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Microbiology and Immunology PDF

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WOMI1.tpgs 5/8/03 6:01 PM Page 1 AM N I C WORLD of D R I O M B M I O U L N O O G L Y O G Y WOMI1.tpgs 5/8/03 6:01 PM Page 3 AM N I C WORLD of D R I O M B M I O U Brigham Narins, Editor L N O Volumes 1 and 2 O G A-Z L Y O G Y womi_fm 5/6/03 1:34 PM Page iv World of Microbiology and Immunology K. Lee Lerner and Brenda Wilmoth Lerner, Editors Project Editor Permissions Product Design Brigham Narins Margaret Chamberlain, Jackie Jones Michael Logusz Editorial Imaging and Multimedia Manufacturing Mark Springer Leitha Etheridge-Sims, Mary K. Grimes, Lezlie Rhonda Williams Light, Dan Newell, David G. Oblender, Christine O’Bryan, Robyn V. Young © 2003 by Gale. Gale is an imprint of The Gale For permission to use material from this While every effort has been made to Group, Inc., a division of Thomson Learning, product, submit your request via Web at ensure the reliability of the information pre- Inc. http://www.gale-edit.com/permissions, or you sented in this publication, The Gale Group, may download our Permissions Request form Inc. does not guarantee the accuracy of the Gale and Design™ and Thomson Learning™ and submit your request by fax or mail to: data contained herein. The Gale Group, Inc. are trademarks used herein under license. accepts no payment for listing; and inclusion Permissions Department in the publication of any organization, For more information, contact The Gale Group, Inc. agency, institution, publication, service, or The Gale Group, Inc. 27500 Drake Road individual does not imply endorsement of the 27500 Drake Road Farmington Hills, MI, 48331-3535 editors or publisher. Errors brought to the Farmington Hills, MI 48331-3535 Permissions hotline: attention of the publisher and verified to the Or you can visit our Internet site at 248-699-8074 or 800-877-4253, ext. 8006 satisfaction of the publisher will be corrected http://www.gale.com Fax: 248-699-8074 or 800-762-4058. in future editions. ALL RIGHTS RESERVED No part of this work covered by the copyright hereon may be reproduced or used in any form or by any means—graphic, electronic, or mechanical, including photocopying, record- ing, taping, Web distribution, or information storage retrieval systems—without the written permission of the publisher. LIBRARY OF CONGRESS CATALOGING-IN-PUBLICATION DATA World of microbiology and immunology / K. Lee Lerner and Brenda Wilmoth Lerner, editors. p. ; cm. Includes bibliographical references and index. ISBN 0-7876-6540-1 (set : alk. paper)— ISBN 0-7876-6541-X (v. 1 : alk. paper)— ISBN 0-7876-6542-8 (v. 2 : alk. paper) 1. Microbiology—Encyclopedias. 2. Immunology—Encyclopedias. [DNLM: 1. Allergy and Immunology—Encyclopedias—English. 2. Microbiology—Encyclopedias—English. QW 13 W927 2003] I. Lerner, K. Lee. II. Lerner, Brenda Wilmoth. QR9 .W675 2003 579’.03—dc21 2002010181 ISBN: 0-7876-6541-X Printed in the United States of America 10 9 8 7 6 5 4 3 2 1 womi_fm 5/6/03 1:34 PM Page v CONTENTS • INTRODUCTION . . . . . . . . . . . . . . . . . .vii HOW TO USE THIS BOOK . . . . . . . . . .ix ACKNOWLEDGMENTS . . . . . . . . . . . . .xiii ENTRIES Volume 1: A-L . . . . . . . . . . . . . . . . . .1 Volume 2: M-Z . . . . . . . . . . . . . . . .359 SOURCES CONSULTED . . . . . . . . . . . .619 HISTORICAL CHRONOLOGY . . . . . . .643 GENERAL INDEX . . . . . . . . . . . . . . . .661 v • womi_fm 5/6/03 1:34 PM Page vii INTRODUCTION • Although microbiology and immunology are fundamen- During the early part of the twentieth century, the science tally separate areas of biology and medicine, they combine to of microbiology developed somewhat independently of other provide a powerful understanding of human health and dis- biological disciplines. Although for many years it did not exist ease—especially with regard to infectious disease, disease as a separate discipline at all—being an “off-shoot” of chem- prevention, and tragically, of the growing awareness that istry (fermentation science) or medicine—with advances in bioterrorism is a real and present worldwide danger. techniques such as microscopy and pure culturing methodolo- World of Microbiology and Immunologyis a collection of gies, as well as with the establishment of the germ theory of 600 entries on topics covering a range of interests—from biog- disease and the rudiments of vaccination, microbiology sud- raphies of the pioneers of microbiology and immunology to denly exploded as a separate discipline. Whereas other biolog- explanations of the fundamental scientific concepts and latest ical disciplines were concerned with such topics as cell struc- research developments. In many universities, students in the ture and function, the ecology of plants and animals, the repro- biological sciences are not exposed to microbiology or duction and development of organisms, the nature of heredity immunology courses until the later half of their undergraduate and the mechanisms of evolution, microbiology had a very dif- studies. In fact, many medical students do not receive their ferent focus. It was concerned primarily with the agents of first formal training in these subjects until medical school. infectious disease, the immune response, the search for Despite the complexities of terminology and advanced knowl- chemotherapeutic agents and bacterial metabolism. Thus, edge of biochemistry and genetics needed to fully explore from the very beginning, microbiology as a science had social some of the topics in microbiology and immunology, every applications. A more detailed historical perspective of the effort has been made to set forth entries in everyday language development of the field may be found in the article “History and to provide accurate and generous explanations of the most of Microbiology” in this volume. important terms. The editors intend World of Microbiology Microbiology established a closer relationship with other and Immunologyfor a wide range of readers. Accordingly, the biological disciplines in the 1940s because of its association articles are designed to instruct, challenge, and excite less with genetics and biochemistry. This association also laid the experienced students, while providing a solid foundation and foundations for the subsequent and still rapidly developing reference for more advanced students. The editors also intend field of genetic engineering, which holds promise of profound that World of Microbiology and Immunology be a valuable impact on science and medicine. resource to the general reader seeking information fundamen- Microorganisms are extremely useful experimental sub- tal to understanding current events. jects because they are relatively simple, grow rapidly, and can Throughout history, microorganisms have spread deadly be cultured in large quantities. George W. Beadle and Edward diseases and caused widespread epidemics that threatened and L. Tatum studied the relationship between genes and enzymes altered human civilization. In the modern era, civic sanitation, in 1941 using mutants of the bread mold Neurospora. In 1943 water purification, immunization, and antibiotics have dramat- Salvador Luria and Max Delbrück used bacterial mutants to ically reduced the overall morbidity and the mortality of dis- show that gene mutations were apparently spontaneous and ease in advanced nations. Yet much of the world is still rav- not directed by the environment. Subsequently, Oswald aged by disease and epidemics, and new threats constantly Avery, Colin M. MacLeod, and Maclyn McCarty provided appear to challenge the most advanced medical and public strong evidence that DNAwas the genetic material and car- health systems. For all our science and technology, we are far ried genetic information during transformation. The interac- from mastering the microbial world. tions between microbiology, genetics, and biochemistry soon vii • womi_fm 5/6/03 1:34 PM Page viii Introduction WORLDOF MICROBIOLOGY & IMMUNOLOGY • led to the development of modern, molecularly oriented made to prevent their spread from sick to healthy people. The genetics. great successes that have taken place in the area of human Recently microbiology has been a major contributor to health in the past 100 years have resulted largely from the rise of molecular biology, the branch of biology dealing advances in the prevention and treatment of infectious disease. with the physical and chemical bases of living matter and its We can consider the eradication of smallpox, a viral disease, function. Microbiologists have been deeply involved in stud- as a prime example. The agent that causes this disease is one ies of the genetic code and the mechanisms of DNA, RNA, of the greatest killers the world has ever known—and was and protein synthesis. Microorganisms were used in many of probably the greatest single incentive towards the formaliza- the early studies on the regulation of gene expression and the tion of the specialized study of immunology. Research into the control of enzyme activity. In the 1970s new discoveries in mechanism of Edward Jenner’s “vaccination” discovery—he microbiology led to the development of recombinant gene found that of a patient injected with cow-pox produces immu- technology and genetic engineering. One indication of the nity to smallpox—laid the foundations for the understanding importance of microbiology today is the number of Nobel of the immune system and the possibility of dealing with other Prizes awarded for work in physiology and medicine during diseases in a similar way. Because of an active worldwide vac- the twentieth century; about a third of these were awarded to cination program, no cases of smallpox have been reported scientists working on microbiological problems. since 1977. (This does not mean, however, that the disease Microorganisms are exceptionally diverse, are found cannot reappear, whether by natural processes or bioterror.) almost everywhere, and affect human society in countless Another disease that had a huge social impact was bubon- ways. The modern study of microbiology is very different ic plague, a bacterial disease. Its effects were devastating in from the chemically and medically oriented discipline pio- the Middle Ages. Between 1346 and 1350, one third of the neered by Louis Pasteur and Robert Koch. Today it is a large entire population of Europe died of bubonic plague. Now gen- discipline with many specialities. It has impact on medicine, erally less than 100 people die each year from this disease. The agricultural and food sciences, ecology, genetics, biochem- discovery of antibiotics in the early twentieth century provid- istry, and many other fields. Today it clearly has both basic and ed an increasingly important weapon against bacterial dis- applied aspects. eases, and they have been instrumental in preventing similar Many microbiologists are interested in the biology of the plague epidemics. microorganisms themselves. They may focus on a specific Although progress in the application of immunological group of microorganisms and be called virologists (scientists research has been impressive, a great deal still remains to be who study viruses), bacteriologists (scientists who study bac- done, especially in the treatment of viral diseases (which do teria), phycologists or algologists (scientists who study algae), not respond to antibiotics) and of the diseases prevalent in mycologists (scientists who study fungi), or protozoologists developing countries. Also, seemingly “new” diseases contin- (scientists who study protozoa). Others may be interested in ue to arise. Indeed, there has been much media coverage in the microbial morphology or particular functional processes and past twenty years in the U.S. of several “new” diseases, work in fields such as microbial cytology, physiology, ecolo- including Legionnaires’disease, toxic shock syndrome, Lyme gy, genetics, taxonomy, and molecular biology. Some microbi- disease, and acquired immunodeficiency syndrome (AIDS). ologists may have a more applied orientation and work on Three other diseases emerged in 1993. In the summer of that problems in fields such as medical microbiology, food and year a mysterious flu-like disease struck the Southwest, result- dairy microbiology, or public health. Because the various ing in 33 deaths. The causative agent was identified as a virus, fields of microbiology are interrelated, an applied microbiolo- hantavirus, carried by deer mice and spread in their droppings. gist must always be familiar with basic microbiology. For In the same year, more than 500 residents of the state of example, a medical microbiologist must have a good under- Washington became ill with a strain of Escherichia colipres- standing of microbial taxonomy, genetics, immunology, and ent in undercooked beef prepared at a fast-food restaurant. The physiology to identify and properly respond to the pathogen of organism synthesized a potent toxin and caused haemolytic- concern. uremic syndrome. Three children died. In 1993, 400,000 peo- It is clear that scientists study the microbial world in ple in Milwaukee became ill with a diarrheal disease, cryp- much the same way as they studied the world of multicellular tosporidiosis, that resulted from the improper chlorination of organisms at the beginning of the twentieth century, when the water supply. microbiology was a young discipline. This is in part due to the It is a great credit to the biomedical research community huge developments and refinements of techniques, which now that the causative agents for all these diseases were identified allow scientists to more closely and fully investigate the world very soon after the outbreaks. The bacteria causing of bacteria and viruses. Legionnaires’disease and Lyme disease have only been iso- One of the focuses of this book is the field of medical lated in the past few decades, as have the viruses that cause microbiology and its connection with immunology. Medical AIDS. Anumber of factors account for the fact that seeming- microbiology developed between the years 1875 and 1918, ly “new” diseases arise almost spontaneously, even in indus- during which time many disease-causing bacteria were identi- trially advanced countries. As people live longer, their ability fied and the early work on viruses begun. Once people realized to ward off infectious agents is impaired and, as a result, the that these invisible agents could cause disease, efforts were organisms that usually are unable to cause disease become viii • womi_fm 5/6/03 1:34 PM Page ix WORLDOF MICROBIOLOGY & IMMUNOLOGY Introduction • potentially deadly agents. Also, lifestyles change and new Because information in immunology often moves rapidly opportunities arise for deadly agents. For example, the use of from the laboratory to the clinical setting, it is increasingly vaginal tampons by women has resulted in an environment in important that scientifically literate citizens—those able to which the Staphylococcus bacterium can grow and produce a participate in making critical decisions regarding their own toxin causing toxic shock syndrome. New diseases can also health care—hold a fundamental understanding of the essen- emerge because some agents have the ability to change abrupt- tial concepts in both microbiology and immunology. ly and thereby gain the opportunity to infect new hosts. It is Alas, as if the challenges of nature were not sufficient, the possible that one of the agents that causes AIDS arose from a evolution of political realities in the last half of the twentieth virus that at one time could only infect other animals. century clearly points toward the probability that, within the Not only are new diseases appearing but many infectious first half of the twenty-first century, biological weapons will diseases that were on the wane in the U.S. have started to surpass nuclear and chemical weapons as a threat to civiliza- increase again. One reason for this resurgence is that thou- tion. Accordingly, informed public policy debates on issues of sands of U.S. citizens and foreign visitors enter the country biological warfare and bioterrorism can only take place when daily. About one in five visitors now come from a country there is a fundamental understanding of the science underpin- where diseases such as malaria, cholera, plague, and yellow ning competing arguments. fever still exist. In developed countries these diseases have The editors hope that World of Microbiology and been largely eliminated through sanitation, vaccination, and Immunology inspires a new generation of scientists who will quarantine. Ironically, another reason why certain diseases are join in the exciting worlds of microbiological and immuno- on the rise is the very success of past vaccination programs: logical research. It is also our modest wish that this book pro- because many childhood diseases (including measles, mumps, vide valuable information to students and readers regarding whooping cough, and diphtheria) have been effectively con- topics that play an increasingly prominent role in our civic trolled in both developed and developing countries, some par- debates, and an increasingly urgent part of our everyday lives. ents now opt not to vaccinate their children. Thus if the disease K. Lee Lerner& Brenda Wilmoth Lerner, editors suddenly appears, many more children are susceptible. St. Remy, France Athird reason for the rise of infectious diseases is that the June 2002 increasing use of medications that prolong the life of the eld- erly, and of treatments that lower the disease resistance of Editor’s note: World of Microbiology and Immunologyis patients, generally weaken the ability of the immune system to not intended to be a guide to personal medical treatment or fight diseases. People infected with human immunodeficiency emergency procedures. Readers desiring information related virus (HIV), the virus responsible for AIDS, are a high-risk to personal issues should always consult with their physician. group for infections that their immune systems would normal- The editors respectfully suggest and recommend that readers ly resist. For this reason, tuberculosis (TB) has increased in the desiring current information related to emergency protocols— U.S. and worldwide. Nearly half the world’s population is especially with regard to issues and incidents related to bioter- infected with the bacterium causing TB, though for most peo- rorism—consult the United States Centers for Disease Control ple the infection is inactive. However, many thousands of new and Prevention (CDC) website at http://www.cdc.gov/. cases of TB are reported in the U.S. alone, primarily among How to Use the Book the elderly, minority groups, and people infected with HIV. Furthermore, the organism causing these new cases of TB is The articles in the book are meant to be understandable resistant to the antibiotics that were once effective in treating by anyone with a curiosity about topics in microbiology or the disease. This phenomenon is the result of the uncontrolled immunology. Cross-references to related articles, definitions, overuse of antibiotics over the last 70 years. and biographies in this collection are indicated by bold-faced Until a few years ago, it seemed possible that the terrible type, and these cross-references will help explain and expand loss of life associated with the plagues of the Middle Ages or the individual entries. Although far from containing a compre- with the pandemic influenza outbreak of 1918 and 1919 would hensive collection of topics related to genetics, World of never recur. However, the emergence of AIDS dramatizes the Microbiology and Immunology carries specifically selected fact that microorganisms can still cause serious, incurable, topical entries that directly impact topics in microbiology and life-threatening diseases. With respect to disease control, there immunology. For those readers interested in genetics, the edi- is still much microbiological research to be done, especially in tors recommend Gale’s World of Geneticsas an accompanying relation to the fields of immunology and chemotherapy. reference. For those readers interested in additional informa- Recent advances in laboratory equipment and techniques tion regarding the human immune system, the editors recom- have allowed rapid progress in the articulation and under- mend Gale’s World of Anatomy and Physiology. standing of the human immune system and of the elegance of This first edition of World of Microbiology and the immune response. In addition, rapidly developing knowl- Immunologyhas been designed with ready reference in mind: edge of the human genome offers hope for treatments designed • Entries are arranged alphabetically rather than to effectively fight disease and debilitation both by directly chronologically or by scientific field. In addition to clas- attacking the causative pathogens, and by strengthening the sical topics, World of Microbiology and Immunology body’s own immune response. contains many articles addressing the impact of ix • womi_fm 5/6/03 1:34 PM Page x Introduction WORLDOF MICROBIOLOGY & IMMUNOLOGY • advances in microbiology and immunology on history, Member, American Society for Microbiology and the ethics, and society. Canadian Society of Microbiologists • Bold-faced termsdirect the reader to related entries. Nova Scotia, Canada • “See also” references at the end of entries alert the Eric v.d. Luft, Ph.D., M.L.S. reader to related entries not specifically mentioned in Curator of Historical Collections the body of the text. SUNYUpstate Medical University • ASources Consultedsection lists the most worthwhile Syracuse, New York print material and web sites we encountered in the com- pilation of this volume. It is there for the inspired read- Danila Morano, M.D. er who wants more information on the people and dis- University of Bologna coveries covered in this volume. Bologna, Italy • The Historical Chronology includes many of the sig- Judyth Sassoon, Ph.D., ARCS nificant events in the advancement of microbiology and Department of Biology & Biochemistry immunology. The most current entries date from just days before World of Microbiology and Immunology University of Bath went to press. Bath, England • Acomprehensive General Index guides the reader to Constance K. Stein, Ph.D. topics and persons mentioned in the book. Bolded page Director of Cytogenetics, Assistant Director of Molecular references refer the reader to the term’s full entry. Diagnostics Although there is an important and fundamental link SUNYUpstate Medical University between the composition and shape of biological molecules Syracuse, New York and their functions in biological systems, a detailed under- standing of biochemistry is neither assumed or required for Acknowledgments World of Microbiology and Immunology. Accordingly, stu- In addition to our academic and contributing advisors, it dents and other readers should not be intimidated or deterred has been our privilege and honor to work with the following by the complex names of biochemical molecules (especially contributing writers, and scientists: Sherri Chasin Calvo; the names for particular proteins, enzymes, etc.). Where nec- Sandra Galeotti, M.S.; Adrienne Wilmoth Lerner; Jill Liske, essary, sufficient information regarding chemical structure is provided. If desired, more information can easily be obtained M.Ed.; and Susan Thorpe-Vargas, Ph.D. from any basic chemistry or biochemistry reference. Many of the advisors for World of Microbiology and Immunology authored specially commissioned articles within Advisory Board their field of expertise. The editors would like to specifically acknowledge the following contributing advisors for their spe- In compiling this edition we have been fortunate in being cial contributions: able to rely upon the expertise and contributions of the follow- ing scholars who served as academic and contributing advisors Robert G. Best, Ph.D. for World of Microbiology and Immunology, and to them we Immunodeficiency disease syndromes would like to express our sincere appreciation for their efforts to Immunodeficiency diseases, genetic ensure that World of Microbiology and Immunologycontains the most accurate and timely information possible: Antonio Farina, M.D., Ph.D. Reproductive immunology Robert G. Best, Ph.D. Director, Division of Genetics, Department of Obstetrics and Brian D. Hoyle, Ph.D. Gynecology Anthrax, terrorist use of as a biological weapon University of South Carolina School of Medicine Eric v.d. Luft, Ph.D., M.L.S. Columbia, South Carolina The biography of Dr. Harry Alfred Feldman Antonio Farina, M.D., Ph.D. Danila Morano, M.D. Visiting Professor, Department of Pathology and Laboratory Rh and Rh incompatibility Medicine Brown University School of Medicine Judyth Sassoon, Ph.D. Providence, Rhode Island BSE and CJD disease, ethical issues and socio-economic Professor, Department of Embryology, Obstetrics, and impact Gynecology University of Bologna Constance K. Stein, Ph.D. Bologna, Italy Genetic identification of microorganisms Brian D. Hoyle, Ph.D. Susan Thorpe-Vargas, Ph.D Microbiologist Immunology, nutritional aspects x • womi_fm 5/6/03 1:34 PM Page xi WORLDOF MICROBIOLOGY & IMMUNOLOGY Introduction • The editors would like to extend special thanks Dr. Judyth complexities and difficulties related to graphics. Most direct- Sassoon for her contributions to the introduction to World of ly, the editors wish to acknowledge and thank the Project Microbiology and Immunology. The editors also wish to Editor, Mr. Brigham Narins for his good nature, goods eyes, acknowledge Dr. Eric v.d. Luft for his diligent and extensive and intelligent sculptings of World of Microbiology and research related to the preparation of many difficult biogra- Immunology. phies. The editors owe a great debt of thanks to Dr. Brian The editors dedicate this book to Leslie Moore, M.D., Hoyle for his fortitude and expertise in the preparation and James T. Boyd, M.D., E.M. Toler, M.D., and to the memory of review of a substantial number of articles appearing in World Robert Moore, M.D. Their professional skills and care provid- of Microbiology and Immunology. ed a safe start in life for generations of children, including our The editors gratefully acknowledge the assistance of own. many at Gale for their help in preparing World of The editors and authors also dedicate this book to the Microbiology and Immunology. The editors thank Ms. countless scientists, physicians, and nurses who labor under Christine Jeryan and Ms. Meggin Condino for their faith in the most dangerous and difficult of field conditions to bring this project. Special thanks are offered to Ms. Robyn Young both humanitarian assistance to those in need, and to advance and the Gale Imaging Team for their guidance through the the frontiers of microbiology and immunology. xi •

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