HARVEY, HILDEBRAND WOLFE

(b. London, United Kingdom, 31 December 1887; d. Plymouth, United Kingdom, 26 November 1970),

biological oceanography, marine analytical chemistry, marine ecosystems and production.

Harvey was one of the guiding spirits of a group of marine scientists at the Plymouth Laboratory of the Marine Biological Association of the United Kingdom who, between the late 1920s and the mid-1950s, established the ways in which marine production is governed by the interplay of the physics of the sea, its chemical composition, and the interactions of the organisms present. Their work involved the development of new techniques for measurement of the nutrient salts necessary for phytoplankton growth; the analysis of how temperature, salinity, light, and organisms were related; and—the crowning achievement of the Plymouth Laboratory during the 1930s—a deductive analysis of the control of marine production by the interplay of environmental factors, plant growth, and grazing by zooplankton. In this, Harvey was the main theoretician as well as a major contributor of new techniques. His books, published in 1928, 1945, and 1955, summarized the state of research in marine plankton production and provided a model of how research in that field could be carried out by impeccable attention to the biology, chemistry, and physics of the ocean.

Background and Training . His parents were Laetitia (née Wolfe), and Henry Allington Harvey, a partner in a manufacturer of high-quality paint. As an undergraduate at the University of Cambridge (1906–1911), Hildebrand Wolfe Harvey studied chemistry. He then went on to postgraduate research under William Hardy in the Cambridge School of Agriculture, working on a series of practical chemical problems, before returning to his family’s paint firm in Surrey until the early years of World War I. During the war he served as a navigating officer on a minesweeper in the Barents Sea, developing a love of the ocean that lasted his whole life. After a brief period in naval research after the war, he was hired in October 1921 by the Plymouth Laboratory to carry out surveys of water properties of the English Channel, part of the laboratory’s contribution to the scientific program of the International Council for the Exploration of the Sea (ICES), and to do the compilation of temperature observations and analyses of salinity and nutrient salts. At Plymouth, Harvey found himself in the research environment created by the laboratory’s director from 1894 to 1936, Edgar Johnson Allen, who had begun to direct the work at Plymouth toward factors controlling marine production and thus underlying coastal fisheries. Equally influential initially was his senior colleague William Ringrose Gelston Atkins, trained in biology, physiology, and chemistry, and appointed to the laboratory only a few months before Harvey, and whose work soon turned toward the analysis of dissolved nutrients essential for plant growth in the sea and to the effects of light and temperature.

The Study of Marine Production . The context into which the work of Harvey and the Plymouth group fitted by the 1930s began in Germany three decades earlier, first in the hands of the physiologist Victor Hensen and then of his younger colleague Karl Brandt, whose research group in the University of Kiel first showed that the production of marine plankton organisms had a seasonal cycle, which could be attributed to changes of light and chemical nutrients, especially phosphorus and nitrogen. By late in the 1920s, the Kiel group had established a theory of plankton production that applied well to seas in temperate latitudes based in large part on applying knowledge of the nitrogen cycle from land to the sea. The group had also developed new analytical techniques for dissolved plant nutrients, and had begun to tackle the problems of how the spring bloom began and why the tropical oceans showed low levels of production. For reasons intrinsic to the German university system, and especially after the devastation of German science by World War I, by the 1920s the German initiative had ceased, and attention to problems of marine production were picked up by E. J. Allen’s Plymouth Laboratory, in which Harvey, Atkins, and a few other young researchers began to play increasingly important roles during the late 1920s and early 1930s.

The early survey work on hydrographic and nutrient properties of the English Channel provided the background for Harvey’s first book, Biological Chemistry and Physics of Sea Water (1928), summarizing what was known of the control of marine plant and animal production, or, as he titled the last section, “Chemical and physical factors controlling the density of population.” Atkins and Harvey had found problems with the nutrient analyses developed at Kiel, and as a result both developed new, sensitive, fast, colorimetric techniques for the determination of nutrients that allowed seasonal and geographical differences to be followed with ease. By 1928, Atkins and Harvey’s new analytical methods had been incorporated into a standard body of techniques endorsed by ICES. Three years later Harvey joined with European marine chemists to establish standard techniques for analysis of the carbon dioxide system in seawater. Each came to play a role in the synthesis of factors governing marine production that Harvey undertook with his colleagues at Plymouth in the early 1930s, based on the concept that phytoplankton provided the food base for a variety of animals, and that all the organisms provided energy to decomposers; these, in their turn, returned inorganic nutrients to the system. In the early years Harvey and his colleagues examined this scheme in greater and greater detail, concentrating on factors that controlled the rate of phytoplankton production and the regeneration of nutrients.

Plankton Production and Its Control . Harvey’s insight by 1928 that the marine system was a closed one, involving producers, grazers, and predators, provided the conceptual background for the Plymouth group’s greatest contribution to biological oceanography. Some technical problems, namely the collection and enumeration of phytoplankton cells, had to be overcome first. A skilled instrument maker as well as experimenter and conceptualist, Harvey designed a small metered phytoplankton net, allowing the cells to be captured quantitatively, also a colorimetric method of estimating phytoplankton abundance, which was far faster than enumeration under a microscope. These advances provided him with the ability to quickly follow changes in abundance of plant cells as the spring bloom developed. The first uses of these techniques indicated that phytoplankton populations varied inversely with the abundance of zooplankton, suggesting that grazing by animals limited the growth of phytoplankton populations. The total growth of the phytoplankton population, estimated from the disappearance of their nutrient salts, was less than the number of cells counted, a further indication that grazing was an integral part of the marine production cycle.

With the help of his Plymouth colleagues Leslie Hugh Norman Cooper, a chemist; Marie Lebour, a specialist in phytoplankton; and Frederick Stratten Russell, a zooplankton specialist, in 1934 Harvey organized a year of collaborative research to bring together all the threads of the marine production cycle in the English Channel. The result was one of the most famous of all papers in biological oceanography: Harvey, Cooper, Lebour, and Russell’s “Plankton Production and Its Control,” published in the Journal of the Marine Biological Association of the United Kingdom in 1935. In it they demonstrated, often in deductive fashion, the control of marine phytoplankton abundance by light, vertical water motion, and nutrients, and the role of grazing zooplankton in holding phytoplankton below their potential level of abundance. This accomplishment rapidly came to be regarded as the classic formulation of the causes of the plankton cycle in the sea and the necessary outcome and extension of the work of Harvey’s continental European predecessors at Kiel and elsewhere.

Factors Controlling Growth of Phytoplankton . Many problems remained after 1935, including details of the marine nitrogen and phosphorus cycles, and the possibility that trace elements, as well as other major nutrients such as iron and manganese, could play a role in controlling production. His approach was to put his formidable experimental skills to work on the effects of the variation of light, temperature, and a number of trace substances and nonclassical plant nutrients on the growth of the phytoplankton cells. Many of these investigations occupied the last twenty years of his career at Plymouth, up to his retirement in 1958. The larger framework of these investigations was what he had referred to in 1928 as the “mosaic of conditions” controlling plankton abundance and ultimately marine production overall (p. 164). By the time his book Recent Advances in the Chemistry and Biology of Sea Water was published in 1945 (the result of enforced “idleness” during World War II, when air raids forced the laboratory to close), Harvey was beginning to think of marine systems as the products of intricate balances among nutrients, plants, and animals. He believed that marine ecosystems were homeostatic, constantly shifting around a maximum level of production that was achieved only when the transfer from one part of the marine production system to another was closely coupled.His last, equally influential book, The Chemistry and Fertility of Sea Waters (1955), took up the emerging relations of bacteria in marine ecosystems, along with new ideas on nutrient cycles, emphasizing that marine systems were constantly passing into and out of balance, involving the formation and decomposition of organic matter, and linking what went on in the overlying water and in the sediments below. It was then, in the 1950s, that Harvey began to put together a scheme of marine production that explicitly involved links between inorganic nutrients, plankton, bottom animals, and fish, a subject that had been suggested by E. J. Allen in the 1920s as one of the goals of research at the Plymouth Laboratory. Only with the accumulation of what he regarded as solid observationally and experimentally based knowledge did Harvey believe that such generalizations could be justified.

Harvey’s Legacy . His work was recognized by the award of a ScD at Cambridge in 1937, his election as a Fellow of the Royal Society of London in 1945, the award of the Alexander Agassiz Medal of the U.S. National Academy of Sciences in 1952, and by his appointment as a Commander of the Order of the British Empire in 1958. When Harvey retired in 1958, research on marine production and the nature of marine ecosystems was being carried on in many other places outside Plymouth. A great deal of new work, much of it in the United States, extended and amplified the grazing model that Harvey, Cooper, Lebour, and Russell had proposed, as well as the detailed laboratory investigations that Harvey carried out during the late 1940s and 1950s. But Plymouth and Harvey’s publications remained the gold standard for work on marine production at least into the 1970s, and the elements of Harvey’s contributions are not difficult to detect, although seldom acknowledged and often not recognized, in a great deal of biological oceanographic research in the twenty-first century.

BIBLIOGRAPHY

WORKS BY HARVEY

Biological Chemistry and Physics of Sea Water. Cambridge, U.K.: Cambridge University Press, 1928.

With Leslie Hugh Norman Cooper, Marie Lebour, and Frederick Stratten Russell. “Plankton Production and Its Control.” Journal of the Marine Biological Association of the United Kingdom 20 (1935): 407–441. One of the great classics of marine science.

“Production of Life in the Sea.” Biological Reviews of the Cambridge Philosophical Society 17 (1942): 221–246.

Recent Advances in the Chemistry and Biology of Sea Water. Cambridge, U.K.: Cambridge University Press, 1945.

“On the Production of Living Matter in the Sea off Plymouth.” Journal of the Marine Biological Association of the United Kingdom 29 (1950): 97–137.

The Chemistry and Fertility of Sea Waters. Cambridge, U.K.: Cambridge University Press, 1955.

OTHER SOURCES

Cooper, Leslie Hugh Norman. “Hildebrand Wolfe Harvey 1887–1970.” Biographical Memoirs of the Fellows of the Royal Society 18 (1972): 331–347. Includes a partial bibliography and significant personal information.

———. “Obituary: Hildebrand Wolfe Harvey.” Journal of the Marine Biological Association of the United Kingdom 52 (1972): 773–775.

Mills, Eric L. Biological Oceanography: An Early History,1870–1960. Ithaca, NY: Cornell University Press, 1989. See especially chapters 8 and 9 for a detailed account of Harvey’s career and work.

Eric L. Mills