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This article has This been published in or collective redistirbution of any portion of this article by photocopy machine, reposting, or other means is permitted only with the approval of The approval portionthe ofwith any permitted articleonly photocopy by is of machine, reposting, this means or collective or other redistirbution A SEA OF MICRO B ES > SECTION I. PROLOGUE > CHAPTER 1. MICROBIAL OCEANOGRAPHY: ROOTS And BR ANCHES Oceanography Foundations of Microbial Oceanography journal of The 20, Number 2, a quarterly , Volume BY DAVID M. K ARL And LITA M. PROCTOR It’s been said, “The farther backwards birth. However, van Leeuwenhoek animalcules,” neither van Leeuwenhoek you look, the farther forward you are improved the art of polishing lenses of nor his contemporaries had any under- likely to see” (Sir Winston Churchill, short focal length, and with a single lens standing of the nature of these tiny O 1874–1965). This is especially true in sci- instrument (“simple” microscope) he life forms. Van Leeuwenhoek was not ceanography ence where knowledge is cumulative and was able to achieve a magnification of a scientist, and his letters to the Royal contemporary challenges are built on the approximately 275 diameters (Porter, Society about his discoveries have been S ociety. ociety. successes of the past. Microbial oceanog- 1976), enough to observe bacteria in described as “tiresome because of repeti- C raphy is a relatively new discipline that seawater (van Leeuwenhoek, 1677). In a tion and discontinuity” (Kofoid, 1934); The 2007 by opyright endeavors to establish a comprehensive recent essay on the discovery of micro- he was even called an “immortal dilet- understanding of sea microbes, from organisms, Gest (2004) suggested that tant” (Becking, 1924). It would take genomes to biomes, thereby coupling the credit for the discovery of microbes more than 150 years until the Prussian O ceanography ceanography biosystems to ecosystems. Microbial should be shared by Robert Hooke microscopist Christian Gottfried O ceanography ceanography oceanography has its historic roots in (1635–1703) and van Leeuwenhoek. Ehrenberg (1795–1876) and others con- S the more established fields of microbiol- Hooke had published a monograph, ducted detailed, laboratory-based inves- ociety. S ogy, oceanography, and marine biology. Micrographia, in 1665 showing draw- tigations of microorganisms, includ- ociety. A ll rights reserved. Permission is granted to copy this article for use in teaching and research. article for use and research. this copy in teaching to granted ll rights reserved. is Permission Many current practitioners, however, are ings of microbes that he had seen with ing culture studies of isolated marine S end all correspondence to: [email protected] or Th e [email protected] to: correspondence all end unaware of some of the key benchmarks his compound microscopes, and includ- microbes. In 1828, Ehrenberg coined and the contributors who helped to ing a section that described—in some the term “bacterium” (from the Greek, establish their discipline. Our introduc- detail—how to construct a microscope meaning “little stick”), and in so doing tion presents a few selected waypoints in from a “very clear piece of Venice glass.” separated them from other microbes. microbial oceanography. The debate over who should get credit for the discovery of microbes is impor- DAVID M. KARL ([email protected]) is DISCOVERY OF MARINE tant, but will not be settled here. Rather Professor, Department of Oceanography, than create a revisionist view, we will MICROORGANISMS University of Hawaii, Honolulu, HI, USA. O ceanography ceanography The earliest study of marine micro- conclude that van Leeuwenhoek prob- LITA M. PROCTOR is Associate Research organisms can be traced to a Dutch ably observed, and certainly published, Scientist, University of California, Santa S draper and amateur lensmaker, Antony the first description of a “sea microbe.” Cruz, CA, USA, and currently Program P ociety, van Leeuwenhoek (1632–1723). Others The authoritative, indeed majestic, Director, Molecular and Cellular Biosciences, O before him, dating back to Leonardo biography of van Leeuwenhoek by Directorate of Biosciences, National Science Box 1931, da Vinci in the fifteenth century, had C. Dobell (1932), published on the Foundation, Arlington, VA, USA. Opinions, R ockville, MD 20849-1931, R employed magnifying lenses for the 300th anniversary of van Leeuwenhoek’s findings, and conclusions or recommenda- reproduction, systemmatic epublication, study of small objects, and the com- birth, should be required reading for tions expressed in this paper are those of the pound microscope had already been all microbiologists. authors and do not necessarily reflect the invented before van Leeuwenhoek’s Despite his ability to observe “little views of the NSF. USA . 16 Oceanography Vol. 20, No. 2 Ehrenberg was also one of the first sci- Golden Age of Microbiology,” was the (1856–1953) and the Dutch microbiolo- entists to systematically describe fossil use of pure cultures. This was often gist Martinus Beijerinck (1851–1931) led microbes (diatoms, radiolaria, foramin- considered a requirement, for example, to the development of the enrichment ifera, coccolithophorids) in rocks. His in the application of Koch’s postulates1 culture technique. This single advance influential monograph, Mikrogeologie, (Grimes, 2006). in technology facilitated the discovery published in 1854, marked the beginning In his writings, Pasteur discussed the of chemolithoautotrophic metabolism of micropaleontology as a scientific dis- importance of the environment in con- (ammonium, sulfur, and iron oxidation cipline and was an important precursor trolling physiology and metabolism, and linked to carbon dioxide fixation), nitro- for modern studies in geobiology. he expressed great interest in the nascent gen fixation, and sulfate reduction, lead- During the nineteenth century, there were several other noteworthy advances in our understanding of the microbial world that helped to promote investi- gations of marine microbes. First was Microbial oceanography is a relatively the notion that bacteria were actually new discipline that endeavors to establish a diverse assemblage of related organ- isms, which lead to various proposals a comprehensive understanding of sea for classification. The Prussian biolo- microbes, from genomes to biomes, gist Ferdinand Julius Cohn (1828–1898) thereby coupling biosystems to ecosystems. identified several unique groups of bacteria based on morphology, cellular structures, pigmentation, and growth characteristics; indeed, he has been credited with founding modern micro- field of microbial ecology, but time did ing to a greater understanding of the role biology (Drews, 1999). Cohn was one not permit his full attention to these of microorganisms, especially bacteria, in of the first to conduct detailed labora- matters. At the diamond jubilee meeting global biogeochemical cycles. Beijerinck tory studies of bacterial growth using of the American Society of Microbiology also was one of the first to recognize defined media, and his work helped to in 1974, René Dubos coined this dichot- “soluble microbes” and coined the term debunk the concept of “spontaneous omy “Pasteur’s dilemma —The road virus to describe them. He also founded generation” of life by documenting that not taken,” and rhetorically asked where the Delft School of Microbiology, which vegetative cells, but not endospores, were the field of microbial ecology (includ- became an important center for research killed by boiling water (Drews, 1999). ing marine microbial ecology) would and training of the next generation of His achievements became an important be today if Pasteur had selected it as his microbial ecologists. foundation for modern microbiology, primary research thrust. Instead, the dis- along with the more well-known stud- cipline lay relatively dormant (at least in ACCESS TO THE SEA ies by Louis Pasteur (1822–1895) and comparison to medical/industrial micro- The other major root of microbial Robert Koch (1843–1910) on infec- biology) for nearly 50 years. Toward the oceanography, namely developments in tious disease and pathology, vaccina- end of the nineteenth century, inde- oceanography and marine biology, can tion, and fermentation. A hallmark of pendent contributions by the Russian be traced back to Kiel and Plymouth this era, sometimes called the “First microbiologist Sergei Winogradsky with the comprehensive studies of 1 Koch’s postulates are four criteria that need to be satisfied to prove that a microorganism causes a disease: the microorganism must found in every case of the disease, the micro- organism must be isolated from the diseased animal and grown in pure culture, the same disease must develop when introduced into healthy individuals, and the microorganism must be re-isolated from the new infection. Oceanography June 2007 17 marine plankton by mostly European of Microbiology was well underway by Sevastopol Biological Station of the scientists, including Victor Hansen, Karl 1872, there were no systematic attempts Academy of Sciences of the USSR, also Brandt, Ernst Haeckel, Louis Agassiz, to collect or study marine bacteria on the founded in 1872, was dedicated pri- and Anton Dohrn, among others. In the HMS Challenger expedition. However, marily to interdisciplinary studies of United States, the establishment of the by the end of the nineteenth century, the Black Sea. The Kristineberg Marine Navy Hydrographic Office following the

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