It is no exaggeration to say that What is Life? is one of the most impor- tant scientific books of all time. —Graeme K. Hunter1p252 On professional biologists the book had probably little or no influence. In so far as they bothered at all to read “What is Life?,” they probably consid- ered the title a piece of colossal nerve. At their most charitable, they must have viewed the book with amused tolerance. —Gunther S. Stent2p3 The Austrian scientist Professor Erwin Schrödinger. He shared the 1933 Nobel Prize for his research into atomic theory. 18 /Autumn 2015 © Hulton-DeutschThe Collection/CORBISPharos The remarkable life of Erwin Schrödinger’s What Is Life? Curtis E. Margo, MD, MPH, and Lynn E. Harman, MD Dr. Margo (AΩA, Emory University, 1974) is Clinical The audience was diverse, ranging from students to dignitaries, Professor of Ophthalmology, and Pathology and Cell including Eamon de Valera, the Irish prime minister. Because Biology at the Morsani College of Medicine, the University so many people had to be turned away, Schrödinger repeated of South Florida. Dr. Harman (AΩA, University of the lecture the following Monday to another packed hall. The South Florida, 1993) is Clinical Assistant Professor of lectures continued weekly into March, with little falloff in at- Ophthalmology at the Morsani College of Medicine, the tendance. Interest in the talks was understandable, given the University of South Florida. novelty of hearing a renowned physicist (who just fifteen years earlier had helped to disassemble the common-sense universe n the evening of February 5, 1943, famed theoretical of Newton) discuss a subject so removed from the physical physicist Erwin Schrödinger delivered the first in a sciences. Even in America, as the nation was transfixed on series of lectures titled “What Is Life?” In the weeks the aerial bombardment of northern Germany and fighting in Oprior to the event, news spread that it would be open to the Guadalcanal, the lectures made news. But coverage in the press public. The lecture was held at Trinity College, Dublin, where was trifling. TIME magazine devoted more space to describing Schrödinger was head of the Institute of Advanced Studies. Schrödinger’s love of the Gaelic language and Irish music than The Pharos/Autumn 2015 19 The remarkable life of Erwin Schrödinger’s What Is Life? to the content of the presentation.3 Schrödinger, however, missed Austria. In 1936, he ac- Following the lecture series, Schrödinger prepared the cepted a professorship at the Karl Franzen University of Graz, material for publication, adding an epilogue on determinism along with an honorary position at the University of Vienna. and free will that had not been presented originally. The book Ostensibly unconcerned about the threat of Nazism, he found What Is Life? was released in December 1944. At $1.85 a copy, himself in considerable danger when Adolf Hitler annexed it sold briskly despite wartime austerity. As the upheaval of Austria in 1938. Schrödinger was viewed by the government as World War II ended, the book remained surprisingly popular. untrustworthy, and ordered by the Nazi rector of the university A second edition without substantive editorial revision was to write a public apology (or confession) to the Führer. The published in 1948. It was eventually translated into seven lan- letter was published in German and Austrian papers, leaving guages and sold over 100,000 copies.4p403 Schrödinger humiliated and his British colleagues bewildered. As the sophistication of biochemical and genetic research Soon thereafter, he was summarily dismissed from the profes- soared in the 1950s and 1960s, opinions concerning What Is sorship at his alma mater. With the help of friends in England Life? among elite scientists became oddly polarized. Some and Ireland, he managed to escape Austria with his family claimed the book was an influential document that ushered before war descended on the continent. in the new epoch of molecular biology, while others viewed Schrödinger was unable to return to Oxford as an émigré of it as factually misleading and unoriginal. Biochemist Graeme a combatant nation once the war began, but he obtained a posi- Hunter proclaimed it “one of the most important scientific tion as head of the newly created Dublin Institute of Advanced books of all time,” 1p252 while bacteriophage biologist Gunther Studies. He remained in Ireland for seventeen years. Stent thought any admiration as unwarranted.2 Few twentieth- century science books have received more divergent appraisals. The book Though no single explanation may adequately address this What Is Life? is a concise manuscript of roughly 27,000 anomaly, exploring the contradictory verdicts may shed light words, slightly shorter than George Orwell’s Animal Farm.5 on the factors that have shaped modern biology. It conveys an excitement about deciphering genetics that conventional textbooks never approach. Schrödinger may Erwin Schrödinger not have expected the intended audience to understand the Erwin Schrödinger (1887–1961) was born and raised in mathematics or physics behind the arguments he proposes, Vienna, Austria. The only child of a wealthy bourgeoisie fam- but he did want readers to sense the power of human intellect ily, he enjoyed a comfortable life insulated from the social to comprehend nature. He covers subjects as different and strife that marked the waning years of the Hapsburg Empire. complex as statistical mechanics, the laws of thermodynamics, His aunt Minnie taught him English at an early age. A brilliant and Mendelian genetics through the liberal use of analogies student, he excelled in mathematics and the humanities at and nearly a dozen line drawings. The book contains scattered Akademisches Gymnasium and the University of Vienna, from footnotes, a rare reference, and no bibliography. In the first which he was awarded a Doctor of Philosophy in physics in three chapters, Schrödinger summarizes the modern physicist’s 1910. Schrödinger rose steadily through the ranks of academia approach to investigating atoms and molecules, the biological to become Professor of Theoretical Physics in Zürich in 1922. mechanism of inheritance, and recent animal experiments His greatest intellectual achievement occurred four years with X-ray induced mutations. Although these reviews are later, with the publication of six papers on wave mechanics in superficial and prone to oversimplification, Schrödinger fuels Annalen der Physik. These works culminated in the wave equa- expectations that some magnificent insight is always lurking tion, which describes the behavior of particles in the atomic on the next page. and subatomic world. A computational cornerstone of quan- Schrödinger accommodates the nonscientist by assuming tum mechanics, the wave equation (which now bears his name) the role of a supportive teacher. This was a particularly exciting catapulted Schrödinger to the most prestigious academic posi- time in biology. The chromosome that biologists were just be- tion in Germany: Professor of Theoretical Physics in Berlin and ginning to explore is what he calls an aperiodic crystal, “a mas- successor to Max Planck.4 terpiece of embroidery” equivalent to a “Raphael tapestry.” 5p5 After Schrödinger arrived in Berlin in 1927 he found the While praising the organic chemist for contributing knowledge political and social atmosphere increasingly disturbing. In 1933, about this tapestry, he laments the physicist for having done he abruptly left Berlin to become a Fellow of Magdalen College, next to nothing to further understanding of the molecule. He Oxford, a decision many friends believed was in protest of is both perplexed and awed by this genetic substance, undoubt- Nazi malevolence. After arriving in Oxford, Schrödinger was edly a polymer, because it reproduces itself with remarkable awarded the Nobel Prize in Physics for his wave equation. This fidelity for generations. To emphasize this key point with a international recognition not only highlighted his awkward tangible example, he selects the infamous Hapsburg lip, faith- departure from Berlin, it insured that the Nazi officials would fully preserved throughout the long dynasty. remember the move as an insult. In the second chapter, Schrödinger introduces the term 20 The Pharos/Autumn 2015 code-script to describe the key function of the chromosome. He among scientists,” yet cautioned that this “meaty exposition” soon apologizes for the inadequacy of even this term, stating was not light reading.7p283 that the material of life is more than just a blueprint. Consider More thorough reviews appeared in professional journals. it instead both an “architect’s plan and builder’s craft—in Distinguished geneticist J. B. S. Haldane, writing in Nature, re- one.” 5p22 His translation of science to the vernacular is artful spectfully disagreed with many of Schrödinger’s “details” and and absolutely necessary to offset the rapid pace at which he “fundamental principles,” but admitted that the work was fas- introduces complicated ideas. Technical words and theorems cinating to read.8 Herman Muller, perhaps the most eminent are seemingly plucked from nowhere. Schrödinger eases the geneticist of the World War II era (and who performed much tension of scientific jargon with the implicit assurance that of the original work on radiation-induced genetic mutations), everything will eventually make sense. complimented Schrödinger for emphasizing the need for From the third to sixth chapters, Schrödinger discusses strengthening the liaison between the physical and biological the quantum nature of genetic mutations and the application sciences. But he, too, warned readers not to take the genetics of the second law of thermodynamics to living matter. Gaps in and chemistry too seriously, because they were incomplete development of concepts are inevitable without mathematics and flawed.9 or sufficient space. The crux of his thesis is found in the fourth After the molecular revolution that followed the discovery chapter: “In the light of present knowledge, the mechanism of of DNA structure in 1953, What Is Life? began to be viewed in heredity is closely related to, nay, founded on, the very basis a different light.