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The Discovery of Revisited A close reading of 17th-century documents shows that Hooke, rather than , was the first to observe a

Howard Gest

wo remarkable geniuses, Robert Preface of and in a 1678 treatise, Hooke and Antonie Leeuwen- Lectures and Collections; Microscopium. hoek, deserve the credit for discover- Hooke’s Preface to Micrographia also describes T ing microorganisms in the 17th cen- how to make a single-lens of the tury. A complex of events and same kind that Leeuwenhoek used in studies a common interest in were instru- that began almost a decade later. mental in leading these two men from very dif- According to my analysis along with the im- ferent backgrounds to codiscover the microbial portant studies of microscopist Brian Ford, . Their separate journeys into that Hooke rather than Leeuwenhoek was the first to were recorded between 1665 and 1678 in pub- observe and document the existence of a micro- lications of the Royal Society of London. . Moreover, his Micrographia pro- Among Hooke’s outstanding achievements is vided the basic model and “trigger” for Leeu- the first published description of a microorgan- wenhoek’s subsequent discoveries of other ism. From Hooke’s excellent drawing in Micro- microbes during the the . Thus, graphia (1665), mycologists identify Hooke’s as as Antonie van Leeuwen- specimen as the microfungus , the com- hoek should be considered responsible for “fa- mon (Fig. 1). In Micrographia, thering” modern . Hooke illustrated microscopic views of diverse biological objects, including , wood, , surfaces, hair, pea- A Glimpse of Robert Hooke’s cock feathers, fly wings, of Illustrious Career silkworms, mites, a flea, and a Robert Hooke (1635–1703) was —as well as that of a mold. Robert Hooke as well as enrolled as an undergraduate at Hooke obtained the microfungus Christ Church College at the Uni- specimen from a “small white spot Antonie van versity of Oxford but apparently of a hairy mould,” many of which Leeuwenhoek never obtained a degree. Nonethe- he observed on the red sheepskin should be less, he became associated with a covers of a small book. He called the considered brilliant group of scholars, includ- organism a “microscopical mush- responsible for ing Christopher Wren and Robert Howard Gest is room.” Although he attempted to “fathering” Boyle, who met regularly to dis- Distinguished Pro- observe the “” (now called spo- cuss a broad range of scientific modern fessor Emeritus of rangiospores) from which it devel- problems. Microbiology and microbiology ops, the relatively low magnifying In due course, this group be- Adjunct Professor power of his microscope made that came the nucleus of the Royal So- of History and Phi- effort impossible. ciety. Hooke, a founding member losophy of Science Hooke advanced the techniques of micros- of the Society, served as “Curator of Experi- at Indiana Univer- copy in many ways that are detailed in the ments” from 1662–1677. His duties included sity, Bloomington.

Volume 70, Number 6, 2004 / ASM News Y 269 FIGURE 1 tenary of Hooke’s , several new biographies were published, describing his and scientific achievements as well as his impor- tant architectural contributions while London was reconstructed following the Great Fire of 1666.

Antonie van Leeuwenhoek’s Unusual Career Leeuwenhoek (1632–1723), who had little formal schooling, opened a shop at age 22 as a draper in , Holland. Indeed, his scientific career likely traces to his use of low-power magnifying glasses to inspect cloth. Leeuwenhoek developed the ability to make superb micro- scopes containing single lenses Microscopic view of a “hairy mould” described by Robert Hooke in 1665 (in Micro- that were about 1 mm in diame- graphia). This image was the first published depiction of a microorganism. The reproductive (sporangia) are characteristic of the microfungus Mucor. Sporangia in different stages ter. Leeuwenhoek was a keen ob- are identified by the letters A, B, C, and D. Hooke included a reference; the length of the server and had extraordinary cu- bar under the diagram represents 1/32 of an inch. Image courtesy of The Lilly , Indiana riosity, which he soon focused on University, Bloomington. the natural world. For instance, he was the first to observe and describe cells of , conducting “considerable ” and do- red cells, , and cells. Al- ing other research projects officially recom- though it was then thought that maggots, fleas, mended to him. Hooke soon became a com- and the like formed through “spontaneous gen- manding intellectual force in the Society and, as eration,” he showed that such creatures hatch Curator, provided the main substance of many from eggs. meetings. During April 1663, Hooke lectured Leeuwenhoek communicated his observa- on the of tissues, such as tions to the scientific world in an unorthodox and , and coined the term “.” In Micro- way. Dutch acquaintances, who were corre- graphia, the structure of cork is detailed in Obs. sponding members of the Royal Society, sug- XVIII: “Texture of cork, and of the cells and gested that he describe his findings in letters pores of some other such frothy bodies.” addressed to the Society. During the rest of his Hooke’s scientific interests were broad, and life, Leeuwenhoek communicated some 200 let- his genius many sided. Physicist J. D. Bernal ters in Dutch, many of great length. Between wrote of Hooke in 1965 that his “interests 1939 and 1983 the “Leeuwenhoek Letters” ranged over the whole of mechanics, physics, were translated into English by large committees chemistry, and biology. He studied elasticity of Dutch , yielding 11 volumes with and discovered what is known as Hooke’s English and Dutch texts on opposing pages. Law. . . .he invented the balance wheel, the use These volumes include extensive annotation. of which made possible accurate watches and Several footnotes show that Leeuwenhoek was chronometers; he wrote Micrographia, the first familiar with Micrographia even before he be- systematic account of the microscopic world, gan sending letters to the Royal Society. including the discovery of cells; he introduced One of Leeuwenhoek’s greatest contributions the telescope into astronomic measurement and to biology was the discovery of in 1676. invented the micrometer.” In 2003, the tercen- As might be expected, his observations and de-

270 Y ASM News / Volume 70, Number 6, 2004 Valuing Impact of History on Science, Witnessing Science’s Impact on History Howard Gest feels very strongly ily to America. He received a B.A. cessful, we felt about the impact of history on sci- in from the University great pride in ence. “I believe that many young of California, Los Angeles (UCLA) our work; we scientists know little about the in 1942, and his Ph.D. from Wash- felt it was very early history of their own research ington University in 1949. While important for areas, and I think this limits their attending UCLA, he spent the sum- the security of understanding of how major dis- mers of 1941 and 1942 assisting the United coveries are made,” he says, adding Max Delbru¨ ck and , States,” Gest that too often they assume that who were studying bacterial vi- says, recalling contemporary biological knowl- ruses. Although Gest began gradu- the Manhattan Howard Gest edge is nearly complete. “History ate work with Delbru¨ ck at Vander- Project. “But we clearly shows that biology is far bilt University, World War II were very disappointed in how the more complicated than each gener- intervened. Later in St. Louis, he bombs were used. We’d hoped they ation thinks it is.” did research with Alfred Hershey, would be used in such a way that

Gest came to this realization using the radioactive isotope P32 to would not lead to the death of many ago, influenced by the late examine what happens to phos- innocent civilians. And we were very J. H. “Jack” Hexter, a noted histo- phorus-containing cell components perturbed that the petition never rian and professor of history at when bacterial replicate. reached President Truman.” Washington University, St. Louis, These studies culminated with their Gest has served on the faculties

Mo., where Gest knew him, and discovery of P32 “suicide” of bacte- of Case Western Reserve University Yale University, New Haven, riophage. in Cleveland, Ohio, Washington Conn. “Hexter once said: ‘History During World War II, Gest was University, and Indiana University, with a capital H deals with major involved in the Manhattan Project and has been a visiting researcher at , large movements, deep run- with physical chemist Charles the California Institute of Technol- ning , portentous rumbles. Coryell—one of Gest’s teachers at ogy, Dartmouth , When a “small h” historian fixes UCLA—first at the University of Stanford University, Oxford Uni- his attention on a fragment of the Chicago, and later at Oak Ridge, versity, Tokyo University, and past washed up on the littered Tenn. His role was to conduct basic UCLA. He was twice named a beach of the present, he is likely to research on the radioactive ele- Guggenheim Fellow and has served ask simple questions about it: How ments formed in fission. on a number of advisory commit- did it get there? What the devil is it? He looks back at that period with tees of the U.S. government. During What was it for? Where is it from? conflicted emotions. While proud his second Guggenheim fellowship, What happened to it?’,” Gest says. of his contribution toward devel- he studied problems of biochemical “Hexter’s remarks made me realize oping the atomic bomb, he was as a member of the Pre- that I had become a ‘small h’ histo- troubled by the real possibility that cambrian Group. rian of microbiology and biochem- its use would result in the needless Gest no longer engages in exper- istry.” loss of life—a fear that ultimately imental research. Instead he spends Gest, 82, is distinguished profes- became a reality. Gest was among much of his writing about the sor emeritus of microbiology and those scientists who signed a peti- history of microbiology and bio- adjunct professor of history and tion that urged President Truman chemistry. He also is a continuing philosophy of science at Indiana to consider the moral implications and familiar presence among grad- University, Bloomington. His re- of dropping the bomb, and asked uate students and postdoctoral fel- search over many years focused on that the Japanese first be offered an lows, with whom he continues to microbial and metabo- opportunity to surrender. Truman work. “I go to the university every lism, especially with photosyn- apparently never saw the document single day,” he says. thetic bacteria. (see http://www.bio.Indiana.edu Marlene Cimons Gest was born in London, and /Gest/). Marlene Cimons is a freelance writer emigrated in infancy with his fam- “When this huge project was suc- in Bethesda, Md.

Volume 70, Number 6, 2004 / ASM News Y 271 scriptions of these very minute were allow me a sight of them. . .. as for at first considered dubious. Unknown to Leeu- the microscopes I looked through, wenhoek, Hooke was appointed Secretary of the they do not magnify much, if any Royal Society in 1677 and in this capacity he thing, more than several glasses I received a letter from Leeuwenhoek, dated 5 have seen, both in England and Ire- October 1677, which described bacteria in a land: but in one particular, I must pepper- infusion. The large number of needs say, they far surpass them all, bacteria in one drop astonished Leeuwenhoek, that is in their extreme clearness, who wrote, “This exceeds belief.” Accordingly, and their representing all objects so he included testimonials of “eight credible per- extraordinary distinctly.” sons” who affirmed his observations. In Microscopium (1678), Hooke confirmed This and other contemporary comments leave Leeuwenhoek’s observations of bacteria, writ- little doubt about Leeuwenhoek’secretive char- ing “much to wonder I discovered vast multi- acter. Other parts of the historical record show tudes of these exceeding small creatures which that he sometimes could also be obscure if not Mr. Leeuwenhoeck had described. . .and some outright misleading. of these so exceeding small that millions of mil- lions might be contained in one drop of water.” Hooke’s confirmation and the authority of the Leeuwenhoek Surely Was Aware of Royal Society proved momentous for Leeuwen- Hooke’s Micrographia hoek and his reputation. In 1680, he was elected More than a decade ago, microscopist Brian to the Royal Society and later became a celeb- Ford, a Fellow of Cardiff University, Cardiff, rity, visited in Delft by notables, including Tsar Wales, United , drew attention to a . passage in the Preface to Micrographia where Hooke writes an exact “prescription” for mak- Intertwined Beginnings of ing what later was called a “Leeuwenhoek mi- and Microbiology croscope.” According to Hooke: There is no doubt that Hooke was the first to “If you take a very clear piece of describe and depict a microorganism in the sci- Venice Glass, and in a Lamp draw it entific literature, and his published descriptions out into very small hairs or threads, of microscopic techniques paved the way for then holding the ends of these improvements. In contrast, Leeuwenhoek was notoriously secretive about his lenses and his threads in the flame, till they melt techniques. In 1685, 20 years after Hooke pub- and run into a small round Globul, lished Micrographia, the Royal Society sent or drop, which will hang at the end Thomas Molyneux to visit Leeuwenhoek and of the thread; and if further you stick obtain information about his methods. Moly- several of these upon the end of a neux’s report describes several of Leeuwen- stick with a little sealing Wax, so as hoek’s low-power microscopes but also com- that the threads stand upwards, and ments: then on a Whetstone first grind off a good part of them, and afterward on “Such were the microscopes, which a smooth Metal plate, with a little I saw, and these are they that he Tripoly, rub them till they come to shews to the curious that come and be very smooth; if one of these be visit him: but besides these, he told fixt with a little soft Wax against a me that he had another sort, which small needle hole, prick’d through a no man living had looked through thin Plate of , Lead, Pewter, or setting aside himself; these he re- any other Metal, and an Object, serves for his own private observa- plac’d very near, be looked at tions wholly, and he assured me they through it, it will both magnifie and performed far beyond any that he make some Objects more distinct had shewed me yet, but would not than any of the great Microscopes.”

272 Y ASM News / Volume 70, Number 6, 2004 This paragraph aptly describes how to build a fied to the point where plain untruths are being single-lens microscope identical to what Leeu- told; at best a fascinating and convoluted story wenhoek later perfected. “The design of the has been reduced to the minimum for easy, hand-held microscope that Leeuwenhoek used uncritical consumption,” Wainwright com- throughout his researches was derived from ments. Hooke’s published account,” Ford concludes. I believe that Hooke’s major contributions Hooke noted in several publications that using have been minimized or ignored because of the this kind of microscope strained his eyes, ex- slanted perspectives fostered by two widely read plaining why he preferred larger microscopes books. The more influential is the scholarly with two lenses. 1932 book by protozoologist , In his 28 April 1673 letter to the Royal Soci- Antony van Leeuwenhoek and his “Little Ani- ety, Leeuwenhoek briefly describes a “mould” mals.” Dobell took a limited view of the scope of and a “lowse”—closely matching Hooke’s microbial life, stressing Leeuwenhoek’s observa- lengthier descriptions of similar objects in Mi- tions of protozoa and bacteria. Ford describes crographia that were published eight years ear- Dobell’s biography as “a monumental, if parti- lier. In that letter, Leeuwenhoek barely alludes san, study.” Dobell’s book, which omits any to similar observations by “others.” That word mention of Hooke’s discovery of “in all probability is a reference to Hooke’s and his contributions to microscopy, inexplica- Micrographia,” according to a footnote by bly dismisses Hooke in a footnote: Dutch scientists in the edited Leeuwenhoek Let- ters. “Dr Robert Hooke, an original Fel- A close comparison no doubt. Indeed, low of the Royal Society, was also additional examples indicate that Leeuwenhoek an original and eccentric genius and studied illustrations in Micrographia and had inventor. His contributions to sci- some of Hooke’s descriptions translated. In a ence are too well-known and nu- 1676 letter to , who preceded merous to mention; though his in- Hooke as Secretary of the Royal Society, Leeu- fluence on his contemporaries, and wenhoek wrote that he had friends who would the part he played in the early days translate French or Latin, adding “I cannot help of the Society, are only just begin- myself with the English language since the death ning to receive their due recogni- of a certain gentleman who was proficient in this language.” However, the Dutch editors of Leeu- tion....It is impossible and unnec- wenhoek’s letters note that, throughout Leeu- essary to discuss this remarkable wenhoek’s lifetime, English-speaking communi- man and his work here” [italics ties thrived in Delft and many of their members added]. were competent in Dutch. Ford also documents Leeuwenhoek’s access to English-Dutch transla- Although Dobell’s book contains 34 citations to tors. Hooke in its extensive index, they are trivial, mainly noting the dates of various letters. The index does not cite Micrographia, which is sim- Why Have Hooke’s Findings, Establishing ply listed among “other References and His Priority, Been Largely Ignored? Sources.” With this background, one may well ask why Dobell’s biography was widely read by gener- Hooke’s primary role in discovering microbial ations of , many of whom were life is barely mentioned in standard accounts. also exposed to ’s popular 1926 Milton Wainwright of Cardiff University points book Microbe Hunters and read its first chapter, out that accounts of the early history of micro- “Leeuwenhoek, first of the microbe hunters.” biology typically begin with the “first sighting” The first paperback edition appeared in 1940 of microorganisms by Leeuwenhoek. For exam- and by 1963 there was a 28th printing! This ple, the Autobiography of Science refers to Leeu- account mentions Hooke only once, simply as a wenhoek as “first of the microbe hunters.” person who confirmed Leeuwenhoek’s discov- “Unfortunately, much of what is taught about ery of bacteria. It is no wonder that over the past the history of microbiology has been oversimpli- 70 years, microbiologists and the general public

Volume 70, Number 6, 2004 / ASM News Y 273 heard little or nothing about Hooke’s seminal during the 17th century. This personal scenario contributions to microbiology. seems tailor-made for de Kruif’s style, which Leeuwenhoek’s career was obviously very un- popularized historical events replete with imag- usual and, as such, had a strong element of ined dialogues. Notwithstanding those vividly drama. He was generally seen as a person of portrayed accounts, Hooke rather than Leeu- genius who made great discoveries while iso- wenhoek was the first to observe and document lated from others pursuing the natural sciences the existence of a microorganism.

ACKNOWLEDGMENT Many of these issues were discussed during the Robert Hooke Tercentenary Commemoration Symposium, which I attended, at the University of Oxford, Oxford, United Kingdom, on 2 October 2003.

SUGGESTED READING Bennett, J, M. Cooper, M. Hunter, and L. Jardine. 2003. London’s Leonardo—the life and work of Robert Hooke. , Oxford. Bernal, J. D. 1965. Science in history. Hawthorn Books, New York. Bulloch, W. 1979. The history of bacteriology. Dover Publications, New York. (Republication of the original published by Oxford University Press in 1938.) de Kruif, P. 1926. Microbe hunters. Harcourt Brace, New York. Dobell, C. 1932. Antony van Leeuwenhoek and his “little animals.” Staples Press, London. Ford, B. J. 1981. The van Leeuwenhoek specimens. Notes Records R. Soc. 36:37–59. Ford, B. J. 1985. Single lens. Harper and Row, New York. Ford, B. J. 1991. The Leeuwenhoek legacy. Biopress and Farrand Press, Bristol and London. Hooke, R. 1665. Micrographia, or some physiological descriptions of minute bodies made by magnifying glasses with observations and inquiries thereupon. J. Martyn and J. Allestry, Printers to the Royal Society, London. Hooke, R. 1678. Lectures and collections; Microscopium. J. Martyn, Printer to the Royal Society, London. Jardine, L. 2004. The curious life of Robert Hooke: the man who measured London. Harper Collins, New York. Lechevalier, H. A., and M. Solotorovsky. 1965. Three centuries of microbiology. McGraw-Hill, New York. Leeuwenhoek, A. v. 1673–1696. Alle de Briefen van Antoni van Leeuwenhoek/The Collected Letters of Antoni van Leeuwenhoek. Eleven volumes published intermittently between 1939 and 1983; edited, illustrated and annotated by committees of Dutch scientists. Swets and Zeitlinger, Amsterdam. Schifferes, J. J. 1960. Autobiography of science, 2nd ed. Doubleday & Company, Garden City, N.Y. Wainwright, M. 2003. An alternative view of the early history of microbiology. Adv. Appl. Microbiol. 52:333–355.

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