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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
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Executive Editor: Leslie Berriman Art Development Editor: Lisa Torri
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Cover Design: Riezebos Holzbaur Design Group Executive Market Development Manager: Josh Frost
Art Coordinator: Jean Lake

Cover Photo Credit: Science Photo Library/Eric Graves

Text art, and photo credits appear after the Glossary.

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
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permission(s) to use material from this work, please submit a written request to Pearson Ed ucation,
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. W here 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 2008044384

ISBN 10: 0-321-55007-2; ISBN 13 : 978-0-321-55007-1

(S tudent edition)
Benjamin Cummings
is an imprint of ISBN 10: 0-321-58202-0; ISBN 13: 978-0-321-58202-7
(Professional copy)

-
PEARSON
_.pearsonhighered.com
I 2 3 4 5 6 7 8 9 IO-DOW-12 II 10 09 08
Manufactured in the United States of America.
Executive Editor: Leslie Berriman Art Development Editor: Lisa Torri
Project Editor: Robin Pille Artists: Precision Graphics
Development Manager: Barbara Yien Copyeditor: Sally Peyrefitte
Assistant Editors: Katie Heimsoth and Kelly Reed Proofreader: Martha Ghent
Managing Editor: Wendy Earl Senior Art and Photo Manager: Travis Amos
Production Supervisor: Janet Vail Photo Researcher: Maureen Spuhler
Director, Media Development: Lauren Fogel Image Rights and Permissions Manager: Zina Arabia
Media Producer: Lucinda Bingham Senior Manufacturing Buyer: Stacey Weinberger
Interior Designer: tani hasegawa Senior Marketing Manager: Neena Bali
Cover Design: Riezebos Holzbaur Design Group Executive Market Development Manager: Josh Frost
Art Coordinator: Jean Lake

Cover Photo Credit: Science Photo Library/Eric Graves

Text art, and photo credits appear after the Glossary.

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, mec hanical, photocopying, recording, or likewise. To obtain
permission(s) to use material from this work, please submit a written request to Pearson Ed ucation,
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. W here 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 2008044384

ISBN 10: 0-321-55007-2; ISBN 13 : 978-0-321-55007-1

(S tudent edition)
Benjamin Cummings
is an imprint of ISBN 10: 0-321-58202-0; ISBN 13: 978-0-321-58202-7
(Professional copy)

-
PEARSON
_.pearsonhighered.com
I 2 3 4 5 6 7 8 9 IO-DOW-12 II 10 09 08
Manufactured in the United States of America.
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 Association (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 Institute 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 student s. 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
 PREFACE

Since the publication of the fi rst edition nearly 30 years ago, edition meets all students at their respective levels of skill and
well over one million students have used Microbiology: An understanding.
Introduction at colleges and universities around the world, mak- The highlights of the tenth edition can be seen in the new
ing it the leading textbook for non-majors microbiology. The Foundation Figures, the new features that help studen ts check
tenth edition continues to be a comprehensive beginning text, their understanding as they progress through each chapter, and
assum ing no previous study of biology or chemistry. The text is the improved boxes that prepare students to start thinking like a
appropriate for students in a wide variety of programs, including cli nician. Content and currency have also been substantially
the allied health sciences, biological science, environmental sci- updated.
ences, animal science, forestry, agric ulture, home economics, and
the liberal arts. Foundation Figures
In order to help students focus on and master the core concepts
HALLMARKS OF MICROBIOLOGY: of microbiology, the authors have integrated text and visuals in to
twenty specially designed Foundation Figures. These Foundation
AN INTRODUCTION Figures include both a key concept statement that ensures stu-
We have retained in this new edition features that made the dents understand the central concept of the figure and an expla-
previous editions so popular: nation of how each figure is foundational to further learning in
the course. In addition, throughout the entire book the illustra-
• An appropriate balance between microbiological funda- tion program has been dramatically revised and updated with
mentals and applications, and between medical applica- new art styles and a brighter color palette that has more con-
tions and other applied areas of microbiology. As in trasting colors and more dimensionality.
previous editions, basic microbiological principles are given
greater emphasis than are applications, and health-related
Features that Help Students Check Their
applications are featured.
Understanding
• Straightforward presentation of complex topics. Each sec-
tion of the text has been written with the student in mind. New Check Your Understanding questions encourage students to
Our book is known for its clear explanations and consistent engage interactively with the material and self-assess their unde r-
pedagogy. standing of the Learning Objectives as they progress through
each chapter. New Draw It questions are now included in the
• Clear, accurate, and pedagogically effective illustrations and
end-of-chapter Review Questions, asking students to sketch a
photos. Step-by-step diagrams that closely coordinate with
rough diagram or fill in missing parts of a figure or graph. The
narrative descriptions aid student comprehension of concepts.
popular Figure Legend Questions have been retained and
Clear and accurate renderings of processes and structures focus
im proved.
students on what they need to learn. The quantity and quality
of electron and light micrographs is unmatched in the market.
Support for Students to Start Thinking like
• Flexible organization. We have organized the book in what
we thin k is a useful fas hion while recognizing that the mate- a Clinician
rial might be effectively p resented in a number of other Revised and redesigned Appl ications of Microbiology boxes
sequences. For instructors who wish to use a different order, describe current and practical uses of microbiology. New and
we have made each cha pter as independent as possible and updated Clinical Focus boxes contain data from the Morbidityalld
have included numerous cross- references. The Instructor's Mortality Weekly Report modified into clinical problem-solving
Guide, written by Christine Case, provides detailed guide- questions that help students develop their critical thinking skills
lines for organizing the material in several other ways. 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
NEW TO THE TENTH EDITION interesting format that provides a helpful study tool for students.
Please see pages x- xviii for a visual introduction to the new
tenth edition . Updates in Content and Currency
The changes in this edition address instructors' biggest chal-
Antimicrobial resistance, biofilms, bioterrorism, and evolution
lenge in the introductory microbiology course: the wide range of
receive special attention and increased emphasis. The immunity
student levels, including student under-preparedness. The tenth
chapters-Chapters 16 and 17- have received a careful and sig-
 PREFACE v

nificant update for increased currency, clarity, and accuracy, with - • Discussion of anaerobic growth media and methods has
out an increase in the level of detail. Taxonomy, nomenclature, been updated.
and d isease incidence data are current through August 2008. • A discussion of Biosafety Levels has been added, including a
figure illustrating Biosafety Level 4.
Cutting-Edge Media Supplements • A new figure showing differential medium is included.
• Figure 6.15 is now a Foundation Figure.
Turn to the inside front cover of this book for an overview of the
• A new Clinical Focus box illustrates the role of biofilms in
new MyMicrobiologyPlace website with its simple th ree-step
causing nosocomial infections.
approach to learning. Pages xiv-xv provide more details about
the exciting new student and instructor media, including the Chapter 7: The Control of Microbial Growth
MP3 Tutor Sessions, the 3D MicroFlix animations, and the • The definition of sterilization has been updated and qualified
Instructor's Resource DVD/CD -ROM. in consideration of the existence of prions.
• Figure 7.1 is now a Foundation Figure.
• Discussion of ultra-h igh temperature (UHT ) has been
CHAPTER-BY-CHAPTER REVISIONS clarified.
Every chapter in this edition has been thoroughly revised, and • New products and newly approved uses are included.
data in the text, tables, and figures have been updated through • A new Clinical Focus box illustrates the relationship between
August 2008 where possible. The main changes for each chapter improper disinfection and nosocomial infection .
are summarized below. Chapter 8: Microbial Genetics
• Figure 8.2 is now a Foundation Figure.
Part One • Discussion of genetic combination by crossing over has been
Fundamentals of Microbiology revised for clarity.
• snRNPs are defined.
Chapter 1: The Microbial Wortd and You • Inducible and repressible operons are explained and com-
• The table "Making Scientific Names Familiar" has been pared in separate figures .
moved to this chapter from Chapter 10.
• Biofilms are introduced . Chapter 9: Biotechnology and Recombinant DNA
• Discussion of emerging infectious diseases has been updated, • Figure 9.1 is now a Foundation Figure.
including a section on antibiotic-resistant bacteria. • Gene silencing, reve rse genetics, and real-time PCR are
discussed.
Chapter 2: Chemical Principles • A new Clincal Focus box describes using reverse-transc ription
• Definitions have been expanded, including definitions of cis PCR to track a norovirus outbreak.
and trans fatty acids.
• Figure 2.16 is now a Foundation Figure.
Part Two
Chapter 3: Observing Microorganisms through a Microscope A Survey of the Microbial World
• Two-photon microscopy is included.
• Several new photos illustrate microscopic images. Chapter 10: Classification of Microorganisms
• Figure 3.2 is now a Foundation Figure. • Figure 10.1 is now a Foundation Figure.
• Photos of fossil and living stromatolites are included.
Chapter 4: Functional Anatomy of Prokaryotic • The use of transport media is explained.
and Eukaryotic Cells
• Figure 4.6 is now a Foundation Figure. Chapter 11 : The Prokaryotes: Domains Bacteria and Archaea
• The discussion of flagella, fimbriae, and pili has been revised, • Several new bacterial groups are discussed: Pelagibacter,
as has the discussion of the lipopolysaccharide. Acinetobacter baumallii, Planctomycetes, Gemmata
• The discussion of facilitated diffusion has been revised, and a obscuriglobus.
new figure compares types of diffusion across membranes, • Discussion of the theoretical minimal size of a bacterium
including aquaporins. and its genetic requirements has been revised.

Chapter 5: Microbial Metabolism Chapter 12: The Eukaryotes: Fungi, Algae, Protozoa,
• The section on biochemical tests has been expanded. and Helminths
• New renditions of enzymes are more realistic. • Examples of new uses of fungi as pesticides are listed.
• Figure 5.11 is now a Foundation Figure. • Discussion of the oomycotes is expanded to include
• A new Clinical Focus box illustrates the use of biochemical introduction of Phytophthora into the United States.
tests to identify slow-growing mycobacteria . The oomycote life cycle is illustrated in a new figure .
• Heartworm is included.
Chapter 6: Microbial Growth • A new Clinical Focus box h ighlights cryptosporidial
• The discussion ofbiofitms previously appearing in Chapter 27 diarrhea, the most common pathogen associated with
has been moved to this chapter and been significantly updated . .
sWlmmmg.
and expanded.
vi PREFACE

Chapter 13: Viruses, Viroids, and Prions • The discussions ofT cells, dendritic cells, and cytokines have
• The chapter begins with the use of retroviridae to genetically been completely revised.
modify cells. • A new Applications of Microbiology box describes the possi-
• Figure 13.15 is now a Foundation Figure. ble use of I L-12 to treat psoriasis.
• Bee colony collapse is mentioned.
Chapter 18: Practical Applications of Immunology
• The Clinical Focus box on the evolution and occurrence of
• Figure 18.2 is now a Foundation Figure.
avian flu has been updated.
• Discussions of DNA vaccines and adjuvants have been
updated and revised.
Part Three • The tables of vaccine schedules have been updated .
Interaction Between Microbe and Host • A new Clinical Focus box illustrates the success of vac-
cination in eliminating measles in the U.S. and highlights the
Chapter 14: Principles of Disease and Epidemiology
importance of measles as a cause of death in developing
• Figure 14.3 is now a Foundation Figure.
countries.
• Statistics on notifiable infectious diseases have been updated.
• A new Clinical Focus box illustrates the emergence of Chapter 19: Disorders Associated with the Immune System
hospital-acquired and community-acquired MRSA. • Coverage of blood groups includes a discussion of the rela-
tionship between certain blood groups and their relative
Chapter 15: Microbial Mechanisms of Pathogenicity
resistance or susceptibility to certain diseases.
• Discussion of A-B toxins has been expanded and clarified.
• A discussion of the autoimmune disease psoriasis and its
• Figure 15.5, action of an exotoxin, has been revised and
associated arthritis has been introduced, along with the
expanded.
current treatments with monoclonal antibodies.
• Figures 15.4 and 15.9 are now Foundation Figures.
• The discussion of stem cells has been updated, and a new
• A new Clinical Focus box illustrates role of biofilms and
figure (Figure 19.10 ) the derivation of stem cells and stem
endotoxins in postoperative infections.
cell lines.
Chapter 16: Innate Immunity: Nonspecific Defenses • The discussion of HIV and AIDS has been revised and
of the Host updated. Especially important is the complete revision of
• Treatment of several topics has been expanded and/or Figure 19.13, which shows the sequence of attachment,
reorganized and clarified: physical and chemical factors in fusion, and entry of the virus into the target CD4+ T cell.
the first line of defense; fo rmed elements in blood; the lym- • Figure 19.16 is now a Foundation Figure.
phatic system (including additional illustrations); adherence,
Chapter 20: Antimicrobial Drugs
acute-phase proteins, complement, iron-binding proteins,
• Figure 20.2 is now a Foundation Figure.
and antimicrobial peptides.
• The historical importance of the sulfa drugs is given more
• The role of biofilms in evading phagocytosis is included.
prommence.
• Figures 16.7 and Figure 16.9 are now Foundation Figures.
• The current methods used for the discovery of new
• The Applications of Microbiology box on serum collection has
antibiotics are discussed, including rapid throughput
been revised to include testing for complement to monitor
methods.
immune complex diseases in patients.
• The discussion of antibiotics has been updated to admit
Chapter 17: Adaptive Immunity: Specific Defenses new antibiotics. The discussion of antivirals for the treatment
of the Host of HIV / AIDS has been especially updated and revised to
• A new photo depicts actual antibody morphology shown by include the latest developments in this constantly changing
atomic force microscopy. area.
• Several important figures have been extensively revised for • The discussion of resistance to antibiotics has been com-
accuracy and clarity: pletely revised and expanded, and a new Foundation Figure
• Figure 17.5. Clonal selection and differentiation of (Figure 20.20) illustrates the most important target areas for
B cells resistance.
• Figure 17.10. Activation ofCD4 + T cells • The concluding discussion on the future of antibiotic devel-
• Figure 17.11. Killing of virus-infected target cell by opment and the prospect for unconventional antibiotics has
cytotoxic T lymphocyte been completely revised and updated.
• Figure 17.19. The dual nature of the immune system
(now a Foundation Figure} Part Four
• A new photo and illustration (Figure 17.9) show M cells
Microorganisms and Human Disease
found within Peyer's patches.
• Discussion of the major histocompatibility complex (M HC) Chapter 21 : Microbial Diseases of the Skin and Eyes
has been revised and improved. • The discussion of Staphylococcus aurerlS has been completely
• Nomenclature conventions have been updated for T cells rewritten to emphasize the importance of MRSA.
(for example, T helper cell, CD4 + T cell).
 PREFACE vii

• The discussion of impetigo and scalded skin syndrome has • Discussion of influenza has been thoroughly revised and
been revised, and discussion of a new disease, Buruli ulcer, updated, especially the means by which mutants arise and
has been added. the infectiveness of the avian flu virus.
• Some of the newer treatments for acne now have an expanded
Chapter 25: Microbial Diseases of the Digestive System
discussion.
• The discussion of traveler's diarrhea has been rewritten to
Chapter 22: Microbial Diseases of the Nervous System include the important pathogen enteroaggregative E. coli.
• A new figure (Figure 22.4) illustrates a spinal ta p. • Recent therapeutic drugs for HBV have been included.
• The discussion of cryptococcosis has been revised to include • The discussion of noroviruses has been updated with special
a newer pathogen. attention to decontamination methods available to deal with
• A brief description of prions has been included to outbreaks.
supplement that given in Chapter 13.
Chapter 26: Microbial Diseases of the Urinary
• The discussion of chronic disease syndrome has been
and Reproductive Systems
completely revised and now includes the CDC's diagnos-
• The discussion of vaginal microbiota has been extensively
tic definition and the alternative name of myalgic
revised.
en cep halo myel it is.
• The introductory discussion of syphilis, especially relating
Chapter 23: Microbial Diseases of the Cardiovascular to recent genetic analysis on its probable origin in the New
and lymphatic Systems World, has been revised.
• The definitions of the similar terms septicemia and sepsis • The discussion of testing for syphilis has been revised.
have been revised. • The TORCH panel of tests is included.
• Discussions of brucellosis and rat-bite fever have been
completely rewritten. Part Five
• The discussion of ehrlichiosis has been revised to include the
Environmental and Applied Microbiology
new terminology of anaplasmosis.
• Discussion of the disease chikungunya fever has been added Chapter 27: Environmental Microbiology
because of its current spread into temperate climates. • The sulfur cycle figure (Figure 27.7)has been completely
• The discussion of malaria has been revised completely to redrawn.
better differentiate between prophylaxis and the rapy. • The discussion of biodegradable plastics has been revised
and updated.
Chapter 24: Microbial Diseases of the Respiratory System
• The discussion of pertussis has been revised to better Chapter 28: Applied and Industrial Microbiology
describe some of the latest developments, especially the • The discussion of biofuels has been expanded .
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.
 ACKNOWLEDG M ENTS

In preparing for this textbook, we have benefited from the guidance Christina Gan
Higldine Comltllmity College
and advice of a large number of microbiology the
country. The reviewers listed provided cnticism and Pete Haddix
valuable suggestions 011 various stages of the reV ISIO n . We gratefully AI/IIIITII University Montgomery
acknowledge our debt to these individuals. Rachel Hirst
Massasoit Co mm/mity College
Dawn Janich
TENTH EDITION REVIEWERS Com lll/mity College of Philadelphia
Cynthia Anderson Judy Kaufman
MI. SaILA/lEOnia College Monroe COlllllllmity College
Rod Anderson MaIda Kocache
Ollio Northern Ulliversity George Maw// University
Terry Austin John M. Lammert
Temple Col/ege GI/stavus Adolphus College
Joan Baird Paul A. leBlanc
Rose State College University of Alabama
Archna Bhasin Michael W. Lema
Valdosta SUIte University Mid/aI/tis Tee/mimI College
Victoria Bingham John Lennox
Daytonll Beach College The Pelll/sylvm/ia Stale University
Phyllis Braun Shawn Lester
Fairfield Ulliversity Montgomery College
Donald P. Breakwell Leslie Lichtenstein
Briglrllm Young University Mlissasoit Commw/ity College
Sandra Burnett Eric Lifson
Brig/ulIlI YOU/IS University Bllch COl/lilY Commllllity College
Susan Capasso Suzanne Long
Sf. ViI/ceil/'s College Mal/roe Commul/ity College
Carol Castaneda William C. Matthai
indillll(l Ullil'ersity Nortlnvest Tnrrallf COWIIY College Northeast
James K. Collins Philip Mixter
University of Arizon(l WashillgtOl1 State University
Lee Couch Rita B. Moyes
University of New Mexico Texas A&M University
Ellen C. Cover Ellyn R. Mulcahy
Lamar Ulliversity Jolmsoll COl m ty Comm/wity College
Jean Cremins Tim R. Mullican
Middlesex COllllfllmity College Dakolll Wesleyml University
Melissa A. Deadmond Richard L Myers
Truckee Meadows Community College Mi550uri Stale Ulliversity
Janet M. D«ker Kabi Neupane
University of Arizona Leeward Comm/mity College
Vivian Elder Lourdes P. Norman
OZ(lrh Trchnical Commlmity College Florit/II COlllllllwily College, Jacksonville

viii
Eric R. Paul Pille, project editor, masterfully managed the development of the
Southwestern Oklahoma Stale University book, keeping communicatlon Ilnes open and ensuring the highest
Judy L. Penn quallty at every stage. Sally Peyrefitte's careful attentlon to continu-
Shordilre Community College ity and detall in her copyedit of both text and art served to keep
concepts and information clear throughout.
Jack Pennington
lanet Vail and Wendy Earl expertly guided the text through the
St. Louis CommrmilY College, Forest Park
production process. lisa Torri and the talented staff at Precision
Indiren Pillay Graphics effectively managed the ambitious overhaul of our large
Culver-Stockton College and complex beautiful new art program. Jean Lake coordinated the
Ronny Priefer many complex stages of the art development and rendering. The
Niagara University photo researcher, Maureen Spuhler, wo rking closely with Senior
Todd P. Primm Photo and Art Manager Travis Amos, made sure we had dear and
Sam HOI/stan State University striking images throughout the book. tani hasegawa created the ele-
gant interior design, and Yvo Riezebos did a wonderful job with the
Mary L. Puglia
cover. The skilled team at Progressive Information Technologies,
Cerl/ral Arizona College
led by Michelle lones, did an outstanding job moving this book
Amy J. Reese quickly and beautifully through composition. Stacey Weinberger
Cedar Crest College guided the book through the manufacturing process.
Lois Sealy Ka tie Heimsoth impeccably handled the instructor supple-
Valencia Community College West ments and also was the editor for the new edition of lohnson/Case
Heather Smith Laboratory Experiments in Microbiology. Kelly Reed brought her
Riverside City College creativity and teaching experience to bear on the development of
the student supplements including the new Get Ready for
Kate Sutton
Microbiology workbook. Lucinda Bingham managed the media
Clark College
program, working many miracles to produce the impressive array
Paul H. Tomasek of resources on the website and Instructor's Resource DVD/CD-
California State University, Northridge ROM including the new MicroFlix. Leslie Austin and lames Bruce
David J. Wartell managed the print and media supplements through the complex
Harrisburg Area Community College production stages.
MJ Weintraub Neena Ball, $enior Marketing Manager, and the entire Pearson
Raymond Walters College Science sales force do a stellar job presenting this book to instruc-
tors and students and ensuring its unwavering status as the best-
Ru th Wrightsman
selling microbiology textbook.
Saddleback College
We would all like to acknowledge our spouses and families,
Anne Zayaitz who have provided invaluable support throughout the writing
Kutztown University process.
Michele Zwolinski FInally, we have an enduring appreciation for our students,
Weber State University whose comments and suggestions provide insight and remi nd us of
their needs. This text is for them.
We also thank the staff at Benjamin Cummings for their dedication
to excellence. Leslie Berriman, our executlve editor, successfully Gera rd ]. Tortora Berdell R. Fun ke Christine L. Case
kept us all focused on where we wanted this revision to go. Robin
A visual approach to teaching ...
Foundation Figures In its Tenth Edition, this best-selling textbook
core concepts in microbiology addresses the #1 challenge of the microbiology
and give students the foundation they
course: the wide variance in student levels, including
need to succeed in the course.
the under-preparedness of many students. New and
•
• highly visual Foundation Figures get students to

• The introductory text explains how the focus on and engage with core microbiology content.
•• figure is foundational to other concepts
students wilileam later.
•
•
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.... 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.

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,
... the foundations of microbiology

A small ver510fl of this OYeMeW figure WIll be included in olher figUres througnoul Ihe
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pieces that make concepts easier to teach and understand. unfamiliar ones with confidence. Molecules such as ATP
are the same color and shape throughout the book.

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

Frequent opportunities for students
to check their understanding. • •
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.

•
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•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
illuminat or to your eye.

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Check Your Understanding

questions appear at the ends of major chapter sections, LEARNING OBJECTIVES
encouraging students to engage interactively with the DISbngUlSh chemcally defll'll!d 80d complex media.
6·'
text and self-assess their understanding of the section.
6·9 the use of e&Ch 01 the following: 8n&efOt)IC techniques.
•
n' cdium o f the same composition. onl y those colo
'IrMg host cells. candle ,ars. selectNe aod dlffe.-en\l81 media.
e"fJnchment medIUm .
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.r Could humans exist on chemiC1l,lJy defin ed media, at least under objectives they have achieved and which require
laboratory conditions? 6-8 .a" further study.

.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
•• .and think like a clinician
• Clinical Focus boxes contain
Morbidity and Mortality Weekly
Report data from the Centers for
Disease Control and Prevention
Delayed Bloodstream Infection (CDC) modified into clinical prob-

-- . ..
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xiii
Unsurpassed online resources • • •
iology Foundation Figure quizzes
-;';;;;'e"," extra practice with core concepts.

will help students get ready for tests with its

simple three-step approach:
place 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. •
FOUNOATION FIGURE OVIZ

See the inside front cover of this book for details.

__._-
- .......
' ..-
Ii ••_ _

....
.. ..
itutor sessions i
o o
_........._.--.._.
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
7 I _ _.......- ..
Quizzes so students can self-assess their understanding.

.-.
{ I. , "_,,,_ - _. _ , , _ _ • "I •••

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-VARIOUS DIATOMS

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

Microbe Reviews- . • • •
gradable multiple-choice -_ I 100_ I

quizzes about micrographs

from the textbook-give c
--
c c .... ,' ......

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students extra practice C
C
with the organisms they C
are studying.

xiv
... for additional
practice and assessment
MicroFlbc
fp

_
"" . . . . . ""'" _ NADII
_ _ ,e
...... d
'00 . . . . . .

. _
• MicroFlix are 3D,
movie-quality animations with
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n.._ .. quizzes that help students master

-.,' ,-_....'"
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.... _ _ 0 1 _
the three toughest topics in
,
microbiology: metabolism, DNA
'" replication, and immunology.

-' ................ _.
fPL _ _ _ '1$ 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.
 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 Your Starting Point
Chapter 2 Math Basics pre-tests students' grasp
of chapter content before
Chapter 3 Tenninology they start the chapter.
Chapter 4 Chemistry Basics •
Chapter 5 Biology Basics
Chapter 6 Cell Biology
• Microbiology
Chapter 7 Microbiology Basics Your Starting Point
Answer the (ollowing questions 10 assess yourchemistr')'

1. The most basic unit of a chemical substance is t he _ _ _ _ __

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
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.
xvi
And for the
microbiology lab • • • ,UOUTOIY UPU, .. INTS IN

MICROBIOLOGY
TnIInsr.r of Bact.rta: NOW IN FULL COLOR
--_._--_.
AHpHc TKhnlqu.
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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PROCEDURE "", ......., _____ ..
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. ____ 0 ,", .. ___. . . . . . __
Rich, vibrant micrographs and other
photographs now appear within the · .. •• .... •·
relevant exercise, right next to the narrative. Clear, colorful, and realistically rendered
allowing students to easily view the images in stepobyostep diagrams walk students through
the context of the exercise and to better inter· each procedure, providing visual instructions in
pret their results, addition to narrative ones.

xvii
The best support for instructors and students
INSTRUCTOR SU PPLEM ENTS STUDENT SUPPLEMENTS
Instructors Resource OVDI MyMicrobiologyPlace Website
CD-ROM (with TestGen ' www.microbiologyplace.com
Computerized Test Bank CD) For a full description of the website, see the inside front cover of this book.
978-0-321-58190-7/0-321-58190-3
Get Ready for Microbiology
This media tool has been overhauled to by Lori K Garrett and Judy Meier Penn
make planning and presenting media easier. 978-0-321-59250-7/0-321-59250-6
It includes: This new brief primer saves classroom time
• All figures from the book with and without and frustration by helping students quickly
labels in JPEG format prepare for their microbiology course.
• All figures from the book with the Label Edit feature (See page xvi for details).
in format
Study Guide by Berdell R. Funke
• Select "process" figures from the book with the Step Edit feature
978-0-321-58195-2 1 0-321-58195-4
in PowerPoint format
Students can master key concepts and earn a better grade with the
• All tables from the book
help of the clear writing and thought-provoking exercises found in this
• Multimedia, including the Microbiology Animations, Microbiology
Study Guide. which includes concise explanations of key conceps. art
Videos, and MicroFlix Animations
labeling exercises. and a variety of self-test questions with answers.
• PowerPoint lecture outlines, including figures and tables from the
book and links to multimedia Study Card
• PRS-enabled Active lecture Clicker Questions 978-0-321-58196-9 1 0-321-58196-2
• PRS-enabled Quiz Show Clicker Questions This six-panel. full-color study card provides students with a quick
• The Instructor's Guide and Testbank as editable Microsofti' Word files reference to the three most challenging topics in microbiology:
• A CD-ROM of the Test Bank in format metabolism. genetics, and immunology.

Instructor's Visual Guide The Microbe Files: Cases in Microbiology

978-0-321-58191-4 1 0-321-58191-1 for the Undergraduate by Marjorie K Cowan
With answers: 978-0-8053-4927-61 0-8053-4927-8
Instructor's GuidelTest Bank by Christine L. Case
Without answers: 978-0-8053-4928-3 I 0-8053-4928-6
978-0-321-58187-7/0-321-58187-3
The Microbe Files provides microbiology students with a fascinating
Transparency Acetates series of short cases that help them apply what they have learned
978-0-321-58189-1/0-321-58189-X in the course.

CourseCompass™/ WebCT I Blackboard Scientific American: Current Issues

Pre-loaded book-specific content and test item files accompanying in Microbiology
the text are available in several course management formats. Vol.l: 978-0-8053-4623-7 / 0-8053-4623-6
Contact your local Benjamin Cummings sales representative for Vo1.2: 978-0-3215-3816- 1/ 0-3215-3816-1
more information. To locate your rep, use the '"Find Your Rep" Accessible, dynamic. and relevant articles from Scientific American
search feature at: www.pearsonhighered.com/educator magazine present key issues in microbiology, and end-of-article
questions help students check their comprehension and make
connections between science and society.

FORTHE LAB
Laboratory Experiments in Techniques in Microbiology:
Microbiology, Ninth Edition A Student Handbook
by Ted R. Johnson and Christine L. Case by John M. Lammert
978-0-321-56028-5 / 0-321-56028-0 978-0-13-224011-6/0-13-224011-4
Containing 57 thoroughly class-tested exercises, This vivid, full-color handbook guides students
the ninth edition features a new full-color design in manipulations and preparations needed in
and art program. (See previous page for details.) the microbiology laboratory. The techniques \ .... "' ...., lI... n""",,-
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.
xviii
 BRIEF CONTENTS

PART ONE Fundamentals of Microbiology PART FOUR Microorganisms

and Human Disease
1 The Microbial World and You I
2 Chemical Principles 26 21 Microbial Diseases of the Skin and Eyes 584
3 Observing Microorganisms through 22 Microbial Diseases of the Nervous System 610
a Microscope 54 23 Microbial Diseases of the Cardiovascular
4 Functional Anatomy of Prokaryotic and Lymphatic Systems 637
and Eukaryotic Cells 76 24 Microbial Diseases of the Respiratory System 674
5 Microbial Metabolism 113 25 Microbial Diseases of the Digestive System 705
6 Microbial Growth 156 26 Microbial Diseases of the Urinary
7 The Control of Microbial Growth 184 and Reproductive Systems 743
8 Microbial Genetics 210
PART FIVE Environmental
9 Biotechnology and Recombinant DNA 246
and Applied Microbiology

PART TWO A Survey of the Microbial World 27 Environmental Microbiology 766

28 Applied and Industrial Microbiology 793
10 Classification of Microorganisms 273
11 The Prokaryotes: Domains Bacteria
and Archaea 299 Answers to Review and Multiple Choice
Study Questions AN-I
12 The Eukaryotes: Fungi, Algae, Protozoa,
and Helminths 329 Appendix A Metabolic Pathways AP-l
13 Viruses, Viroids, and Prions 367
Appendix B Exponents, Exponential Notation,
Logarithms, and Generation Time AP-7
PART THREE Interaction between
Microbe and Host Appendix C Methods for Taking Clinical
Samples AP-S
14 Principles of Disease and Epidemiology 399
Appendix D Pronunciation of Scientific Names
15 Microbial Mechanisms of Pathogenicity 428
AP-9
16 Innate Immunity: Nonspecific Defenses
of the Host 449 Appendix E Word Roots Used in Microbiology
AP-13
17 Adaptive Immunity: Specific Defenses
of the Host 476 Appendix F Classification of Bacteria According
18 Practical Applications of Immu nology 500 to Bergey's Manual AP-16

19 Disorders Associated with the Immune Glossary G-l

System 522
Credits C-l
20 Antimicrobial Drugs 553
Index 1-1

xix
 CONTENTS

PART ONE Fundamentals of Microbiology How Atoms Form Molecules: Chemical

Bonds 28

1 The Microbial World and You 1 Ionic Bonds 28

Covalent Bonds 30
H ydrogen Bonds 31
Microbes in OUf Lives 2
Molecular Weight and Moles 32
Naming and ClassifYing Microorganisms 2
Chemical Reactions 32
Nomenclature 2
Energy in Chemical Reactions 32
Types of Microorganisms 3
Synthesis Reactions 32
Classification of Microorganisms 6
Decomposition Reactions 32
A Brief History of Microbiology 6 Exchange Reactions 33
The First Observations 7 The Reversibility of Reactions 34
The Debate Over Spontaneous Generation 8
IMPORTANT BIOLOGICAL
The Golden Age of Microbiology 9 MOLECULES 34
The Birth of Modern Chemotherapy: Dreams
of a "Magic Bullet" 12 Inorganic Compounds 34
Modern Developments in Microbiology 13 Water 34
Acids, Bases, and Salts 35
Microbes and Human Welfare 16
Acid-Base Balance: The Concept of pH 35
Recycling Vital Elements 16
Sewage Treatment: Using Microbes to Recycle Organic Compounds 37
Water 17 Structure and Chemistry 37
Bioremediation: Using Microbes to Clean Up Carbohydrates 39
Pollutant s 17 Lipids 40
Insect Pest Control by Microorganisms 17 Proteins 42
Modern Biotechnology and Recombinant DNA Nucleic Acids 47
Technology 17 Adenosine Triphosphate (ATP) 47
Microbes and Human Disease 18 Study Outline 49
Normal Microbiota 18
Study Questions 51
Biofilms ] 8
Infectious Diseases 19
Observing Microorganisms
Emerging Infectious Diseases ]9
Study Outline 22 3 through a Microscope 54
Study Questions 23 Units of Measurement 55
Microscopy: The Instruments 55

2 Chemical Principles 26 Light Microscopy 56

Two-Photon Microscopy 62
Scanning Acoustic Microscopy 63
The Structure of Atoms 27 Electron Microscopy 63
Chemical Elements 27 Scanned-Probe Microscopy 65
Electronic Configurations 28

"
 CONTENTS xxi

Preparation of Specimens for Light Microscopy 68 Golgi Complex 104

Preparing Smears for Staining 68 Lysosomes 104
Sim ple Stains 69 Vacuoles 104
Differential Stains 69 Mitochondria 104
Special Stains 71 Chloroplasts 105
Study Outline 73 Peroxisomes 105
Centrosome 105
Study Questions 74
The Evolution of Eukaryotes ] 06
Study Outline 108
Functional Anatomy of Prokaryotic
4 and Eukaryotic Cells 76 Study Questions 110

Comparing Prokaryotic and Eukaryotic Cells:

An Overview 77
THE PROKARYOTIC CELL 77
5 Microbial Metabolism 113
The Size, Shape, and Arrangement of Bacterial Cells 77 Catabolic and Anabolic Reactions 114
Structures External to the Cell Wall 79 Enzymes 115
Glycocalyx 79 Collision Theory 11 5
Flagella 81 Enzymes and Chemical Reactions lIS
Axial Filaments 82 Enzyme Specificity and Efficiency 116
Fimbriae and Pili 83 Naming Enzymes 11 6
The Cell Wall 84 Enzyme Components I 16
Composition and Characteristics 85 The Mechanism of Enzymatic Action 117
Cell Walls and The Gram Stain Mechanism 87 Factors Influencing Enzymatic Activity 11 8
Atypical Cell Walls 87 Feedback Inhibition 120
Damage to the Cell Wall 88 Ribozymes 121
Structures Internal to the Cell Wall 89 Energy Production 12l
The Plasma (Cytoplasmic) Membrane 89 Oxidation-Reduction Reactions 122
The Movement of Materials across The Generation of ATP 122
Membranes 91 Metabolic Pathways of Energy
Cytoplasm 94 Production 123
The Nucleoid 94 Carbohydrate Metabolism 124
Ribosomes 95 Glycolysis 124
Inclusions 95 Alternatives to Glycolysis 125
Endospores 96 Cellular Respiration 127
THE EUKARYOTIC CELL 98 Fermentation 132
Flagella and Cilia 98 Lipid and Protein Catabolism 136
The Cell Wall and Glycocalyx 98 Biochemical Tests and Bacterial
Identification 137
The Plasma (Cytoplasmic) Membrane 100
Photosynthesis 140
Cytoplasm 100
The Light-Dependent Reactions:
Ribosomes 10] Photophosphorylation 140
Organelles l02 The Light-Independent Reactions: The Calvin-Benson
Cycle 140
The Nucleus 102
Endoplasmic Reticulum 103 Chaprer 5 continues
xxii CONTENTS

Chapter 5, continued
A Summary of Energy Production Mechanisms 141 7 The Control of Microbial Growth 184
Metabolic Diversity among Organisms 142
The Terminology of Microbial C ontrol 185
Photoautotrophs 143
The Rate of Microbial Death 186
Photoheterotrophs 145
Chemoautotrophs 145 Actions of Microbial Control Agents 186
Chemoheterotrophs 145 Alteration of Membrane Permeability 186
Damage to Proteins and Nucleic Acids 187
Metabolic Pathways of Energy Use 146
Polysaccharide Biosynthesis 146 Physical Methods of Microbial Control 187
Lipid Biosynthesis 146 Heat 188
Amino Acid and Protein Biosynthesis 146 Filtration 191
Purine and Pyrimidine Biosynthesis 147 Low Temperatures 19 1
High Pressure 192
The Integration of Metabolism 147
Desiccation 192
Study Outline 150 Osmotic Pressure 192
Study Questions 153 Radiation 192
Chemical Methods of Microbial Control 195
Principles of Effective Disinfection 195
6 Microbial Growth 156 Evaluating a Disinfect ant 195
Types of Disinfectants 195
T he Requirements for Growth 157 Microbial Characteristics and Microbial
Physical Requirements 157 Control 202
Chemical Requirements 160 Stud y Outline 205
Biofilms 162
Study Questions 207
C ulture Media 164
Chemically Defined Media 165
Complex Medi a 165
Anaerobic Growth Media and Methods 166
Special C ulture Techniques 167
8 Microbial Genetics 210
Structure and Function of the Genetic Material 211
Selective and Differential Media 168
G enotype and Phenot ype 21 I
Enrichment Culture 169
D NA and Chromosomes 21 1
Obtaining Pure C ultures 170 T he Flow of Genetic Information 2 12
Preserving Bacterial C ultures 170 D NA Replicat ion 2 12
T he Growth of Bacterial C ultures 17 1 RNA and Protein Synthesis 216
Bact erial Division 171 The Regulation of Bacterial Gene Expression 221
Generation T ime 17 1 Repression and Induction 224
Logarithmic Representation of Bact erial T he Operon Model of G ene Expression 224
Populations 17 1 Positive Regulation 225
Phases of Growth I 72
Mutation: C hange in the Genetic Material 226
Direct Measurement of Microbial Growth 174
Types of Mutations 227
Estimating Bacterial Numbers by Indirect
Methods 178 Mutagens 229
T he Frequency of Mutation 23 1
Study Outline 180
Identifying Mutants 231
Study Questions 18 1 Identifying Chemical Carcinogens 232
 CONTENTS xxiii

Genetic Transfer and Recombination 233 C lassification of Organisms 278

Transformation in Bacteria 234 Scientific Nomenclature 278
Conjugation in Bacteria 236 The Taxonomic Hierarchy 279
Transduction in Bacteria 237 Classification of Prokaryotes 279
Plasmids and Transposons 237 Classification of Eukaryotes 281
Genes and Evolution 241 Classification of Viruses 282

Study Outline 242 Methods of C lassifying and IdentifYing

Microorganisms 282
Study Questions 244
Morphological Characteristics 284
Differential Staining 285
Biotechnology and Recombinant
9
Biochemical Tests 285
DNA 246 Serology 287
Phage Typing 288
Introduction to Biotechnology 247 Fatty Acid Profiles 288
Recombinant DNA Technology 247 Flow Cytometry 288
An Overview of Recombinant DNA Procedures 247 DNA Base Composition 288
Tools of Biotechnology 247 DNA Fingerprinting 289
Selection 249 The Polymerase Chain Reaction 290
Mutation 249 Nucleic Acid Hybridization 291
Restriction Enzymes 249 Putting Classification Methods Together 293
Vectors 250 Study Outline 295
Polymerase Chain Reaction 25 I
Study Questions 296
Techniques of Genetic Modification 253
Inserting Foreign DNA into Cells 253
The Prokaryotes: Domains
Obtaining DNA 254
Selecting a Clone 256 11 Bacteria and Archaea 299
Making a Gene Product 257
The Prokaryotic Groups 300
Applications of rO NA 258
DOMAIN BACTERIA 302
Therapeutic Applications 258
The Proteobacteria 302
The Human Genome Project 261
The Alphaproteobacteria 303
Scientific Applications 261
The Betaproteobacteria 305
Agricultural Applications 264
The Gammaproteobacteria 306
Safety Issues and the Ethics of Using rONA 268
The Deltaproteobacteria 312
Study Outline 269 The Epsilonproteobacteria 312
Study Questions 271 The Nonproteobacteria Gram-Negative Bacteria 313
Cyanobacteria (The Oxygenic Photosynthetic
PART TWO A Survey of the Microbial World Bacteria) 313
Purple and Green Photosynthetic Bacteria (The
Classification of
10 Microorganisms 273
Anoxygenic Photosynthetic Bacteria) 315
The Gram-Positive Bacteria 3 15
Firmicutes (Low G + C Gram-Positive Bacteria) 316
The Study of Phylogenetic Relationships 274
Actinobacteria (High G + C Gram-Positive
The Three Domains 274
Bacteria) 320
A Phylogenetic Hierarchy 277
Chapter II continues
xxvi CONTENTS

Chapter 16, continued Practical Applications

Anti microbial Substances 463
The Complement System 463
18 of Immunology 500
Interferons 468 Vaccines so I
Iron-Binding Proteins 470 Principles and Effects of Vacci nation 501
Antimicrobial Peptides 470 Types of Vaccines and Their Characteristics 501
The Development of New Vaccines 504
Study Outli ne 472
Safety of Vaccines 506
Study Questions 474
Diagnostic Immunology 507
Im munologic-Based Diagnostic Tests 507
Adaptive Immunity: Specific
17 Defenses of the Host 476
Monoclonal Antibodies 507
Precipitation Reactions 509
Agglutination Reactions 5 10
The Adaptive Immune System 477 Neutralization Reactions 5 12
Dual Nature of the Adaptive Immune Complement-Fixation Reactions 512
System 477 Fluorescent-Antibody Techniques 5) 3
Humoral Immunity 477
Enzyme-Linked Immunosorbent Assay
Cellular Immunity 477 (E LI SA) 514
Antigens and Antibodies 478 Western Blotting (Immunoblotting) 516
The Nature of Antigens 4 78 The Future of Diagnostic Immunology 516
The Nature of Antibodies 479 Study Outline 5 I 9
B Cells and Humoral Immunity 482 Stud y Questions 520
Clon al Selection of Antibody-Producing
Cells 482
Disorders Associated with
The Diversity of Antibodies 484
Antigen-Antibody Binding and Its Results 484 19 the Immune System 522
T Cells and Cellular Immunity 486 Hypersensitivity 523
Classes ofT Cells 487 Type I (Anaphylactic) Reactions 523
T Helper Cells (CD4+ T Cells) 487 Type " (Cytotoxic) Reactions 526
T Cytotox ic Cells (CD8+ T Cells) 488 Type III (Immune Complex) Reactions 528
T Regulatory Cells 489 Type IV (Delayed Cell-Mediated) Reactions 529
Antigen-Presenting Cells (APCs) 489 Autoimmune Diseases 532
Dendritic Cells 490 Cytotoxic Autoimmune Reactions 53 2
Macrophages 490 Immune Complex Autoimmune Reactions 532
Extracellular Killing by the Immune System 491 Cell-Mediated Autoimmune Reactions 532
Antibody-Dependent Cel l-MLxl iated Reactions Related to the Human Leukocyte Antigen
Cytotoxicity 491 (HLA) Complex 533
Cytokines: C hemical Messengers of Immune Reactions to Transplantation 534
Cells 491 Immunosuppression 536
Immunological Memory 493 The Immune System and Cancer 537
Types of Adaptive Immunity 494 Immunotherapy for Cancer 538

Study Outline 497 Immunodeficiencies 538

Congenital Immunodefi ciencies 538
Study Questions 498
Acquired Immunodeficiencies 538
 CONTENTS xxv

The Etiology of Infectious Diseases 404 How Bacterial Pathogens Damage Host Cells 434
Koch's Postulates 404 Using the Host 's Nutrients: Siderophores 434
Exceptions to Koch's Postulates 404 Direct Damage 434
Classifying Infectious Diseases 406 The Production of Toxins 434
Occurrence of a Disease 406 Plasmids, Lysogeny, and Pathogenicity 439
Severity or Duration of a Disease 406 Pathogenic Properties of Viruses 441
Extent of Host Involvement 407 Viral Mechanisms for Evading Host
Defenses 441
Patterns of Disease 408
Cytopathic Effects of Viruses 441
Predisposing Factors 408
Development of Disease 408 Pathogenic Properties of Fungi, Protozoa, Helminths,
and Algae 442
The Spread of Infection 409
Fungi 443
Reservoirs of Infection 409
Protozoa 443
Transmission of Disease 409
Helminths 444
Nosocomial (Hospital-Acquired) Infections 413 Algae 444
Microorganisms in the Hospital 414
Portals of Exit 444
Compromised Host 41 5
Study Outline 445
Chain of Transmission 415
Control of Nosocomial Infections 416 Study Questions 447

Emerging Infectious Diseases 416

Innate Immunity: Nonspecific
Epidemiology 4 J 8
Descriptive Epidemiology 419
Analytical Epidemiology 41 9
16 Defenses of the Host 449
The Concept of Immunity 450
Experimental Epidemiology 420
Case Reporting 420 FIRST LINE OF DEFENSE: SKIN AND MUCOUS
MEMBRANES 450
The Centers for Disease Control and Prevention 420
Physical Factors 45 J
Study Outline 423
Chemical Factors 453
Study Questions 425
Normal Microbiota and Innate Immunity 453
SECOND LINE OF DEFENSE 454
Microbial Mechanisms
15 of Pathogenicity 428 Formed Elements in Blood 454
The Lymphatic System 456
How Microorganisms Enter a Host 429
Phagocytes 457
Portals of Ent ry 429
Actions of Phagocytic Cells 457
The Preferred Portal of Entry 429
The Mechanism of Phagocytosis 458
Numbers of Invading Microbes 429
Microbial Evasion of Phagocytosis 459
Adherence 431
Inflammation 460
How Bacterial Pathogens Penetrate Host Defenses 432
Vasodilation and Increased Permeability
Capsules 432 of Blood Vessels 460
Cell Wall Components 432 Phagocyte Migration and Phagocytosis 462
Enzymes 432 Tissue Repair 462
Antigenic Variation 433
Fever 463
Penetration into the Host Cell Cytoskeleton 433
Chapter 16 continues
 CONTENTS xxvii

Acquired Immunodefkiency Syndrome Effects of Combinations of Drugs 578

(AlDS) 539
The Future of Chemotherapeutic Agents 578
The Origin of AIDS 540
Antimicrobial Peptides 578
HIV Infection 540
Antisense Agents 579
Diagnostic Methods 545
Study Outline 580
HIV Transmission 545
AIDS Worldwide 546 Study Questions 582
Preventing and Treating AIDS 547
The AIDS Epidemic and the Im portance PART FOUR Microorganisms
of the Scientific Research 548 and Human Disease
Study Outline 549
Microbial Diseases of the Skin
Study Questions 551
21 and Eyes 584
Structure and Function of the Skin 585

20 Antimicrobial Drugs 553 Mucous Membranes 585

Normal Microbiota of the Skin 585
The History of Chemotherapy 554 Microbial Diseases of the Skin 586
Antibiotic Discovery Today 554 Bacterial Diseases of the Skin 586
The Spectrum of Antimicrobial Activity 555 Viral Diseases of the Skin 595
The Action of Antimicrobial Drugs 555 Fungal Diseases of the Skin and Nails 600
Inhibiting Cell Wall Synthesis 556 Parasitic Infestation of the Skin 602
Inh ibiting Protein Synthesis 556 Microbial Diseases of the Eye 603
Injuring the Plasma Membrane 558 Infla mmation of the Eye Membranes:
Inhibiting Nucleic Acid Synthesis 558 Conjunctivitis 603
Inhibiting the Synthesis of Essential Metabolites 558 Bacterial Diseases of the Eye 603
Other Infectious Diseases of the Eye 605
A Survey of Commonly Used Antimicrobial Drugs 559
Antibacterial Antibiotics: Inhibitors of Cell Wall Study Outline 606
Synthesis 559 Study Questions 608
Antimycobacterial Antibiotics 563
Inh ibitors of Protein Synthesis 563
Microbial Diseases of the Nervous
Injury to the Plasma Membrane 566
Inh ibitors of Nucleic Acid (DNA/RN A) Synthesis 567 22 System 610
Competitive Inhibitors of the Synthesis of
Structure and Function of the Nervous
Essential Metabolites 567
System 611
Antifungal Drugs 567
Bacterial Diseases of the Nervous System 611
Antiviral Drugs 569
Bacterial Meningitis 612
Antiprotozoan and Antihelminthic Drugs 57 1
Tetanus 615
Tests to Guide Chemotherapy 572
Botulism 616
The Diffusion Methods 572
Leprosy 619
Broth Dilution Tests 572
Viral Diseases of the Nervous System 620
Resistance to Antimicrobial Drugs 573
Poliomyelitis 620
Mechanisms of Resistance 574
Rabies 622
Antibiotic Misuse 575
Arboviral Encephalitis 624
Cost and Prevention of Resistance 576
Chapter 22 continues
Antibiotic Safety 576
xxviii CONTENTS

Chapter 22, continued Protozoan Diseases of the C ardiovascular and Lymphatic

Systems 660
Fungal Disease of the Nervous Syst em 626
Chagas' Disease (American Trypanosomiasis) 661
Cryptococcus neoformans Meningit is
(Cryptococcosis) 626 Toxoplasmosis 661
Malaria 663
Protozoan Diseases of the Nervous System 627
Leishmaniasis 665
African Trypanosomiasis 627
Babesiosis 666
Amebic Meningoencephalitis 629
Helminthic Diseases of the Cardiovascular and Lymphatic
Nervous System Diseases C aused by Prions 629
Systems 666
Bovine Spongifonn Encephalopathy
Schistosomiasis 666
and Variant Creutzfeldt-Jakob
Disease 63 1 Swimmer's Itch 667

Disease C aused by Unidentified Agents 633 Study Outline 669

Chronic Fatigue Syndrome 633 Study Questions 671
Study Outline 633
Microbial Diseases of the
24
Study Questions 635
Respiratory System 674
Microbial Diseases of the Structure and Function of the Respi ratory System 675
Cardiovascular and Lymphatic
23 Systems 637
Normal Microbiota of the Respiratory System 675
MI C ROBIAL DISEASES OF TH E UPPER
Structure and Function of the C ardiovascular RESPIRATORY SYSTEM 676
and Lymphatic Systems 638 Bacterial Diseases of the Upper Respiratory System 677
Bacterial Diseases of the C ardiovascular and Lymphatic Streptococcal Pharyngitis (Strep Throat) 677
Systems 638 Scarlet Fever 677
Sepsis and Septic Shock 639 Diphtheria 677
Bact erial Infections of the Heart 64 1 Otitis Media 679
Rheumat ic Fever 64 1 Viral Diseases of the Upper Respiratory System 679
Tularemia 64 2 T he Common Cold 679
Brucellosis (Undulant Fever) 643
MI C ROBIAL DISEASES OF TH E LOWER
Anthrax 645 RESPIRATORY SYSTEM 680
Gangrene 646
Bacterial Diseases of the Lower Respiratory
Syst emic Diseases Caused by Bites System 680
and Scratches 647
Pertussis (Whooping Cough) 680
Vector-Transmitted Diseases 648
Tubercu losis 682
Viral Diseases of the C ardiovascular and Lymphatic Bacterial Pneumonias 684
Systems 655
Meliodosis 690
Burkitt's Lymphoma 655
Viral Diseases of the Lower Respiratory System 692
Infectious Mononucleosis 656
Other Diseases and Epstein-Barr Viral Pneumonia 692
Virus 657 Respiratory Syncytial Virus (RSV) 692
Cytomegalovirus Infections 658 Influenza (Flu) 692
Chikungunya Fever 658 Fungal Diseases of the Lower Respiratory System 695
Classic Viral Hemorrhagic Fevers 658 Histoplasmosis 695
Emerging Viral Hemorrhagic Fevers 659 Coccidiomycosis 696
 CONTENTS xxix

Pneumocystis Pneumonia 697 Helminthic Diseases of the Digestive System 732

Blast omycosis (North American Blastomycosis) 697 Tapeworms 732
O t her Fungi Involved in Respiratory Disease 69 7 H ydatid Disease 733
Study Outline 700 Nematodes 734

Study Questions 702 Study Outline 738

Study Questions 741

Microbial Diseases of
25 the Digestive System 705
26 Microbial Diseases of the Urinary
and Reproductive Systems 743
Structure and Function of the Digestive System 706
Normal Microbiota of the Digestive Syst em 706 Structure and Function of the U rinary System 744

Bacterial Diseases of the Mouth 707 Structure and Function of the Reproductive
Systems 744
Dental Caries (Tooth Decay) 707
Periodontal Disease 709 Normal Microbiota of the Urinary and Reproductive
Systems 745
Bacterial Diseases of the Lower Digestive
System 710 DISEASES OF THE URINARY SYSTEM 746
St aphylococcal Food Poisoning (Staphylococcal Bacterial Diseases of the Urinary System 746
Ent erotoxicosis) 7 11 Cystitis 746
Sh igellosis (Bacillary Dysent ery) 71 2 Pyelonephritis 746
Salmonellosis (Salmonella Gastroenteritis) 71 2 Leptospirosis 746
Typhoid Fever 7 14
DISEASES OF THE REPROD UCTIVE
Cholera 7 16 SYSTEMS 747
Noncholera Vibrios 717
Bacterial Diseases of the Reproductive Systems 747
Escherichia coli Gastroenteritis 717
Gonorrhea 747
Ca mpylobacter Gastroenteritis 718
Nongonococcal Urethritis (NGU) 750
Helicobacter Peptic Ulcer Disease 7 18
Pelvic Inflammatory Disease (PID) 751
Yersinia Gastroenteritis 720
Syphilis 752
Clostridium peifrjngens Gastroent eritis 720
Lymphogranuloma Venereum 75 5
Clostridium difficile- Associated Diarrhea 720
Chancroid (Soft Chancre) 756
Bacillus cereus Gastroenteritis 720
Bacterial Vaginosis 756
Viral Diseases of the Digestive System 721
Viral Diseases of the Reproductive System 757
Mumps 721
Genital Herpes 757
Hepatitis 721
Genital Warts 758
Viral G ast roenteritis 728
AID S 758
Fungal Diseases of the Digestive System 729
Fungal Disease of the Reproductive Systems 758
Ergot Poisoning 729
Candidiasis 758
Aflatoxin Poisoning 730
Protozoan Disease of the Reproductive
Protozoan Diseases of the Digestive Systems 760
System 730
Trichomoniasis 760
Giardiasis 730
T he TO RCH Panel of Tests 760
Cryptosporidiosis 73 1
Study Outline 762
Cyclospora Diarrheal Infection 731
Amoebic Dysentery (Amoebiasis) 73 1 Study Questions 764