John Tyler Bonne
1920-2019
Richard H. Kessin
Professor Emeritus of Pathology and Cell Biology
Columbia University
John Tyler Bonner died on February 7. He was a biologist, a teacher, and a writer who could tell a fine story. John was fascinated by evolution and development, and particularly by his beloved social amoebae. He stood on a salient of biology that connected with the classical biologists of the 19th century, where the object was to examine the unusual, experiment with it, and then to explain it in an engaging way. His experiments were decidedly low tech and did not require frenetic grant writing; it was a path that left time for teaching and thinking and books, which he wrote during summers in Margaree Harbour, Nova Scotia. From the perspective of the world of constant grant writing to which I belonged, I saw him as less reductionist than the molecular biologists like me, but with a curiosity and serenity that made me want to step into his world.
John was an undergraduate at Harvard, graduating with honors in 1941. He began a PhD with Thomas Weston, working of the social amoeba (or slime mold) Dictyostelium discoideum. He received a Master’s degree in 1942, but the Second World War intervened and he enlisted in the Army Air Force, where he spent the better part of four years working in a laboratory, rising from private to lieutenant.
Leaving the Air Force in 1946, he returned to Harvard for a year and finished his PhD. Decades ago, I found it in Harvard’s Biolabs. Typed on onionskin paper, it was a systematic examination of all of the mechanisms by which starving amoebae might aggregate in waves. Eventually he worked his way to chemotaxis. We know a lot now about how aggregation in Dictyostelium discoideum and related species occurs. Some of us have made careers studying its biochemical components, but John’s interrupted thesis was the beginning.
John made a film of the aggregation of the Dictyostelium amoebae and their subsequent development into a migrating slug-like creature and then a fruiting body, with approximately 80,000 spores mounted on a stalk of dead cells. It is a film of extraordinary quality, as if the viewer had shrunk to the size of an aggregate and stood on the agar surface as the aggregates collected and developed and the slugs migrated by. The video is on the Princeton University homepage where hundreds of thousands of people have seen it. It attracted me to the field when I saw it in an undergraduate biology class.
The film had an important early viewer. After John became an Assistant Professor and moved to Princeton in 1947, he got a call from Albert Einstein’s secretary, who asked him to bring his projector and film to the Einstein house. John recounted this story at a Dictyostelium meeting in Bar Harbor, Maine in 1999. A bed sheet was hung and John, then 27, showed his film. There was animated discussion by Einstein and his guests. Physicists have always been excited by the creation of order out of a uniform field of cells and Dictyostelium aggregation had waves, which made it irresistible. When asked what Einstein and his friends said, John grinned and told his audience that most of discussion was in German, so he did not know.
In the 1950s and 1960s, the properties of the chemotactic molecule were described, if not yet its chemical identity. It was small molecule that attracted starving amoebae and was originally named acrasin, after a witch named Acrasia, who, according to Edmund Spenser in the Fairie Queene, lured sailors (Odysseus was one) to her island. Brian Shaffer, on sabbatical in John’s lab, established the details of attraction and learned that acrasin was constantly degraded by an acrasinase. In 1968, the Bonner laboratory tried cyclic AMP, which turned out be the acrasin for D. discoideum. The acrasinase was thus cAMP phosphodiesterase. An induced inhibitor glycoprotein appears to right the situation when too much phophodiesterase is made. There are several cAMP receptors and associated regulatory molecules that control chemotaxis and development.
These possibilities attracted biochemists, geneticists and cell biologists—people with a more reductionist approach to biology than John. He was delighted and labored on in his own way, producing a series of papers on various properties of the cellular slime molds, almost all of which he reported in single author publications, (about 160) and books (20).
He and Kenneth Raper did the initial experiments that turned the cellular slime molds into the experimental organisms that we know. They are now used to study problems in cell biology including chemotaxis, motility, phagocytosis, autophagy and mechanisms of bacterial pathogenesis, and self-recognition.
The amoebae are used to study many problems of ecology and evolution, among them: What preys on amoebae in the soil? How are cheaters avoided in the competition to become live spore rather than dead stalk? How are slime molds dispersed around the world? How important are they in the turnover of soil bacteria? How are the many species related?
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John Bonner was a serious teacher. Dr. Robert Klitzman, Professor of Clinical Psychiatry at Columbia and Director of Columbia’s Masters program in Bioethics, has this memory:
He was an inspiring teacher, and his course remains one of the favorites in my entire education. From him, I first learned of books by D’Arcy Thompson (On Growth and Form) and Richard Dawkins (The Selfish Gene, which had just been published the year before). I remember fondly him taking the class to the woods around the Institute for Advanced Study in Princeton—one of the few remaining areas of virgin woods in the Northeast US, to see firsthand how biological processes played out over time. He described how—in the very classroom in which he now spoke to us—the Nobel Laureate Karl von Frisch presented his research on the dance language of bees, with Einstein sitting attentively in the front row. Dr. Bonner pointed to the chair in which Einstein had sat. Bonner made the facts, excitement and history of biology come alive for us.
His service to students continued through his books. He was a charming writer and these lucid books cover fundamental issues for budding scientists including biological concerns about evolution and morphogenesis. Some of the books have been translated into multiple languages, fourteen different ones by my count. His memoir of summers on Cape Breton was equally charming.
I am grateful to Denise Valenti of the Princeton University Communications Office, for this partial list of John’s books : “Morphogenesis: An Essay on Development” (1952), “Cells and Societies” (1955), “The Ideas of Biology” (1962), “Size and Cycle” (1965), “The Cellular Slime Molds” (1967), “Life Cycles: Reflections of an Evolutionary Biologist” (1993), “Why Size Matters: From Bacteria to Blue Whales” (2006), “The Social Amoebae: The Biology of Cellular Slime Molds” (2009), and “Randomness in Evolution” (2013).
John Bonner’s service to Princeton University, its faculty and students spanned almost 65 years. As emeritus professor he was active until 2012, giving his final lecture in Introductory Biology in 2009. He was the George M. Moffett Professor of Biology, was Department Chair (several times) and was awarded numerous honorary degrees. John was a member of The National Academy of Sciences. The American Academy of Arts and Sciences, The American Philosophical Society and the Indian Academy of Sciences. Princeton awarded him an honorary degree in 2006. More information on John’s extraordinary service can be found in a tribute by Denise Valenti, which appears on the Princeton homepage.
John Bonner had four children, a daughter and three sons, and five grandchildren, who survive him. His wife Mary died before him.