HARVEY, EDMUND NEWTON

(b. Germantown, Pennsylvania, 25 November 1887; d. Woods Hole, Massachusetts, 21 July 1959)

Physiology.

The son of William Harvey and Althea Ann Newton, Harvey was raised by his mother and three older sisters after his father’s early death. In 1909 he received the B.S. in general science from the University of Pennsylvania and in 1911 the Ph.D. from Columbia University with a dissertation on cell permeability. In 1911 Harvey was appointed instructor in Edwin Grant Conklin’s department of biology at Princeton, where he became assistant professor in 1915, full professor in 1919, and Henry Fairfield Osborn research professor in 1933. Harvey occupied this post until he retired in 1956. On 12 March 1916 he married Ethel Nicholson Browne, who had received her Ph.D. in zoology from Columbia in 1913; they had two sons. The couple shared laboratories during much of their lives.

Harvey never lost the interest in natural history fieldwork developed in his youth, but as an undergraduate he was drawn to cell physiology and the experimental laboratory by the teachings of Ralph Lillie. Both he and Lillie, Harvey later asserted, were influenced by Jacques Loeb and his program for the physicochemical analysis of life, and early in his career Harvey looked to Loeb’s work on artificial parthenogenesis as a model of what biological experimentation should be. Alfred Goldsborough Mayor invited Harvey to be a collector at the Dry Tortugas Marine Biological Laboratory of the Carnegie Institution in the summer of 1909 and, later that season, at the Marine Biological Laboratory at Woods Hole, where Harvey investigated membrane formation and artificial parthenogenesis.

Harvey’s visit to Woods Hole initiated a lifelong association with marine biological stations and an emphasis on marine organisms in much of his research. From 1909 until the end of his life he spent part of every summer doing research at Woods Hole. It was in 1913, while on an expedition with Mayor to Australia’s Great Barrier Reef, that Harvey’s interest was directed to bioluminescence, the phenomenon that would occupy the greatest part of his research career. In that year he began publishing a series of studies on light emission in luminous bacteria, but his first important paper on the mechanism of light production in animals did not appear until 1916. In it Harvey first drew attention to the work of Raphael Dubois, who in the 1880’s had shown that the luminous organs of Pyrophorus moctilucus contained two separable substances, luciférine and luciférase, that, when mixed, produced light. Harvey proceeded to demonstrate the existence of a luciferin-luciferase system in three other organisms. With bold optimism he asserted that “the problem of bioluminescence has been solved at least in its broad aspects” and predicted that the exact chemical nature of luciferin soon would be determined.

While visiting Japan in 1916 to study a luminous squid, Harvey was introduced to the organism most central to his research from then on, the ostracod crustacean Cypridina hilgendorfii. What made this tiny luminescent organism especially intriguing is that when it is dried after collecting, its luminescence system not only remains stable but can be easily activated, even after years of storage, by aqueous solutions containing molecular oxygen. Between 1916 and 1919, working with dried material shipped to his Princeton laboratory, Harvey found that Cypridina also possesses a two-component lightemitting system and developed techniques for separating and purifying its substrate, luciferin, and enzyme, luciferase. Luciferin emits light when oxidized, he showed, and the oxidation product can be reduced to luciferin again. Cypridina proved exceedingly useful in kinetic studies of its enzymesubstrate system as well, for bioluminescence provided a visible indicator of its own reaction velocity. Harvey continued to pursue chemical analyses of animal light throughout his career, after the late 1920’s focusing particularly on how bioluminescent activity is affected by alterations in the chemical and physical environment.

By the late 1920’s Harvey’s elucidation of bioluminescence had established him as doyen of the field, yet the expectations so confidently proclaimed in 1916 remained discouragingly unfulfilled. He systematically sought the Cypridina-type luciferinluciferase system in every kind of luminescent organism he could examine but found the reaction in only a few, which frustrated his hopes of disclosing a general pattern, This, along with the seeming intractability of the precise biochemistry of luminescence, helped rechannel his energies toward other physiological interests. Harvey had never set aside his research on cell permeability and between 1930 and 1934 he actively investigated cell surface tension. He was also managing editor from 1932 to 1939, of Journal of Cellular and Comparative Physiology.

Harvey was encouraged to diversify his research interests about 1927 by his friendship with the physicist Alfred Lee Loomis, who had just opened a private laboratory at Tuxedo Park, New York, within driving distance of Princeton. At the Loomis Laboratory they conducted experiments on the biological effects of high-frequency sound waves and about 1930 constructed the centrifuge microscope, which Harvey used to study the effects of centrifugal force on cells. In 1935 they entered the new field of electroencephalography, and for the remainder of the 1930’s studied the electrical potentials of the human brain during sleep.

The outbreak of World War II redirected Harvey’s research efforts. At the Loomis Laboratory radio communications research supplanted the biophysics work of interest to Harvey. While at Princeton he was among those scientists who diverted their energies to military research. In 1942 he started working for the Committee on Medical Research of the Office of Scientific Research and Development. Of the two major programs he directed, the more important investigated bubble formation in animal blood and tissues and the pathogenesis of decompression sickness; the other, which involved firing high-velocity missiles at anesthetized animals, studied the mechanisms of wounding. For his work on decompression sickness Harvey was awarded the Armed Forces Certificate of Merit in 1948.

After the war Harvey published many reports on his military research, but his laboratory activity slackened. He was vice president of the corporation of the Marine Biological Laboratory at Woods Hole from 1942 to 1952, and helped shape its reorganization and growth after the war. Harvey continued his research on bioluminescence, for which he received the Rumford Medal of the American Academy of Arts and Sciences in 1947, and published the book Bioluminescence in 1952. Perhaps in part out of frustration at never finding the simple physicochemical solutions to the problems of bioluminescence he had once anticipated, however, he increasingly shifted his attention to broader questions of the evolution of luminescent organisms, and he wrote on the history of luminescence.

BIBLIOGRAPHY

I. Original Works. Harvey’s published works are listed in poggendroff, VI, 1038–1040, and VIIb, 1875– 1878. His books include The Nature of Animal Light (Philadelphia and London, 1920); Living Light (Princeton, 1940; repr. New York, 1965); Bioluminescence (New York, 1952); and A History of Luminescence from the Earliest Times to 1900 (Philadelphia, 1957). Among more than 250 scientific papers Harvey authored or coauthored, particularly significant are “The Mechanism of Light Production in Animals, “in Science, 44 (1916), 208–209; “Potential Rhythms of the Cerebral Cortex during Sleep,” ibid., 81 (1935), 597–598, with Alfred L.Loomis and Bubble Formation in Blood and Tissues,” in Harvey Lectures, 40 (1944–1945), 41–76.

The American Philosophical Society Library, Philadelphia, has a collection of Harvey’s papers (1923–1959, about 5, 600 items plus 19 volumes), which is described in Margaret Miller, “The Papers of Edmund Newton Harvey (1887–1959),” in Survey of Sources for the History of Biochemistry and Molecular Biology, no.6 (November 1977), 8–9.

II. Secondary Literature. The fullest biographical sketch of Harvey is Frank H. Johnson, “Edmund Newton Harvey,” in Biographical Memoirs, National Academy of Sciences, 39 (1967), 193–266, which contains extracts from an unpublished autobiographical memoir, Other obituary notices that supplement this are by Elmer GrimShaw Butler, in American Philosophical Society, Year Book 1959 (1960), 127–130; and by Aurin M. Chase, in Biological Bulletin, 119 (1960), 9–10.

John Harley Warner