Journal of Cave and Karst Studies Editor Malcolm S
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August 2006 JOURNAL OF Volume 68 Number 2 ISSN 1090-6924 A Publication of the National CAVE AND KARST Speleological Society STUDIES CAVE SURVEYOR AS MICROBIOLOGIST Journal of Cave and Karst Studies Editor Malcolm S. Field of the National Speleological Society National Center of Environmental Assessment (8623D) Volume 68 Number 2 August 2006 Office of Research and Development U.S. Environmental Protection Agency 1200 Pennsylvania Avenue NW CONTENTS Washington, DC 20460-0001 202-564-3279 Voice 202-565-0079 FAX Article [email protected] Introduction to cave microbiology: Production Editor A review for the non-specialist James A. Pisarowicz Hazel A. Barton 43 Wind Cave National Park Hot Springs, SD 57747 Article 605-673-5582 Potential effects of recurrent low oxygen conditions [email protected] on the Illinois cave amphipod BOARD OF EDITORS S.V. Panno, K.C. Hackley, W.R. Kelly, H.H. Hwang, Anthropology F.M. Wilhelm, S.J. Taylor, and B.J. Stiff 55 Patty Jo Watson Department of Anthropology Washington University Article St. Louis, MO 63130 Architecture of air-filled caves within the karst of the [email protected] Brooksville Ridge, west-central Florida Conservation-Life Sciences Lee J. Florea 64 Julian J. Lewis & Salisa L. Lewis Lewis & Associates LLC Article Cave, Karst & Groundwater Biological Consulting 17903 State Road 60 Age constraints on cave development and landscape evolution Borden, IN 47106-8608 in the Bighorn Basin of Wyoming, USA 812-283-6120 Greg M. Stock, Catherine A. Riihimaki, and Robert S. Anderson 76 [email protected] Earth Sciences-Journal Index Article Ira D. Sasowsky A revision of the genus Typhlogastrura in North American caves Department of Geology University of Akron with description of five new species Akron, OH 44325-4101 Kenneth Christiansen and Hanghang Wang 85 330-972-5389 [email protected] World Karst Science Reviews 99 Exploration Paul Burger Cave Resources Office Cave Science News 103 3225 National Parks Highway Carlsbad, NM 88220 (505)785-3106 [email protected] Paleontology Greg McDonald Geologic Resource Divison National Park Service P.O. Box 25287 Denver, CO 80225 303-969-2821 [email protected] Social Sciences Joseph C. Douglas History Department Volunteer State Community College 1480 Nashville Pike Gallatin, TN 37066 615-230-3241 [email protected] The Journal of Cave and Karst Studies (ISSN 1090-6924, CPM Number #40065056) is a multi-disciplinary, refereed Book Reviews journal published three times a year by the National Speleological Society, 2813 Cave Avenue, Huntsville, Alabama Arthur N. Palmer & Margaret V. Palmer 35810-4431 USA; (256) 852-1300; FAX (256) 851-9241, e-mail: [email protected]; World Wide Web: Department of Earth Sciences http://www.caves.org/pub/journal/. The annual subscription fee is $23 US, $44 US for 2 years, and $65 US for 3 years. State University of New York Check the Journal website for international rates. Back issues and cumulative indices are available from the NSS Oneonta, NY 13820-4015 office. POSTMASTER: send address changes to the Journal of Cave and Karst Studies, 2813 Cave Avenue, Huntsville, 607-432-6024 Alabama 35810-4431 USA. [email protected] The Journal of Cave and Karst Studies is covered by the following ISI Thomson Services: Science Citation Index Proofreader Expanded, ISI Alerting Services, and Current Contents/Physical, Chemical, and Earth Sciences. Donald G. Davis Copyright © 2006 by the National Speleological Society, Inc. Printed on recycled paper by American Web, 4040 JOURNAL ADVISORY BOARD Dahlia Street, Denver, Colorado 80216 USA Calvin Alexander Barbara am Ende Front cover: Microbiological activity in caves. See Hazel A. Barton, p. 42–53. Hazel A. Barton Carol Hill Back cover: Lance Elder, Kali Pace-Graczyk, and Lee Florea survey the main passage of Blowing Hole Cave in the Horton Hobbs III Julia James Withlacoochee State Forest in Citrus County, Florida (photo by Tom Turner). See Lee J. Florea, p. 64–75. Kathy Lavoie Joyce Lundberg Stephen Worthington Hazel A. Barton – Introduction to cave microbiology: A review for the non-specialist. Journal of Cave and Karst Studies, v. 68, no. 2, p. 43–54. INTRODUCTION TO CAVE MICROBIOLOGY: A REVIEW FOR THE NON-SPECIALIST HAZEL A. BARTON Department of Biological Sciences, Northern Kentucky University, Highland Heights, KY 41099 Within the past decade there has been an increasing interest in cave microbiota. Such interest has helped many speleologists both recognize and understand the importance of microbial species in caves, which has led to improvements in cave conservation practices to better conserve these unseen ecosystems. While much information on the metabolic properties and functions of such subterranean ecosystems has been published in the microbial ecology literature, is it sometimes unusable by the non-specialist due to technical “jargon” and unexplained background information. It is the aim of this review to provide such background information and to explain the current technologies available to study cave microbiota. In doing so, it is hoped that this material will make the microbiology literature more accessible to interest- ed non-specialists, and open new areas of inquiry in the study of microbial-mineral interactions. INTRODUCTION it would take two thousand of them stacked end-to-end to tra- verse the head of a pin, which Leeuwenhoek accurately The study of cave microbiology deals with the microscopic life described in great detail. that resides in caves. Microscopic life has historically been Leeuwenhoek’s animalcules were considered nothing more grouped into the Monera, one of the five kingdoms of life, and than a curiosity for almost 200 years, until the golden age of a pretty broad group that includes any microscopic organism microbiology and the work of such luminaries as Pasteur and that lacks a nucleus; also historically known as the prokaryotes Koch. Louis Pasteur wanted to understand the then-accepted (Whittaker, 1969). The protozoa, which are also microscopic notion of spontaneous generation – that a vital force allowed and include such organisms as the amoeba, do contain a nucle- life to arise spontaneously in food or wine. In attempting to us and are classed within the eukaryotes (Whittaker, 1969). define the existence of this vital force, in 1861 Pasteur suc- While viruses are also of microscopic, they are not technically cessfully demonstrated that this “vital force” was comprised of alive and, although fascinating, will not be discussed within microorganisms in the air. Pasteur subsequently went on to the scope of this review. Animals, plants and fungi all consti- design ways of killing these organisms in food to prevent tute the forms of life that can be seen with the naked eye spoilage and developed the technique of pasteurization. Robert (macroscopic) and are nucleus-containing eukaryotes. Koch took the principles of Pasteur and other investigators fur- ther, to show that the same microorganisms that could spoil ABRIEF HISTORY OF MICROBIOLOGY food could also cause disease in humans. In 1876 Koch devel- oped the techniques to identify the causative agent of anthrax. During the Dark Ages (c.400-700 AD) devastating dis- These techniques became an accepted scientific principle, eases, such as the Black Death, ravaged humanity. At that time, termed Koch’s postulates, that can be applied to any pathogen- there was only a vague inkling in medicine that these maladies ic microbe today, including as recently as in the case of severe were transmitted by an infectious agent. Rather, physicians acute respiratory syndrome (SARS) virus. Following the work believed in mystical spirits and lethal miasmas, which were of Pasteur, Koch and other eminent microbiologists, responsible for such pestilence. Patients would attempt to ward Leeuwenhoek’s animalcules went from being simply a curios- off these evil spirits with charms and even flowers; the “pock- ity to the basis of the accepted science of microbiology. et-full of posies” of the nursery rhyme. These attempts were With its founding in the medical sciences, the study of somewhat beneficial, although not for immediately obvious microorganisms has historically concentrated on the impact of reasons – breathing through containers of dried flowers effec- microorganisms on humans, whether through disease or food tively filtered out the bacteria that were responsible for the spoilage. Nonetheless, in 1887 Sergei Winogradsky made a spread of pneumonic plague, perpetuating the myth of evil groundbreaking discovery; he identified microorganisms that vapors. did not obtain food through photosynthesis, but could use min- The first awareness of a microscopic world occurred eral energy to literally pull food out of thin air, a process around 300 years ago, when Dutch linen merchant Antoni van termed chemolithotrophy (chemical energy – chemo, food Leeuwenhoek devised a rudimentary, hand-made microscope from the environment – litho, to eat – trophy) (Winogradsky, to look at linen fibers. Leeuwenhoek also used his microscope 1887). Despite the identification of microorganisms that defied to look at other materials; in pond water he saw tardigrades (for slow-walker), small insect-like organisms he termed ani- Corresponding author address: Hazel A. Barton, Department of Biological malcules. Eventually, by refining his microscope lens, Sciences, Northern Kentucky University, SC 204D, Nunn Drive, Highland Leeuwenhoek went on to describe the first bacteria, which he Heights, KY 41099. Email: [email protected],