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Page 1: Microbiology 10th edition by Tortora
Page 2: Microbiology 10th edition by Tortora

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AN INTRODUCTION

TENTH EDITION

Gerard J. Tortora BERGEN COMMUNITY COLLEGE

Berdell R. Funke NORTH DAKOTA STATE UNIVERSITY

Christine L. Case SKYLINE COLLEGE

Benjamin Cummings San Francisco Boston New York

Cape Town Hong Kong London Madrid Mexico City Montreal Munich Paris Singapore Sydney Thkyo Thronto

Page 3: Microbiology 10th edition by Tortora

Executive Editor: Leslie Berriman Project Editor: Robin Pille Development Manager: Barbara Yien Assistant Editors: Katie Heimsoth and Kelly Reed Managing Editor: Wendy Earl Production Supervisor: Janet Vail Director, Media Development: Lauren Fogel Media Producer: Lucinda Bingham Interior Designer: tani hasegawa Cover Design: Riezebos Holzbaur Design Group Art Coordinator: Jean Lake

Cover Photo Credit: Science Photo Library/Eric Graves Text art, and photo credits appear after the Glossary.

Art Development Editor: Lisa Torri Artists: Precision Graphics Copyeditor: Sally Peyrefitte Proofreader: Martha Ghent Senior Art and Photo Manager: Travis Amos Photo Researcher: Maureen Spuhler Image Rights and Permissions Manager: Zina Arabia Senior Manufacturing Buyer: Stacey Weinberger Senior Marketing Manager: Neena Bali Executive Market Development Manager: Josh Frost

Copyright © 2010 Pearson Education, [nc., publishing as Pearson Benjamin Cummings, 1301 Sansome St., San Francisco, CA 94111. All rights reserved. Manufactured in the United States of America. This publication is protected by Copyright and permission should be obtained from the publisher prior to any prohibited reproduction, storage in a retrieval system, or transmission in any form or by any means, electronic, mechanical, photocopying, recording, or likewise. To obtain permission(s) to use material from this work, please submit a written request to Pearson Education, Inc., Permissions Department, 1900 E. Lake Ave., Glenview, IL 60025. For information regarding permissions, call (847) 486-2635.

Many of the designations used by manufacturers and sellers to distinguish their products are claimed as trademarks. Where those designations appear in this book, and the publisher was aware of a trademark claim, the designations have been printed in initial caps or all caps.

Pearson Benjamin Cummings is a trademark, in the U.S. and/or other countries, of Pearson Education, Inc. or its afffiliates.

Library of Congress Cataloging-in-Publication Data Tortora, Gerard J. Microbiology: an introduction / Gerard J. Tortora, Berdell R. Funke, Christine L. Case. - 10th ed.

p. cm. Includes bibliographical references and index. ISBN-13: 978-0-321-55007-1 (Student ed.) ISBN-IO: 0-321-55007-2 (Student ed.) ISBN-13: 978-0-321-58202-7 (Professional copy) ISBN-IO: 0-321-58202-0 (Professional copy) I. Microbiology-Textbooks. I. Funke, Berdell R. II. Case, Christine L., 1948- III . Title. [DNLM: I . Microbiology. QW 4 T712m 20 I 0 I

QR41.2.T672010 579-dc22

Benjamin Cummings is an imprint of

PEARSON -

2008044384

_.pearsonhighered.com

ISBN 10: 0-321-55007-2; ISBN 13: 978-0-321-55007-1 (Student edition) ISBN 10: 0-321-58202-0; ISBN 13: 978-0-321-58202-7 (Professional copy)

I 2 3 4 5 6 7 8 9 IO-DOW-12 II 10 09 08 Manufactured in the United States of America.

Page 4: Microbiology 10th edition by Tortora

Executive Editor: Leslie Berriman Project Editor: Robin Pille Development Manager: Barbara Yien Assistant Editors: Katie Heimsoth and Kelly Reed Managing Editor: Wendy Earl Production Supervisor: Janet Vail Director, Media Development: Lauren Fogel Media Producer: Lucinda Bingham Interior Designer: tani hasegawa Cover Design: Riezebos Holzbaur Design Group Art Coordinator: Jean Lake

Cover Photo Credit: Science Photo Library/Eric Graves Text art, and photo credits appear after the Glossary.

Art Development Editor: Lisa Torri Artists: Precision Graphics Copyeditor: Sally Peyrefitte Proofreader: Martha Ghent Senior Art and Photo Manager: Travis Amos Photo Researcher: Maureen Spuhler Image Rights and Permissions Manager: Zina Arabia Senior Manufacturing Buyer: Stacey Weinberger Senior Marketing Manager: Neena Bali Executive Market Development Manager: Josh Frost

Copyright © 2010 Pearson Education, [nc., publishing as Pearson Benjamin Cummings, 1301 Sansome St., San Francisco, CA 94111. All rights reserved. Manufactured in the United States of America. This publication is protected by Copyright and permission should be obtained from the publisher prior to any prohibited reproduction, storage in a retrieval system, or transmission in any form or by any means, electronic, mechanical, photocopying, recording, or likewise. To obtain permission(s) to use material from this work, please submit a written request to Pearson Education, Inc., Permissions Department, 1900 E. Lake Ave., Glenview, IL 60025. For information regarding permissions, call (847) 486-2635.

Many of the designations used by manufacturers and sellers to distinguish their products are claimed as trademarks. Where those designations appear in this book, and the publisher was aware of a trademark claim, the designations have been printed in initial caps or all caps.

Pearson Benjamin Cummings is a trademark, in the U.S. and/or other countries, of Pearson Education, Inc. or its afffiliates.

Library of Congress Cataloging-in-Publication Data Tortora, Gerard J. Microbiology: an introduction / Gerard J. Tortora, Berdell R. Funke, Christine L. Case. - 10th ed.

p. cm. Includes bibliographical references and index. ISBN-13: 978-0-321-55007-1 (Student ed.) ISBN-IO: 0-321-55007-2 (Student ed.) ISBN-13: 978-0-321-58202-7 (Professional copy) ISBN-IO: 0-321-58202-0 (Professional copy) I. Microbiology-Textbooks. I. Funke, Berdell R. II. Case, Christine L., 1948- III . Title. [DNLM: I . Microbiology. QW 4 T712m 20 I 0 I

QR41.2.T672010 579-dc22

Benjamin Cummings is an imprint of

PEARSON -

2008044384

_.pearsonhighered.com

ISBN 10: 0-321-55007-2; ISBN 13: 978-0-321-55007-1 (Student edition) ISBN 10: 0-321-58202-0; ISBN 13: 978-0-321-58202-7 (Professional copy)

I 2 3 4 5 6 7 8 9 IO-DOW-12 II 10 09 08 Manufactured in the United States of America.

Page 5: Microbiology 10th edition by Tortora

ABOUT THE AUTHORS

Gerard J. Tortora Jerry Tortora is a professor of biology and teaches microbiology, human anatomy, and physiology at Bergen Community College in Paramus, New Jersey. He received his M.A. in Biology from Montclair State College in 1965. He belongs to a number of biology/microbiology organizations. such as the American Society of Microbiology (ASM). Human Anatomy and Physiology Society (HAPS),

American Association for the Advancement of Science (AAAS), National Education Association (NEA), New Jersey Educational Associat ion (NJEA), and the Metropolitan Association of College and University Biologists (MAGUB). Jerry is the author of numerous biological science textbooks. In 1995, he was selected as one of the finest faculty scholars of Bergen Community College and was named Distinguished Faculty Scholar. In 1996. Jerry received a National Inst itute for Staff and Organizational Development (NISOD) excellence award from the University of Texas and was selected to represent Bergen Community College in a campaign to increase awareness of the contributions of community colleges to higher education.

Berdell R. Funke Bert Funke received his Ph.D .• M.S .• and B.S. in microbiology from Kansas State University. He has spent his professional years as a professor of microbiology at North Dakota State University. He taught introductory microbiology. including laboratory sections. general microbiology. food microbiology. soil microbiology. clinical parasitology. and pathogenic microbiology. As a research scientist in the Experiment

Station at North Dakota State. he has published numerous papers in soil microbiology and food microbiology.

Christine l. Case Chris Case is a registered microbiologist and a professor of microbiology at Skyline College in San Bruno. California. where she has taught for the past 38 years. She received her Ed.D. in curriculum and instruction from Nova Southeastern University and her M.A. in microbiology from San Francisco State University. She was Director for the Society for Industrial Microbiology (S IM) and is an active

member of the ASM and Northern California SIM. She received the ASM and California Hayward outstanding educator awards. In 2008. Chris received the SACNAS Distinguished CommunityfTribal College Mentor Award for her commitment to her students. several of whom have presented at undergraduate research conferences and won awards. In addition to teaching. Chris contributes regularly to the professional literature. develops innovative educational methodologies. and maintains a personal and professional commitment to conservation and the importance of science in society. Chris is also an avid photographer. and many of her photographs appear in this book.

iii

Page 6: Microbiology 10th edition by Tortora

PREFACE

Since the publication of the fi rst edition nearly 30 years ago, well over one million students have used Microbiology: An Introduction at colleges and universities around the world, mak-ing it the leading textbook for non-majors microbiology. The tenth edition continues to be a comprehensive beginning text, assum ing no previous study of biology or chemistry. The text is appropriate for students in a wide variety of programs, including the allied health sciences, biological science, environmental sci-ences, animal science, forestry, agriculture, home economics, and the liberal arts.

HALLMARKS OF MICROBIOLOGY: AN INTRODUCTION We have retained in this new edition features that made the previous editions so popular:

• An appropriate balance between microbiological funda-mentals and applications, and between medical applica-tions and other applied areas of microbiology. As in previous editions, basic microbiological principles are given greater emphasis than are applications, and health-related applications are featured.

• Straightforward presentation of complex topics. Each sec-tion of the text has been written with the student in mind. Our book is known for its clear explanations and consistent pedagogy.

• Clear, accurate, and pedagogically effective illustrations and photos. Step-by-step diagrams that closely coordinate with narrative descriptions aid student comprehension of concepts. Clear and accurate renderings of processes and structures focus students on what they need to learn. The quantity and quality of electron and light micrographs is unmatched in the market.

• Flexible organization. We have organized the book in what we think is a useful fas hion while recognizing that the mate-rial might be effectively p resented in a number of other sequences. For instructors who wish to use a different order, we have made each chapter as independent as possible and have included numerous cross- references. The Instructor's Guide, written by Christine Case, provides detailed guide-lines for organizing the material in several other ways.

NEW TO THE TENTH EDITION Please see pages x- xviii for a visual introduction to the new tenth edition .

The changes in this edition address instructors' biggest chal-lenge in the introductory microbiology course: the wide range of student levels, including student under-preparedness. The tenth

edition meets all students at their respective levels of skill and understanding.

The highlights of the tenth edition can be seen in the new Foundation Figures, the new features that help studen ts check their understanding as they progress through each chapter, and the improved boxes that prepare students to start thinking like a cli nician. Content and currency have also been substantially updated.

Foundation Figures In order to help students focus on and master the core concepts of microbiology, the authors have integrated text and visuals in to twenty specially designed Foundation Figures. These Foundation Figures include both a key concept statement that ensures stu-dents understand the central concept of the figure and an expla-nation of how each figure is foundational to further learning in the course. In addition, throughout the entire book the illustra-tion program has been dramatically revised and updated with new art styles and a brighter color palette that has more con-trasting colors and more dimensionality.

Features that Help Students Check Their Understanding New Check Your Understanding questions encourage students to engage interactively with the material and self-assess their under-standing of the Learning Objectives as they progress through each chapter. New Draw It questions are now included in the end-of-chapter Review Questions, asking students to sketch a rough diagram or fill in missing parts of a figure or graph. The popular Figure Legend Questions have been retained and im proved.

Support for Students to Start Thinking like a Clinician Revised and redesigned Appl ications of Microbiology boxes describe current and practical uses of microbiology. New and updated Clinical Focus boxes contain data from the Morbidityalld Mortality Weekly Report modified into clinical problem-solving questions that help students develop their critical thinking skills and give them an active role while reading. Diseases in Focus boxes replace disease comparison tables, organizing comparative infor-mation about similar diseases in a d iscovery-oriented and visually interesting format that provides a helpful study tool for students.

Updates in Content and Currency Antimicrobial resistance, biofilms, bioterrorism, and evolution receive special attention and increased emphasis. The immunity chapters-Chapters 16 and 17- have received a careful and sig-

Page 7: Microbiology 10th edition by Tortora

nificant update for increased currency, clarity, and accuracy, with-out an increase in the level of detail. Taxonomy, nomenclature, and d isease incidence data are current through August 2008.

Cutting-Edge Media Supplements Turn to the inside front cover of this book for an overview of the new MyMicrobiologyPlace websi te with its simple th ree-step approach to learning. Pages xiv-xv provide more details about the exciting new student and instructor media, including the MP3 Tutor Sessions, the 3D MicroFlix animations, and the Instructor's Resource DVD/CD -ROM.

CHAPTER-BY-CHAPTER REVISIONS Every chapter in this edition has been thoroughly revised, and data in the text, tables, and figures have been updated through August 2008 where possible. The main changes for each chapter are summarized below.

Part One Fundamentals of Microbiology Chapter 1: The Microbial Wortd and You • The table "Making Scientific Names Familiar" has been

moved to this chapter from Chapter 10. • Biofilms are introduced . • Discussion of emerging infectious diseases has been updated,

including a section on antibiotic-resistant bacteria.

Chapter 2: Chemical Principles • Definitions have been expanded, including definitions of cis

and trans fatty acids. • Figure 2.16 is now a Foundation Figure.

Chapter 3: Observing Microorganisms through a Microscope • Two-photon microscopy is included. • Several new photos illustrate microscopic images. • Figure 3.2 is now a Foundation Figure.

Chapter 4: Functional Anatomy of Prokaryotic and Eukaryotic Cells • Figure 4.6 is now a Foundation Figure. • The discussion of flagella, fimbriae, and pili has been revised,

as has the discussion of the lipopolysaccharide. • The discussion of facilitated diffusion has been revised, and a

new figure compares types of diffusion across membranes, including aquaporins.

Chapter 5: Microbial Metabolism • The section on biochemical tests has been expanded. • New renditions of enzymes are more realistic. • Figure 5.11 is now a Foundation Figure. • A new Clinical Focus box illustrates the use of biochemical

tests to identify slow-growing mycobacteria .

Chapter 6: Microbial Growth • The discussion ofbiofitms previously appearing in Chapter 27

has been moved to this chapter and been significantly updated and expanded.

PREFACE v

• Discussion of anaerobic growth media and methods has been updated.

• A discussion of Biosafety Levels has been added, including a figure illustrating Biosafety Level 4.

• A new figure showing differential medium is included. • Figure 6.15 is now a Foundation Figure. • A new Clinical Focus box illustrates the role of biofilms in

causing nosocomial infections.

Chapter 7: The Control of Microbial Growth • The definition of sterilization has been updated and qualified

in consideration of the existence of prions. • Figure 7.1 is now a Foundation Figure. • Discussion of ultra-h igh temperature (UHT) has been

clarified. • New products and newly approved uses are included. • A new Clinical Focus box illustrates the relationship between

improper disinfection and nosocomial infection .

Chapter 8: Microbial Genetics • Figure 8.2 is now a Foundation Figure. • Discussion of genetic combination by crossing over has been

revised for clarity. • snRNPs are defined. • Inducible and repressible operons are explained and com-

pared in separate figures .

Chapter 9: Biotechnology and Recombinant DNA • Figure 9.1 is now a Foundation Figure. • Gene silencing, reverse genetics, and real-time PCR are

discussed. • A new Clincal Focus box describes using reverse-transcription

PCR to track a norovirus outbreak.

Part Two A Survey of the Microbial World Chapter 10: Classification of Microorganisms • Figure 10.1 is now a Foundation Figure. • Photos of fossil and living stromatolites are included. • The use of transport media is explained.

Chapter 11 : The Prokaryotes: Domains Bacteria and Archaea • Several new bacterial groups are discussed: Pelagibacter,

Acinetobacter baumallii, Planctomycetes, Gemmata obscuriglobus.

• Discussion of the theoretical minimal size of a bacterium and its genetic requirements has been revised.

Chapter 12: The Eukaryotes: Fungi, Algae, Protozoa, and Helminths • Examples of new uses of fungi as pesticides are listed. • Discussion of the oomycotes is expanded to include

introduction of Phytophthora into the United States. The oomycote life cycle is illustrated in a new figure .

• Heartworm is included. • A new Clinical Focus box h ighlights cryptosporidial

diarrhea, the most common pathogen associated with . . sWlmmmg.

Page 8: Microbiology 10th edition by Tortora

vi PREFACE

Chapter 13: Viruses, Viroids, and Prions • The chapter begins with the use of retroviridae to genetically

modify cells. • Figure 13.15 is now a Foundation Figure. • Bee colony collapse is mentioned. • The Clinical Focus box on the evolution and occurrence of

avian flu has been updated.

Part Three Interaction Between Microbe and Host Chapter 14: Principles of Disease and Epidemiology • Figure 14.3 is now a Foundation Figure. • Statistics on notifiable infectious diseases have been updated. • A new Clinical Focus box illustrates the emergence of

hospital-acquired and community-acquired MRSA.

Chapter 15: Microbial Mechanisms of Pathogenicity • Discussion of A-B toxins has been expanded and clarified. • Figure 15.5, action of an exotoxin, has been revised and

expanded. • Figures 15.4 and 15.9 are now Foundation Figures. • A new Clinical Focus box illustrates role of biofilms and

endotoxins in postoperative infections.

Chapter 16: Innate Immunity: Nonspecific Defenses of the Host • Treatment of several topics has been expanded and/or

reorganized and clarified: physical and chemical factors in the first line of defense; fo rmed elements in blood; the lym-phatic system (including additional illustrations); adherence, acute-phase proteins, complement, iron-binding proteins, and antimicrobial peptides.

• The role of biofilms in evading phagocytosis is included. • Figures 16.7 and Figure 16.9 are now Foundation Figures. • The Applications of Microbiology box on serum collection has

been revised to include testing for complement to monitor immune complex diseases in patients.

Chapter 17: Adaptive Immunity: Specific Defenses of the Host • A new photo depicts actual antibody morphology shown by

atomic force microscopy. • Several important figures have been extensively revised for

accuracy and clarity: • Figure 17.5. Clonal selection and differentiation of

B cells • Figure 17.10. Activation ofCD4 + T cells • Figure 17.11. Killing of virus-infected target cell by

cytotoxic T lymphocyte • Figure 17.19. The dual nature of the immune system

(now a Foundation Figure} • A new photo and illustration (Figure 17.9) show M cells

found within Peyer's patches. • Discussion of the major histocompatibility complex (M HC)

has been revised and improved. • Nomenclature conventions have been updated for T cells

(for example, T helper cell, CD4 + T cell).

• The discussions ofT cells, dendritic cells, and cytokines have been completely revised.

• A new Applications of Microbiology box describes the possi-ble use of I L-12 to treat psoriasis.

Chapter 18: Practical Applications of Immunology • Figure 18.2 is now a Foundation Figure. • Discussions of DNA vaccines and adjuvants have been

updated and revised. • The tables of vaccine schedules have been updated . • A new Clinical Focus box illustrates the success of vac-

cination in eliminating measles in the U.S. and highlights the importance of measles as a cause of death in developing countries.

Chapter 19: Disorders Associated with the Immune System • Coverage of blood groups includes a discussion of the rela-

tionship between certain blood groups and their relative resis tance or susceptibility to certain diseases.

• A discussion of the autoimmune disease psoriasis and its associated arthritis has been introduced, along with the current treatments with monoclonal antibodies.

• The discussion of stem cells has been updated, and a new figure (Figure 19.10) the derivation of stem cells and stem cell lines.

• The discussion of HIV and AIDS has been revised and updated. Especially important is the complete revision of Figure 19.13, which shows the sequence of attachment, fusion, and entry of the virus into the target CD4+ T cell.

• Figure 19.16 is now a Foundation Figure.

Chapter 20: Antimicrobial Drugs • Figure 20.2 is now a Foundation Figure. • The historical importance of the sulfa drugs is given more

prommence. • The current methods used for the discovery of new

antibiotics are discussed, including rapid throughput methods.

• The discussion of antibiotics has been updated to admit new antibiotics. The discussion of antivirals for the treatment of HIV / AIDS has been especially updated and revised to include the latest developments in this constantly changing area.

• The discussion of resistance to antibiotics has been com-pletely revised and expanded, and a new Foundation Figure (Figure 20.20) illustrates the most important target areas for resis tance.

• The concluding discussion on the future of antibiotic devel-opment and the prospect for unconventional antibiotics has been completely revised and updated.

Part Four Microorganisms and Human Disease Chapter 21 : Microbial Diseases of the Skin and Eyes • The discussion of Staphylococcus aurerlS has been completely

rewritten to emphasize the importance of MRSA.

Page 9: Microbiology 10th edition by Tortora

• The discussion of impetigo and scalded skin syndrome has been revised, and discussion of a new disease, Buruli ulcer, has been added.

• Some of the newer treatments for acne now have an expanded discussion.

Chapter 22: Microbial Diseases of the Nervous System • A new figure (Figure 22.4) illustrates a spinal tap. • The discussion of cryptococcosis has been revised to include

a newer pathogen. • A brief description of prions has been included to

supplement that given in Chapter 13. • The discussion of chronic disease syndrome has been

completely revised and now includes the CDC's diagnos-tic definition and the alternative name of myalgic en cep halo myel it is.

Chapter 23: Microbial Diseases of the Cardiovascular and lymphatic Systems • The definitions of the similar terms septicemia and sepsis

have been revised. • Discussions of brucellosis and rat-bite fever have been

completely rewritten. • The discussion of ehrlichiosis has been revised to include the

new terminology of anaplasmosis. • Discussion of the disease chikungunya fever has been added

because of its current spread into temperate climates. • The discussion of malaria has been revised completely to

better differentiate between prophylaxis and therapy.

Chapter 24: Microbial Diseases of the Respiratory System • The discussion of pertussis has been revised to better

describe some of the latest developments, especially the recent increase in cases.

• The discussion of tuberculosis has been updated and revised to include more on extensively resistant strains of the pathogen and some of the more recent testing methods.

PREFACE vii

• Discussion of influenza has been thoroughly revised and updated, especially the means by which mutants arise and the infectiveness of the avian flu virus.

Chapter 25: Microbial Diseases of the Digestive System • The discussion of traveler's diarrhea has been rewritten to

include the important pathogen enteroaggregative E. coli. • Recent therapeutic drugs for HBV have been included. • The discussion of noroviruses has been updated with special

attention to decontamination methods available to deal with outbreaks.

Chapter 26: Microbial Diseases of the Urinary and Reproductive Systems • The discussion of vaginal microbiota has been extensively

revised. • The introductory discussion of syphilis, especially relating

to recent genetic analysis on its probable origin in the New World, has been revised.

• The discussion of tes ting for syphilis has been revised. • The TORCH panel of tests is included.

Part Five Environmental and Applied Microbiology Chapter 27: Environmental Microbiology • The sulfur cycle figure (Figure 27.7)has been completely

redrawn. • The discussion of biodegradable plastics has been revised

and updated.

Chapter 28: Applied and Industrial Microbiology • The discussion of biofuels has been expanded .

Page 10: Microbiology 10th edition by Tortora

ACKNOWLEDG M ENTS

In preparing for this textbook, we have benefited from the guidance and advice of a large number of microbiology the country. The reviewers listed provided cnticism and valuable suggestions 011 various stages of the reVISIOn . We gratefully acknowledge our debt to these individuals.

TENTH EDITION REVIEWERS Cynthia Anderson

MI. SaIL A/lEOnia College Rod Anderson

Ollio Northern Ulliversity

Terry Austin Temple Col/ege

Joan Baird Rose State College

Archna Bhasin Valdosta SUIte University

Victoria Bingham Daytonll Beach College

Phyllis Braun Fairfield Ulliversity

Donald P. Breakwell Briglrllm Young University

Sandra Burnett Brig/ulIlI YOU/IS University

Susan Capasso Sf. ViI/ceil/'s College

Carol Castaneda indillll(l Ullil'ersity Nortlnvest

James K. Collins University of Arizon(l

Lee Couch University of New Mexico

Ellen C. Cover Lamar Ulliversity

Jean Cremins Middlesex COllllfllmity College

Melissa A. Deadmond Truckee Meadows Community College

Janet M. D«ker University of Arizona

Vivian Elder OZ(lrh Trchnical Commlmity College

viii

Christina Gan Higldine Comltllmity College

Pete Haddix AI/IIIITII University Montgomery

Rachel Hirst Massasoit Comm/mity College

Dawn Janich Comlll/mity College of Philadelphia

Judy Kaufman Monroe COlllllllmity College

MaIda Kocache George Maw// University

John M. Lammert GI/stavus Adolphus College

Paul A. leBlanc University of Alabama

Michael W. Lema Mid/aI/tis Tee/mimI College

John Lennox The Pelll/sylvm/ia Stale University

Shawn Lester Montgomery College

Leslie Lichtenstein Mlissasoit Commw/ity College

Eric Lifson Bllch COl/lilY Commllllity College

Suzanne Long Mal/roe Commul/ity College

William C. Matthai Tnrrallf COWIIY College Northeast

Philip Mixter WashillgtOl1 State University

Rita B. Moyes Texas A&M University

Ellyn R. Mulcahy Jolmsoll COlmty Comm/wity College

Tim R. Mullican Dakolll Wesleyml University

Richard L Myers Mi550uri Stale Ulliversity

Kabi Neupane Leeward Comm/mity College

Lourdes P. Norman Florit/II COlllllllwily College, Jacksonville

Page 11: Microbiology 10th edition by Tortora

Eric R. Paul Southwestern Oklahoma Stale University

Judy L. Penn Shordilre Community College

Jack Pennington St. Louis CommrmilY College, Forest Park

Indiren Pillay Culver-Stockton College

Ronny Priefer Niagara University

Todd P. Primm Sam HOI/stan State University

Mary L. Puglia Cerl/ral Arizona College

Amy J. Reese Cedar Crest College

Lois Sealy Valencia Community College West

Heather Smith Riverside City College

Kate Sutton Clark College

Paul H. Tomasek California State University, Northridge

David J. Wartell Harrisburg Area Community College

MJ Weintraub Raymond Walters College

Ruth Wrightsman Saddleback College

Anne Zayaitz Kutztown University

Michele Zwolinski Weber State University

We also thank the staff at Benjamin Cummings for their dedication to excellence. Leslie Berriman, our executlve editor, successfully kept us all focused on where we wanted this revision to go. Robin

Pille, project editor, masterfully managed the development of the book, keeping communicatlon Ilnes open and ensuring the highest quallty at every stage. Sally Peyrefitte's careful attentlon to continu-ity and detall in her copyedit of both text and art served to keep concepts and information clear throughout.

lanet Vail and Wendy Earl expertly guided the text through the production process. lisa Torri and the talented staff at Precision Graphics effectively managed the ambitious overhaul of our large and complex beautiful new art program. Jean Lake coordinated the many complex stages of the art development and rendering. The photo researcher, Maureen Spuhler, working closely with Senior Photo and Art Manager Travis Amos, made sure we had dear and striking images throughout the book. tani hasegawa created the ele-gant interior design, and Yvo Riezebos did a wonderful job with the cover. The skilled team at Progressive Information Technologies, led by Michelle lones, did an outstanding job moving this book quickly and beautifully through composition. Stacey Weinberger guided the book through the manufacturing process.

Katie Heimsoth impeccably handled the instructor supple-ments and also was the editor for the new edition of lohnson/Case Laboratory Experiments in Microbiology. Kelly Reed brought her creativity and teaching experience to bear on the development of the student supplements including the new Get Ready for Microbiology workbook. Lucinda Bingham managed the media program, working many miracles to produce the impressive array of resources on the website and Instructor's Resource DVD/CD-ROM including the new MicroFlix. Leslie Austin and lames Bruce managed the print and media supplements through the complex production stages.

Neena Ball, $enior Marketing Manager, and the entire Pearson Science sales force do a stellar job presenting this book to instruc-tors and students and ensuring its unwavering status as the best-selling microbiology textbook.

We would all like to acknowledge our spouses and families, who have provided invaluable support throughout the writing process.

FInally, we have an enduring appreciation for our students, whose comments and suggestions provide insight and remind us of their needs. This text is for them.

Gerard ]. Tortora Berdell R. Funke Christine L. Case

Page 12: Microbiology 10th edition by Tortora

A visual approach to teaching ...

,

Foundation Figures core concepts in microbiology

and give students the foundation they need to succeed in the course.

In its Tenth Edition, this best-selling textbook addresses the #1 challenge of the microbiology course: the wide variance in student levels, including the under-preparedness of many students. New and highly visual Foundation Figures get students to focus on and engage with core microbiology content.

• • • • • • •

The introductory text explains how the figure is foundational to other concepts students wilileam later.

.... shows typical structures thai may be fouod In boetena Eactl of the labeled structures WIll be dlSC\lSS :d indMdually In thiS cnapler As you will see in later chapters. some 0' these structures contnbute to bactenal wulence. play 8 role ,n bactenal I(\enIJliCII\JOf1. and are Ulrge\.S of 8I11imicrobial agenlS.

"

Key Concept

, ' iii .

, d 7

• • • •

__ 0"

"

• "

The Key Concept box presents the big picture, helping to ensure students understand the central concept presented by the figure.

Clear and consistent TEM/SEM/lM icons appear for all micrograph images, showing at a glance which type of microscope was used.

Page 13: Microbiology 10th edition by Tortora

... the foundations of microbiology

A small ver510fl of this OYeMeW figure WIll be included in olher figUres througnoul Ihe chapter to indicate the relationships 01 different reactions to the 0\IeftI11 procMses 01 respiration and fermentatIOn.

RUPIRATION FERMeNTATION o GI',coIt'!e pmdo or F 7 A TP I NACt 10 NADH

• oxIIlaIg gluu.11 10 pyruvic acid In responllion. the WwIc acid Ie conve.1ed 10 the :1\nI reactanlin Ih& Krebs cycle.

I-------/- 0 QI 7 , . ' ,

• • • • • • • • • • • • • • • • • • • while IJM"V 011 CO:z.

PO'" C8fTY eledrlll'l$

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Easy-Io-find blue step numbers guide the eye through complex processes, breaking them down into clear, manageable pieces that make concepts easier to teach and understand.

For a complete list of Foundation Figures, turn to page xxxi.

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Consistent use of symbols and colors enables students to progress from familiar parts of illustrated processes to unfamiliar ones with confidence. Molecules such as ATP are the same color and shape throughout the book.

Page 14: Microbiology 10th edition by Tortora

Frequent opportunities for students to check their understanding.

: --

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Check Your Understanding questions appear at the ends of major chapter sections, encouraging students to engage interactively with the text and self-assess their understanding of the section.

• n' cdium o f the same composition. onl y those colo o' ganisms capable of using phenol should grow. A rem

Draw It questions give students an opportunity to ;;;;;;;;;ct with figures and develop a deeper understanding of the content. Suggested answers to the Draw It questions are provided in the Answers section at the back of the book and replicate how an actual student's work might look.

• • • 3. DRAW IT I .. ,bel the parts of the compound light microscope in the figure below, and th.::n draw the path of light fTOllllhe illuminator to your eye.

o

LEARNING OBJECTIVES 6·' DISbngUlSh chemcally defll'll!d 80d complex media. 6·9 the use of e&Ch 01 the following: 8n&efOt)IC techniques .

'IrMg host cells. candle ,ars. selectNe aod dlffe.-en\l81 media. e"fJnchment medIUm .

• 6-10 Differentiate btosafety levels 1. 2. 3. aod 4 •

of this particular technique is that phenol is norma y • • to most bacteria. • • • • • •

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•• • • • • questions helps students determine which . ' .. ' • • • • objectives they have achieved and which require .r Could humans exist on chemiC1l,lJy defined media, at least under

.a" further study. laboratory conditions? 6-8 .r Could louis Pasteur, in the 1800s, have grown rabies viruses in

cell culture instead of in living animals? 6-9 .r What BSL is your laboratory? 6-10

xii

• •

Page 15: Microbiology 10th edition by Tortora

•• .and think like a clinician

Delayed Bloodstream Infection Following Catheterization

• Clinical Focus boxes contain Morbidity and Mortality Weekly Report data from the Centers for Disease Control and Prevention (CDC) modified into clinical prob-lem-solving scenarios and questions to help students develop their criti-cal thinking skills.

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Diseases in Focus ••• boxes organize comparative information about similar diseases and encourage students to think like a clinician.

Disease tables are organized •••• around similar symptoms, chunking information in a way that is relevant to clinical situations,

There are 23 Diseases in Focus boxes in this edition, 17 of them new, For a complete listing of topics. see page xxxi,

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Page 16: Microbiology 10th edition by Tortora

Unsurpassed online resources • • • iology

place will help students get ready for tests with its simple three-step approach:

Foundation Figure quizzes -;';;;;'e"," extra practice with core concepts.

Each gradable quiz includes multiple choice questions based on the figure as well as an essay question. o Take a Pre-Test and obtain a personalized study plan. e Learn and Practice with animations, tutoria ls, and

MP3 Tutor Sessions. .. Test Yourself with quizzes and a chapter post-test.

See the inside front cover of this book for details.

.. .. i tutor sessions i

MP3 Tutor Sessions are downloadable study guides for each chapter of the textbook,

allowing students to study on the go. They include mini· lectures about the toughest topics together with audio Quizzes so students can self-assess their understanding.

VARIOUS DIATOMS -

xiv

• • FOUNOATION FIGURE OVIZ

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25 Microbiology Videos .J feature live-action footage of microorganisms as they move and interact with their environments.

Microbe Reviews- . • • • gradable multiple-choice quizzes about micrographs from the textbook-give students extra practice with the organisms they are studying.

c c .... ,' . ..... c -C -C -C .. -

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Page 17: Microbiology 10th edition by Tortora

... for additional practice and assessment

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• MicroFlbc

MicroFlix are 3D, movie-quality animations with self-paced tutorials and grad able quizzes that help students master the three toughest topics in microbiology: metabolism, DNA replication, and immunology. Students can review the fundamentals by viewing the animations, completing the tutorial, printing a personal review sheet, and taking the quiz. Students also have access to BioFlix animation-stha! help them review relevant con-cepts from general biology_

• 115 multi-step Microbiology Animations explain and visually demonstrate core concepts, providing an additional chance for students to learn. They are accompanied by gradable quizzes. References to the Microbiology Animations appear throughout the chapters of the book.

A gradebook feature allows instructors to track and record their students' performance on website quizzes and tests.

Page 18: Microbiology 10th edition by Tortora

Get Ready for Microbiology quickly prepares your students for the microbiology course, helping them brush up on the skills they need to succeed.

CONTENTS Chapter 1 Study Skills Chapter 2 Math Basics Chapter 3 Tenninology Chapter 4 Chemistry Basics Chapter 5 Biology Basics Chapter 6 Cell Biology Chapter 7 Microbiology Basics

• • • • •

• •

Your Starting Point pre-tests students' grasp of chapter content before they start the chapter.

• • Microbiology Your Starting Point

Answer the (ollowing questions 10 assess yourchemistr')'

1. The most basic unit of a chemical substance is the _____ _

2. What the three SialH of maner? __________ _

3. An atom is made: orwh,1t three wb'llomic

• Chapters include textbook quality photographs and illustrations .

T I"" TO T il'

"""mont thfft drawonp Mlow. ones that you ar. crlls.

Engaging features like Time to Try •• provide a simple experiment or quick question that gives students a chance

xvi

to practice what they just learned.

The MyMicrobiologyPlace website includes additional practice and assessment material for Get Ready for Microbiology. including a Diagnostic Test to get students started.

Page 19: Microbiology 10th edition by Tortora

And for the microbiology lab • • •

,UOUTOIY UPU, .. INTS IN

MICROBIOLOGY TnIInsr.r of Bact.rta: AHpHc TKhnlqu. --_._--_. .::;;=,.=.=. • •

_ .. ... '", ... '--____ a ----_ .. , ---_ .. --

NOW IN FULL COLOR Laboratory Experiments in Microbiology, Ninth Edition by Ted R. Johnson I Christine L. Case

-1- ...... The new full-color design makes each lab exercise easier to navigate. The quick reference sections (Objectives, Materials, Cultures, Techniques Required) are clearly distinguished from the instructive and procedural sections (Background. Procedure) through colored headings .

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Rich, vibrant micrographs and other photographs now appear within the · .. •• .... •· relevant exercise, right next to the narrative. allowing students to easily view the images in the context of the exercise and to better inter· pret their results,

Clear, colorful, and realistically rendered stepobyostep diagrams walk students through each procedure, providing visual instructions in addition to narrative ones.

xvii

Page 20: Microbiology 10th edition by Tortora

The best support for instructors and students INSTRUCTOR SU PPLEM ENTS Instructors Resource OVDI CD-ROM (with TestGen ' Computerized Test Bank CD) 978-0-321-58190-7/0-321-58190-3

This media tool has been overhauled to make planning and presenting media easier. It includes: • All figures from the book with and without

labels in JPEG format • All figures from the book with the Label Edit feature

in format • Select "process" figures from the book with the Step Edit feature

in PowerPoint format • All tables from the book • Multimedia, including the Microbiology Animations, Microbiology

Videos, and MicroFlix Animations • PowerPoint lecture outlines, including figures and tables from the

book and links to multimedia • PRS-enabled Active lecture Clicker Questions • PRS-enabled Quiz Show Clicker Questions • The Instructor's Guide and Testbank as editable Microsofti' Word files • A CD-ROM of the Test Bank in format

Instructor's Visual Guide 978-0-321-58191-4 1 0-321-58191-1

Instructor's GuidelTest Bank by Christine L. Case 978-0-321-58187-7/0-321-58187-3

Transparency Acetates 978-0-321-58189-1/0-321-58189-X

CourseCompass™/ WebCT I Blackboard Pre-loaded book-specific content and test item files accompanying the text are available in several course management formats. Contact your local Benjamin Cummings sales representative for more information. To locate your rep, use the '"Find Your Rep" search feature at: www.pearsonhighered.com/educator

FORTHE LAB Laboratory Experiments in Microbiology, Ninth Edition by Ted R. Johnson and Christine L. Case 978-0-321-56028-5 / 0-321-56028-0 Containing 57 thoroughly class-tested exercises, the ninth edition features a new full-color design and art program. (See previous page for details.)

xviii

STUDENT SUPPLEMENTS MyMicrobiologyPlace Website www.microbiologyplace.com For a full description of the website, see the inside front cover of this book.

Get Ready for Microbiology by Lori K Garrett and Judy Meier Penn 978-0-321-59250-7/0-321-59250-6 This new brief primer saves classroom time and frustration by helping students quickly prepare for their microbiology course. (See page xvi for details).

Study Guide by Berdell R. Funke 978-0-321-58195-2 1 0-321-58195-4 Students can master key concepts and earn a better grade with the help of the clear writing and thought-provoking exercises found in this Study Guide. which includes concise explanations of key conceps. art labeling exercises. and a variety of self-test questions with answers.

Study Card 978-0-321-58196-9 1 0-321-58196-2 This six-panel. full-color study card provides students with a quick reference to the three most challenging topics in microbiology: metabolism. genetics, and immunology.

The Microbe Files: Cases in Microbiology for the Undergraduate by Marjorie K Cowan With answers: 978-0-8053-4927-61 0-8053-4927-8 Without answers: 978-0-8053-4928-3 I 0-8053-4928-6 The Microbe Files provides microbiology students with a fascinating series of short cases that help them apply what they have learned in the course.

Scientific American: Current Issues in Microbiology Vol.l: 978-0-8053-4623-7 / 0-8053-4623-6 Vo1.2: 978-0-3215-3816-1/ 0-3215-3816-1 Accessible, dynamic. and relevant articles from Scientific American magazine present key issues in microbiology, and end-of-article questions help students check their comprehension and make connections between science and society.

Techniques in Microbiology: A Student Handbook by John M. Lammert 978-0-13-224011-6/0-13-224011-4 This vivid, full-color handbook guides students in manipulations and preparations needed in the microbiology laboratory. The techniques are the ones that are used frequently for study-ing microbes in the laboratory and include those identified by the American Society for Microbiology (ASM) in its recommendations for the Microbiology Lab Core Curriculum.

\ .... "' .... , lI ... n""",,-

Page 21: Microbiology 10th edition by Tortora

BRIEF CONTENTS

PART ONE Fundamentals of Microbiology 1 The Microbial World and You I 2 Chemical Principles 26 3 Observing Microorganisms through

a Microscope 54 4 Functional Anatomy of Prokaryotic

and Eukaryotic Cells 76 5 Microbial Metabolism 113 6 Microbial Growth 156 7 The Control of Microbial Growth 184 8 Microbial Genetics 210 9 Biotechnology and Recombinant DNA 246

PART TWO A Survey of the Microbial World 10 Classification of Microorganisms 273 11 The Prokaryotes: Domains Bacteria

and Archaea 299 12 The Eukaryotes: Fungi, Algae, Protozoa,

and Helminths 329 13 Viruses, Viroids, and Prions 367

PART THREE Interaction between Microbe and Host 14 Principles of Disease and Epidemiology 399 15 Microbial Mechanisms of Pathogenicity 428 16 Innate Immunity: Nonspecific Defenses

of the Host 449 17 Adaptive Immunity: Specific Defenses

of the Host 476 18 Practical Applications of Immunology 500 19 Disorders Associated with the Immune

System 522 20 Antimicrobial Drugs 553

PART FOUR Microorganisms and Human Disease 21 Microbial Diseases of the Skin and Eyes 584 22 Microbial Diseases of the Nervous System 610 23 Microbial Diseases of the Cardiovascular

and Lymphatic Systems 637 24 Microbial Diseases of the Respiratory System 674 25 Microbial Diseases of the Digestive System 705 26 Microbial Diseases of the Urinary

and Reproductive Systems 743

PART FIVE Environmental and Applied Microbiology 27 Environmental Microbiology 766 28 Applied and Industrial Microbiology 793

Answers to Review and Multiple Choice Study Questions AN-I

Appendix A Metabolic Pathways AP-l

Appendix B Exponents, Exponential Notation, Logarithms, and Generation Time AP-7

Appendix C Methods for Taking Clinical Samples AP-S

Appendix D Pronunciation of Scientific Names AP-9

Appendix E Word Roots Used in Microbiology AP-13

Appendix F Classification of Bacteria According to Bergey's Manual AP-16

Glossary G-l

Credits C-l

Index 1-1

xix

Page 22: Microbiology 10th edition by Tortora

CONTENTS

PART ONE Fundamentals of Microbiology

1 The Microbial World and You 1 Microbes in OUf Lives 2

Naming and ClassifYing Microorganisms 2 Nomenclature 2 Types of Microorganisms 3 Classification of Microorganisms 6

A Brief History of Microbiology 6 The First Observations 7 The Debate Over Spontaneous Generation 8 The Golden Age of Microbiology 9 The Birth of Modern Chemotherapy: Dreams of a "Magic Bullet" 12 Modern Developments in Microbiology 13

Microbes and Human Welfare 16 Recycling Vital Elements 16 Sewage Treatment: Using Microbes to Recycle Water 17 Bioremediation: Using Microbes to Clean Up Pollutants 17 Insect Pest Control by Microorganisms 17 Modern Biotechnology and Recombinant DNA Technology 17

Microbes and Human Disease 18 Normal Microbiota 18 Biofilms ] 8 Infectious Diseases 19 Emerging Infectious Diseases ]9

Study Outline 22

Study Questions 23

2 Chemical Principles The Structure of Atoms 27

Chemical Elements 27 Electronic Configurations 28

"

26

How Atoms Form Molecules: Chemical Bonds 28

Ionic Bonds 28 Covalent Bonds 30 Hydrogen Bonds 31 Molecular Weight and Moles 32

Chemical Reactions 32 Energy in Chemical Reactions 32 Synthesis Reactions 32 Decomposition Reactions 32 Exchange Reactions 33 The Reversibility of Reactions 34

IMPORTANT BIOLOGICAL MOLECULES 34 Inorganic Compounds 34

Water 34 Acids, Bases, and Salts 35 Acid-Base Balance: The Concept of pH 35

Organic Compounds 37 Structure and Chemistry 37 Carbohydrates 39 Lipids 40 Proteins 42 Nucleic Acids 47 Adenosine Triphosphate (ATP) 47

Study Outline 49

Study Questions 51

3 Observing Microorganisms through a Microscope 54

Units of Measurement 55

Microscopy: The Instruments 55 Light Microscopy 56 Two-Photon Microscopy 62 Scanning Acoustic Microscopy 63 Electron Microscopy 63 Scanned-Probe Microscopy 65

Page 23: Microbiology 10th edition by Tortora

Preparation of Specimens for Light Microscopy 68 Preparing Smears for Staining 68 Sim ple Stains 69 Differential Stains 69 Special Stains 71

Study Outline 73

Study Questions 74

4 Functional Anatomy of Prokaryotic and Eukaryotic Cells 76

Comparing Prokaryotic and Eukaryotic Cells: An Overview 77

THE PROKARYOTIC CELL 77

The Size, Shape, and Arrangement of Bacterial Cells 77

Structures External to the Cell Wall 79 Glycocalyx 79 Flagella 81 Axial Filaments 82 Fimbriae and Pili 83

The Cell Wall 84 Composition and Characteristics 85 Cell Walls and The Gram Stain Mechanism 87 Atypical Cell Walls 87 Damage to the Cell Wall 88

Structures Internal to the Cell Wall 89 The Plasma (Cytoplasmic) Membrane 89 The Movement of Materials across Membranes 91 Cytoplasm 94 The Nucleoid 94 Ribosomes 95 Inclusions 95 Endospores 96

THE EUKARYOTIC CELL 98

Flagella and Cilia 98

The Cell Wall and Glycocalyx 98

The Plasma (Cytoplasmic) Membrane 100

Cytoplasm 100 Ribosomes 10]

Organelles l02 The Nucleus 102 Endoplasmic Reticulum 103

Golgi Complex 104 Lysosomes 104 Vacuoles 104 Mitochondria 104 Chloroplasts 105 Peroxisomes 105 Centrosome 105

The Evolution of Eukaryotes ] 06

Study Outline 108

Study Questions 110

CONTENTS xxi

5 Microbial Metabolism 113 Catabolic and Anabolic Reactions 114

Enzymes 115 Collision Theory 11 5 Enzymes and Chemical Reactions lIS Enzyme Specificity and Efficiency 116 Naming Enzymes 11 6 Enzyme Components I 16 The Mechanism of Enzymatic Action 117 Factors Influencing Enzymatic Activity 11 8 Feedback Inhibition 120 Ribozymes 121

Energy Production 12l Oxidation-Reduction Reactions 122 The Generation of ATP 122 Metabolic Pathways of Energy Production 123

Carbohydrate Metabolism 124 Glycolysis 124 Alternatives to Glycolysis 125 Cellular Respiration 127 Fermentation 132

Lipid and Protein Catabolism 136

Biochemical Tests and Bacterial Identification 137

Photosynthesis 140 The Light-Dependent Reactions: Photophosphorylation 140 The Light-Independent Reactions: The Calvin-Benson Cycle 140

Chaprer 5 continues

Page 24: Microbiology 10th edition by Tortora

xxii CONTENTS

Chapter 5, continued

A Summary of Energy Production Mechanisms 141

Metabolic Diversity among Organisms 142 Photoautotrophs 143 Photoheterotrophs 145 Chemoautotrophs 145 Chemoheterotrophs 145

Metabolic Pathways of Energy Use 146 Polysaccharide Biosynthesis 146 Lipid Biosynthesis 146 Amino Acid and Protein Biosynthesis 146 Purine and Pyrimidine Biosynthesis 147

The Integration of Metabolism 147

Study Outline 150

Study Questions 153

6 Microbial Growth 156 The Requirements for Growth 157

Physical Requirements 157 Chemical Requirements 160 Biofilms 162

C ulture Media 164 Chemically Defined Media 165 Complex Medi a 165 Anaerobic Growth Media and Methods 166 Special Culture Techniques 167 Selective and Differential Media 168 Enrichment Culture 169

Obtaining Pure C ultures 170

Preserving Bacterial C ultures 170

The Growth of Bacterial C ultures 17 1 Bacterial Division 171 Generation T ime 171 Logarithmic Representation of Bacterial Populations 171 Phases of Growth I 72 Direct Measurement of Microbial Growth 174 Estimating Bacterial Numbers by Indirect Methods 178

Study Outline 180

Study Questions 18 1

7 The Control of Microbial Growth The Terminology of Microbial Control 185

The Rate of Microbial Death 186

Actions of Microbial Control Agents 186 Alteration of Membrane Permeability 186 Damage to Proteins and Nucleic Acids 187

Physical Methods of Microbial Control 187 Heat 188 Filtration 191 Low Temperatures 19 1 High Pressure 192 Desiccation 192 Osmotic Pressure 192 Radiation 192

Chemical Methods of Microbial Control 195 Principles of Effective Disinfection 195 Evaluating a Disinfectant 195 Types of Disinfectants 195

Microbial Characteristics and Microbial Control 202

Stud y Outline 205

Study Questions 207

8 Microbial Genetics 210 Structure and Function of the Genetic Material 211

Genotype and Phenotype 21 I D NA and Chromosomes 21 1 The Flow of Genetic Information 2 12 D NA Replicat ion 2 12 RNA and Protein Synthesis 216

The Regulation of Bacterial Gene Expression 221 Repression and Induction 224 The Operon Model of Gene Expression 224 Positive Regulation 225

Mutation: C hange in the Genetic Material 226 Types of Mutations 227 Mutagens 229 The Frequency of Mutation 23 1 Identifying Mutants 231 Identifying Chemical Carcinogens 232

184

Page 25: Microbiology 10th edition by Tortora

Genetic Transfer and Recombination 233 Transformation in Bacteria 234 Conjugation in Bacteria 236 Transduction in Bacteria 237 Plasmids and Transposons 237

Genes and Evolution 241

Study Outline 242

Study Questions 244

9 Biotechnology and Recombinant DNA 246

Introduction to Biotechnology 247 Recombinant DNA Technology 247 An Overview of Recombinant DNA Procedures 247

Tools of Biotechnology 247 Selection 249 Mutation 249 Restriction Enzymes 249 Vectors 250 Polymerase Chain Reaction 25 I

Techniques of Genetic Modification 253 Inserting Foreign DNA into Cells 253 Obtaining DNA 254 Selecting a Clone 256 Making a Gene Product 257

Applications of rONA 258 Therapeutic Applications 258 The Human Genome Project 261 Scientific Applications 261 Agricultural Applications 264

Safety Issues and the Ethics of Using rONA 268

Study Outline 269

Study Questions 271

PART TWO A Survey of the Microbial World

1 0 Classification of Microorganisms 273

The Study of Phylogenetic Relationships 274 The Three Domains 274 A Phylogenetic Hierarchy 277

C lassification of Organisms 278 Scientific Nomenclature 278 The Taxonomic Hierarchy 279 Classification of Prokaryotes 279 Classification of Eukaryotes 281 Classification of Viruses 282

Methods of C lassifying and IdentifYing Microorganisms 282

Morphological Characteristics 284 Differential Staining 285 Biochemical Tests 285 Serology 287 Phage Typing 288 Fatty Acid Profiles 288 Flow Cytometry 288 DNA Base Composition 288 DNA Fingerprinting 289 The Polymerase Chain Reaction 290 Nucleic Acid Hybridization 291

CONTENTS xxiii

Putting Classification Methods Together 293

Study Outline 295

Study Questions 296

11 The Prokaryotes: Domains Bacteria and Archaea 299

The Prokaryotic Groups 300

DOMAIN BACTERIA 302

The Proteobacteria 302 The Alphaproteobacteria 303 The Betaproteobacteria 305 The Gammaproteobacteria 306 The Deltaproteobacteria 312 The Epsilonproteobacteria 312

The Nonproteobacteria Gram-Negative Bacteria 313 Cyanobacteria (The Oxygenic Photosynthetic Bacteria) 313 Purple and Green Photosynthetic Bacteria (The Anoxygenic Photosynthetic Bacteria) 315

The Gram-Positive Bacteria 3 15 Firmicutes (Low G + C Gram-Positive Bacteria) 316 Actinobacteria (High G + C Gram-Positive Bacteria) 320

Chapter II continues

Page 26: Microbiology 10th edition by Tortora

xxvi CONTENTS

Chapter 16, continued

Anti microbial Substances 463 The Complement System 463 Interferons 468 Iron-Binding Proteins 470 Antimicrobial Peptides 470

Study Outli ne 472

Study Questions 474

1 7 Adaptive Immunity: Specific Defenses of the Host 476

The Adaptive Immune System 477

Dual Nature of the Adaptive Immune System 477

Humoral Immunity 477 Cellular Immunity 477

Antigens and Antibodies 478 The Nature of Antigens 478 The Nature of Antibodies 479

B Cells and Humoral Immunity 482 Clon al Selection of Antibody-Producing Cells 482 The Diversity of Antibodies 484

Antigen-Antibody Binding and Its Results 484

T Cells and Cellular Immunity 486 Classes ofT Cells 487 T Helper Cells (CD4+ T Cells) 487 T Cytotox ic Cells (CD8+ T Cells) 488 T Regulatory Cells 489

Antigen-Presenting Cells (APCs) 489 Dendritic Cells 490 Macrophages 490

Extracellular Killing by the Immune System 491

Antibody-Dependent Cel l-MLxl iated Cytotoxicity 491

Cytokines: C hemical Messengers of Immune Cells 491

Immunological Memory 493

Types of Adaptive Immunity 494

Study Outline 497

Study Questions 498

1 8 Practical Applications of Immunology 500

Vaccines so I Principles and Effects of Vacci nation 501 Types of Vaccines and Their Characteristics 501 The Development of New Vaccines 504 Safety of Vaccines 506

Diagnostic Immunology 507 Im munologic-Based Diagnostic Tests 507 Monoclonal Antibodies 507 Precipitation Reactions 509 Agglutination Reactions 5 10 Neutralization Reactions 5 12 Complement-Fixation Reactions 512 Fluorescent-Antibody Techniques 5) 3 Enzyme-Linked Immunosorbent Assay (E LI SA) 514 Western Blotting (Immunoblotting) 516 The Future of Diagnostic Immunology 516

Study Outline 5 I 9

Stud y Questions 520

1 9 Disorders Associated with the Immune System 522

Hypersensitivity 523 Type I (Anaphylactic) Reactions 523 Type " (Cytotoxic) Reactions 526 Type III (Immune Complex) Reactions 528 Type IV (Delayed Cell-Mediated) Reactions 529

Autoimmune Diseases 532 Cytotoxic Autoimmune Reactions 532 Immune Complex Autoimmune Reactions 532 Cell-Mediated Autoimmune Reactions 532

Reactions Related to the Human Leukocyte Antigen (HLA) Complex 533

Reactions to Transplantation 534 Immunosuppression 536

The Immune System and Cancer 537 Immunotherapy for Cancer 538

Immunodeficiencies 538 Congenital Immunodeficiencies 538 Acquired Immunodeficiencies 538

Page 27: Microbiology 10th edition by Tortora

The Etiology of Infectious Diseases 404 Koch's Postulates 404 Exceptions to Koch's Postulates 404

Classifying Infectious Diseases 406 Occurrence of a Disease 406 Severity or Duration of a Disease 406 Extent of Host Involvement 407

Patterns of Disease 408 Predisposing Factors 408 Development of Disease 408

The Spread of Infection 409 Reservoirs of Infection 409 Transmission of Disease 409

Nosocomial (Hospital-Acquired) Infections 413 Microorganisms in the Hospital 414 Compromised Host 41 5 Chain of Transmission 415 Control of Nosocomial Infections 416

Emerging Infectious Diseases 416

Epidemiology 4 J 8 Descriptive Epidemiology 419 Analytical Epidemiology 41 9 Experimental Epidemiology 420 Case Reporting 420 The Centers for Disease Control and Prevention 420

Study Outline 423

Study Questions 425

1 5 Microbial Mechanisms of Pathogenicity 428

How Microorganisms Enter a Host 429 Portals of Entry 429 The Preferred Portal of Entry 429 Numbers of Invading Microbes 429 Adherence 431

How Bacterial Pathogens Penetrate Host Defenses 432 Capsules 432 Cell Wall Components 432 Enzymes 432 Antigenic Variation 433 Penetration into the Host Cell Cytoskeleton 433

CONTENTS xxv

How Bacterial Pathogens Damage Host Cells 434 Using the Host 's Nutrients: Siderophores 434 Direct Damage 434 The Production of Toxins 434 Plasmids, Lysogeny, and Pathogenicity 439

Pathogenic Properties of Viruses 441 Viral Mechanisms for Evading Host Defenses 441 Cytopathic Effects of Viruses 441

Pathogenic Properties of Fungi, Protozoa, Helminths, and Algae 442

Fungi 443 Protozoa 443 Helminths 444 Algae 444

Portals of Exit 444

Study Outline 445

Study Questions 447

16 Innate Immunity: Nonspecific Defenses of the Host 449

The Concept of Immunity 450

FIRST LINE OF DEFENSE: SKIN AND MUCOUS MEMBRANES 450

Physical Factors 45 J

Chemical Factors 453

Normal Microbiota and Innate Immunity 453

SECOND LINE OF DEFENSE 454 Formed Elements in Blood 454

The Lymphatic System 456

Phagocytes 457 Actions of Phagocytic Cells 457 The Mechanism of Phagocytosis 458 Microbial Evasion of Phagocytosis 459

Inflammation 460 Vasodilation and Increased Permeability of Blood Vessels 460 Phagocyte Migration and Phagocytosis 462 Tissue Repair 462

Fever 463

Chapter 16 continues

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Acquired Immunodefkiency Syndrome (AlDS) 539

The Origin of AIDS 540 HIV Infection 540 Diagnostic Methods 545 HIV Transmission 545 AIDS Worldwide 546 Preventing and Treating AIDS 547 The AIDS Epidemic and the Im portance of the Scientific Research 548

Study Outline 549

Study Questions 551

20 Antimicrobial Drugs

The History of Chemotherapy 554 Antibiotic Discovery Today 554

553

The Spectrum of Antimicrobial Activity 555

The Action of Antimicrobial Drugs 555 Inhibiting Cell Wall Synthesis 556 Inh ibiting Protein Synthesis 556 Injuring the Plasma Membrane 558 Inhibiting Nucleic Acid Synthesis 558 Inhibiting the Synthesis of Essential Metabolites 558

A Survey of Commonly Used Antimicrobial Drugs 559 Antibacterial Antibiotics: Inhibitors of Cell Wall Synthesis 559 Antimycobacterial Antibiotics 563 Inh ibitors of Protein Synthesis 563 Injury to the Plasma Membrane 566 Inh ibitors of Nucleic Acid (DNA/RN A) Synthesis 567 Competitive Inhibitors of the Synthesis of Essential Metabolites 567 Antifungal Drugs 567 Antiviral Drugs 569 Antiprotozoan and Antihelminthic Drugs 57 1

Tests to Guide Chemotherapy 572 The Diffusion Methods 572 Broth Dilution Tests 572

Resistance to Antimicrobial Drugs 573 Mechanisms of Resistance 574 Antibiot ic Misuse 575 Cost and Prevention of Resistance 576

Antibiotic Safety 576

CONTENTS xxvii

Effects of Combinations of Drugs 578

The Future of Chemotherapeutic Agents 578 Antimicrobial Peptides 578 Antisense Agents 579

Study Outline 580

Study Questions 582

PART FOUR Microorganisms and Human Disease

2 1 Microbial Diseases of the Skin and Eyes 584

Structure and Function of the Skin 585 Mucous Membranes 585

Normal Microbiota of the Skin 585

Microbial Diseases of the Skin 586 Bacterial Diseases of the Skin 586 Viral Diseases of the Skin 595 Fungal Diseases of the Skin and Nails 600 Parasitic Infestation of the Skin 602

Microbial Diseases of the Eye 603 Inflammation of the Eye Membranes: Conjunctivitis 603 Bacterial Diseases of the Eye 603 Other Infectious Diseases of the Eye 605

Study Outline 606

Study Questions 608

2 2 Microbial Diseases of the Nervous System 610

Structure and Function of the Nervous System 611

Bacterial Diseases of the Nervous System 611 Bacterial Meningitis 612 Tetanus 615 Botulism 616 Leprosy 619 Viral Diseases of the Nervous System 620

Poliomyelitis 620 Rabies 622 Arboviral Encephalitis 624

Chapter 22 continues

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xxviii CONTENTS

Chapter 22, continued

Fungal Disease of the Nervous System 626 Cryptococcus neoformans Meningit is (Cryptococcosis) 626

Protozoan Diseases of the Nervous System 627 African Trypanosomiasis 627 Amebic Meningoencephalitis 629

Nervous System Diseases Caused by Prions 629 Bovine Spongifonn Encephalopathy and Variant Creutzfeldt-Jakob Disease 63 1

Disease Caused by Unidentified Agents 633 Chronic Fatigue Syndrome 633

Study Outline 633

Study Questions 635

23 Microbial Diseases of the Cardiovascular and Lymphatic Systems 637

Structure and Function of the Cardiovascular and Lymphatic Systems 638

Bacterial Diseases of the Cardiovascular and Lymphatic Systems 638

Sepsis and Septic Shock 639 Bacterial Infections of the Heart 64 1 Rheumat ic Fever 64 1 Tularemia 64 2 Brucellosis (Undulant Fever) 643 Anthrax 645 Gangrene 646 Systemic Diseases Caused by Bites and Scratches 647 Vector-Transmitted Diseases 648

Viral Diseases of the Cardiovascular and Lymphatic Systems 655

Burkitt's Lymphoma 655 Infectious Mononucleosis 656 Other Diseases and Epstein-Barr Virus 657 Cytomegalovirus Infections 658 Chikungunya Fever 658 Classic Viral Hemorrhagic Fevers 658 Emerging Viral Hemorrhagic Fevers 659

Protozoan Diseases of the Cardiovascular and Lymphatic Systems 660

Chagas' Disease (American Trypanosomiasis) 661 Toxoplasmosis 661 Malaria 663 Leishmaniasis 665 Babesiosis 666

Helminthic Diseases of the Cardiovascular and Lymphatic Systems 666

Schistosomiasis 666 Swimmer's Itch 667

Study Outline 669

Study Questions 671

2 4 Microbial Diseases of the Respiratory System 674

Structure and Function of the Respi ratory System 675

Normal Microbiota of the Respiratory System 675

MICROBIAL DISEASES OF TH E UPPER RESPIRATORY SYSTEM 676

Bacterial Diseases of the Upper Respiratory System 677 Streptococcal Pharyngitis (Strep Throat) 677 Scarlet Fever 677 Diphtheria 677 Otitis Media 679

Viral Diseases of the Upper Respiratory System 679 The Common Cold 679

MICROBIAL DISEASES OF TH E LOWER RESPIRATORY SYSTEM 680 Bacterial Diseases of the Lower Respiratory System 680

Pertussis (Whooping Cough) 680 Tubercu losis 682 Bacterial Pneumonias 684 Meliodosis 690

Viral Diseases of the Lower Respiratory System 692 Viral Pneumonia 692 Respiratory Syncytial Virus (RSV) 692 Influenza (Flu) 692

Fungal Diseases of the Lower Respiratory System 695 Histoplasmosis 695 Coccidiomycosis 696

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Pneumocystis Pneumonia 697 Blastomycosis (North American Blastomycosis) 697 O ther Fungi Involved in Respiratory Disease 697

Study Outline 700

Study Questions 702

2 5 Microbial Diseases of the Digestive System 705

Structure and Function of the Digestive System 706

Normal Microbiota of the Digestive System 706

Bacterial Diseases of the Mouth 707 Dental Caries (Tooth Decay) 707 Periodontal Disease 709

Bacterial Diseases of the Lower Digestive System 710

Staphylococcal Food Poisoning (Staphylococcal Enterotoxicosis) 711 Sh igellosis (Bacillary Dysentery) 71 2 Salmonellosis (Salmonella Gastroenteritis) 71 2 Typhoid Fever 714 Cholera 716 Noncholera Vibrios 717 Escherichia coli Gastroenteritis 717 Ca mpylobacter Gastroenteritis 718 Helicobacter Peptic Ulcer Disease 718 Yersinia Gastroenteritis 720 Clostridium peifrjngens Gastroenteritis 720 Clostridium difficile- Associated Diarrhea 720 Bacillus cereus Gastroenteritis 720

Viral Diseases of the Digestive System 721 Mumps 721 Hepatitis 721 Viral Gast roenteritis 728

Fungal Diseases of the Digestive System 729 Ergot Poisoning 729 Aflatoxin Poisoning 730

Protozoan Diseases of the Digestive System 730

Giardiasis 730 Cryptosporidiosis 73 1 Cyclospora Diarrheal Infection 731 Amoebic Dysentery (Amoebiasis) 73 1

CONTENTS xxix

Helminthic Diseases of the Digestive System 732 Tapeworms 732 Hydatid Disease 733 Nematodes 734

Study Outline 738

Study Questions 741

2 6 Microbial Diseases of the Urinary and Reproductive Systems 743

Structure and Function of the Urinary System 744

Structure and Function of the Reproductive Systems 744

Normal Microbiota of the Urinary and Reproductive Systems 745

DISEASES OF THE URINARY SYSTEM 746 Bacterial Diseases of the Urinary System 746

Cystitis 746 Pyelonephritis 746 Leptospirosis 746

DISEASES OF THE REPROD UCTIVE SYSTEMS 747 Bacterial Diseases of the Reproductive Systems 747

Gonorrhea 747 Nongonococcal Urethritis (NGU) 750 Pelvic Inflammatory Disease (PID) 751 Syphilis 752 Lymphogranuloma Venereum 755 Chancroid (Soft Chancre) 756 Bacterial Vaginosis 756

Viral Diseases of the Reproductive System 757 Genital Herpes 757 Genital Warts 758 AIDS 758

Fungal Disease of the Reproductive Systems 758 Candidiasis 758

Protozoan Disease of the Reproductive Systems 760

Trichomoniasis 760 The TO RCH Panel of Tests 760

Study Outline 762

Study Questions 764