his untimely death. While some speakers feel fatigue or non-connectivity with the audience after a talk, this photo reveals him ready and keen for more engagement. Said differently, it is “so him”. Photo reproduced with permission of the Cold Spring Harbor Laboratory.