Make the Connection Between
Lecture, Lab, and the Real World
In its Eleventh Edition, Microbiology: An Introduction helps you make the connection
between microbiological theory presented in the text and real-world applications,
encouraging you to see the connection between human health and microbiology.
▶

FOUNDATION FIGURE 16.7

The Phases of Phagocytosis

Foundation Figures focus on especially
important topics in microbiology.
Clearly marked step-numbers make
process-oriented figures easy to follow,
while the “Key Concepts” highlight the
take-away lessons for easy review. In
MasteringMicrobiology®, Foundation
Figures are highly interactive activities,
designed to guide you through the essential
concepts and processes of microbiology
with in-depth, self-paced tutorials.

A phagocytic macrophage
uses a pseudopod to engulf
nearby bacteria.

Pseudopods

Phagocyte

Cytoplasm

3
2
1

CHEMOTAXIS
and
ADHERENCE
of phagocyte to
microbe

INGESTION
of microbe by phagocyte

Details of
adherence

Formation of phagosome
(phagocytic vesicle)

4

Microbe
or other
particle

Fusion of phagosome
with a lysosome
to form a phagolysosome


Lysosome

5

Digestive
enzymes

PAMP
(peptidoglycan
in cell wall)

Partially
digested
microbe

Stunningly Revised
Foundation Figures

DIGESTION
of ingested
microbes by
enzymes in the
phagolysosome

Indigestible
material

TLR
(Toll-like receptor)

6

Plasma membrane

Formation of
the residual body
containing
indigestible
material

KEYCONCEPTS

•

Chemotaxis, adherence, ingestion, and digestion are phases of
phagocytosis.

•

Chemotaxis allows phagocytes to migrate to infection sites and destroy
invading bacteria.

•

Phagocytosis is an important second line of immune defense. Phagocytes
can also stimulate T and B cells.

•

Toll-like receptors (TLRs) are a focus of current immunological research.


7

DISCHARGE of
waste materials

634

634

diSeaSeS iN FOCuS 22.2
PArT ONE Part Title

Types of Arboviral Encephalitis
461

7497_Ch16_pp0451-0477.indd 461

14/09/11

Disease in Focus
These boxes encourage you to think
like a clinician by making a differential
diagnosis based on a brief clinical
overview. Diseases in Focus include
disease tables, focusing on similar
diseases or infections. These tables
are organized around symptoms
and pathogens in order to be as
clinically relevant as possible.

Disease in Focus activities in
MasteringMicrobiology help you
see the practical applications of
microbiology to your future career.

▶

Arboviral encephalitis is usually characterized by fever, headache, and altered mental status
ranging from confusion to coma. Vector control to decrease contacts between humans and
mosquitoes is the best prevention. Mosquito control includes removing standing water and
using insect repellent while outdoors. An 8-year-old girl in rural Wisconsin has chills, headache,
12:50 PM
and fever and reports having been bitten by mosquitoes. Use the table below to determine
which types of encephalitis are most likely. How would you confirm your diagnosis? For the
solution, go to www.masteringmicrobiology.com
Mosquito
Vector

Reservoir

U.S. Distribution

Culex mosquito engorged with human blood.

Disease

Pathogen

Epidemiology


Mortality

Western
Equine
Encephalitis

WEE virus
(Togavirus)

Culex

Birds,
horses

Severe disease; frequent
neurological damage,
especially in infants

5%

Eastern
Equine
Encephalitis

EEE virus
(Togavirus)

Aedes,
Culiseta


Birds,
horses

More severe than WEE; affects
mostly young children and
younger adults; relatively
uncommon in humans

.30%

St. Louis
Encephalitis

SLE virus
(Flavivirus)

Culex

Birds

Mostly urban outbreaks; affects
mainly adults over 40

20%

California
Encephalitis

CE virus
(Bunyavirus)


Aedes

Small
mammals

Affects mostly 4- to 18-year age
groups in rural or suburban
areas; La Crosse strain medically
most important. rarely fatal;
about 10% have neurological
damage

1% of those
hospitalized

West Nile
Encephalitis

WN virus
(Flavivirus)

Primarily
Culex

Primarily
birds,
assorted
rodents,
and large

mammals

Most cases asymptomatic—
otherwise symptoms vary from
mild to severe; likelihood of
severe neurological symptoms
and fatality increases with age

4–18% of
those
hospitalized

exposure to them is apparently widespread; many in the population carry antibodies—fortunately, symptomatic disease is
rare. Naegleria fowleri is a protozoan (ameba) that causes a neurological disease, primary amebic meningoencephalitis (PAM)
(Figure 22.17). Although scattered cases are reported in most
parts of the world, only a few cases are reported in the United

States annually. The most common victims are children who
swim in warm ponds or streams. The organism initially infects
the nasal mucosa and later penetrates to the brain and proliferates, feeding on brain tissue. The fatality rate is nearly 100%,
death occurring within a few days after symptoms appear.
Because of the rarity of the disease, there is a low “index of


(10-6 m). The prefix
micro indicates that the unit following it should be divided by
1 million, or 106 (see the “Exponential Notation” section in Appen­
dix B). A nanometer (nm) is equal to 0.000000001 m (10-9 m).
Angstrom (Å) was previously used for 10-10 m, or 0.1 nm.
Table 3.1 presents the basic metric units of length and some

of their U.S. equivalents. In Table 3.1, you can compare the mi­
NEW! Clinical Cases
croscopic units of measurement with the commonly known
macroscopic
units in
of measurement,
centimeters,
Clinical Cases
every chaptersuch
helpasmotivate
youmeters,
to
and
kilometers.
If
you
look
ahead
to
Figure
3.2,
you
will
see the
think critically about the chapter content and provide
relative
sizespractical
of variousapplications
organisms onto
theyour

metric
scale.allied
you with
future

▶

clinical case: Microscopic Mayhem
Maryanne, a 42-year-old marketing executive and mother
of three occasionally works from home, but she always feels
that she isn’t getting as much done at home as she does in

healthYoUr
career.Understanding
Each case segment includes a criticalcheck

the office. She has been experiencing recurrent stomach

question
related
tolength,
the chapter
material.
In
✓thinking
If a microbe
measures
10 μm in
how long
is it in

nanometers? 3-1
MasteringMicrobiology®
, additional case studies come
alive with images and questions, leading you through
the process of disease diagnosis.

pain, which seems to be getting worse. She jokes with her
husband that he should buy stock in Pepto-Bismol, because
she buys so much of it. At her husband’s urging, she finally

Microscopy: the instruments

makes an appointment to see
her primary care physician.
After hearing that Maryanne

learning objectives

feels better immediately

3-2 Diagram the path of light through a compound microscope.

after taking Pepto-Bismol, the

3-3 Define total magnification and resolution.

doctor suspects Maryanne

3-4 Identify a use for darkfield, phase-contrast, differential interference
contrast, fluorescence, confocal, two-photon, and scanning

acoustic microscopy, and compare each with brightfield illumination.

may have a peptic ulcer
associated with Helicobacter
5 μm

pylori.

CliniCal FoCUS

Morbidity
and Mortality Weekly Report
PArT onehow
Fundamentals
of Microbiology
3-5 142
Explain
electron
microscopy differsFrom
fromthe
light
microscopy.

What is Helicobacter pylori? read on to find out.

54
▲

64


69

3-6 Identify one use for the TEM, SEM, and scanned-probe microscopes.

Human Tuberculosis–Dallas, Texas

71

The simple microscope used by van Leeuwenhoek in the seven­
As you read through this box, you will
species. Speciation of the M. tuberculosis
teenth century
had
onlythat
one
lens and was
to a magnifying
encounter a series
of questions
laboratory
complexsimilar
is done by biochemical
testing
technicians ask themselves as they identify
bacteria. Try to answer each question before
going on to the next one.
1. Daria, a 12-month-old African American girl,

in reference laboratories (Figure A). The
bacteria need to be grown in culture media.

Slow-growing mycobacteria may take up to
6 weeks to form colonies.

After colonies have been isolated, what is
is brought by her parents to the emergency
the next step?
department of a Dallas, Texas, hospital. She
4.
Two weeks later, the laboratory results
has a fever of 39°C, a distended abdomen,
show that the bacteria are slow-growing.
some abdominal pain, and watery diarrhea.
U.S.
Equivalent
According to the identification scheme, the
Daria is admitted to the pediatric wing of
urease test should be performed.
the hospital, pending results of laboratory
What
is the result shown in Figure B?
3
1000 m = 10 m and radiologic tests. Test results suggest 3280.84 ft or
0.62 mi; 1 mi = 1.61 km
5. Because the urease test is positive, the
peritoneal tuberculosis. Caused by one
nitrate
reduction test is performed. It
of several closely related species in the
Standard unit of Mycobacterium
length tuberculosis complex, TB is39.37 inshows

or 3.28
or 1.09
ydproduce the
that theftbacteria
do not
enzyme nitrate reductase. Daria’s physician
a reportable condition in the United States.
-1
lets
her
parents
know
that
they
are very
Peritoneal
TB
is
a
disease
of
the
intestines
3.94 in
0.1 m = 10 m
close to identifying the pathogen that is
and abdominal cavity.
causing Daria’s illness.
What organ is usually associated with


3.1 Metric Units of Length and U.S. Equivalents
▶
Clinical Focus
Metric Unit
Meaning of Prefix
Metric Equivalent

Table

Clinical Focus boxes contain
kilo = 1000
Morbidity and Mortality Weekly
Report data from the Centers for
deci = 1/10Control and Prevention
Disease
(CDC)
0.394 in; 1 in = 2.54 cm
centi
= 1/100 modified into clinical
0.01 m = 10-2 m
6. M. bovis is a pathogen that primarily infects
problem-solving scenarios with
2. Pulmonary TB is contracted by inhaling the
-3
cattle. However, humans can become
milli = 1/1000
0.001 m = 10 m
bacteria; ingesting the bacteria can result
infected by consuming unpasteurized
questions to help you develop

in peritoneal TB. A laparoscopy reveals that
dairy products or inhaling infectious
-6
nodules
are present in Daria’s abdominal
m
microyour
= 1/1,000,000
0.000001
m
=
10
critical-thinking skills.
cavity. A portion of a nodule is removed

1 kilometer (km)
1 meter (m)
1 decimeter (dm)
1 centimeter (cm)

tuberculosis? How might someone get
peritoneal TB?

1 millimeter (mm)
1 micrometer (μm)

What is the bacterium?

Acid-fast mycobacteria


1 nanometer (nm)

for biopsy so that it can be observed

-9
m of acid-fast bacteria.
0.000000001 m =for10
the presence

nano = 1/1,000,000,000

1 picometer (pm)

Based on the presence of the abdominal
-12physician begins
Daria’s
m
0.000000000001nodules,
m = 10
conventional antituberculosis treatment.
This long-term treatment can last up to
12 months.

pico = 1/1,000,000,000,000
1

Adult tapeworm
releases eggs.
Intermediate
host


6

2

3. The lab results confirm that acid-fast

Human intermediate host
ingests eggs. Dead end.

bacteria are indeed present in Daria’s
abdominal cavity. The laboratory now
needs to identify the Mycobacterium

Egg (30–38 µm)
Adult
tapeworm
LM

0.7 mm

Figure A An identification scheme
for selected species of slow-growing
mycobacteria.

Intermediate
host

Rapid-growing


Urease test

What is the next step?

Scoleces from cyst
attach to intestine and
grow into adults.

7497_Ch03_pp0053-0074.indd 54

Slow-growing

+
Nitrate reductase test
+
M. tuberculosis

–
M. avium

–
M. bovis

Test

Control

Figure B The urease test. In a positive
test, bacterial urease hydrolyzes urea,
producing ammonia. The ammonia raises

the pH, and the indicator in the medium
turns to fuchsia.

droplets from cattle. Human-to-human
transmission occurs only rarely. The clinical
and pathologic characteristics of M. bovis TB
are indistinguishable from M. tuberculosis TB,
but identification of the bacterium is
important for prevention and treatment.
Children may be at higher risk. In one study,
almost half of the culture-positive pediatric
TB cases were caused by M. bovis.
Unfortunately, Daria does not recover
from her illness. Her cadiovascular system
collapses, and she dies. The official cause of
death is peritoneal tuberculosis caused by
M. bovis. Everyone should avoid consuming
products from unpasteurized cow’s milk,
which carry the risk of transmitting M. bovis
if imported from countries where the
bacterium is common in cattle.
Source: Adapted from Rodwell T.C., Moore M., Moser K.S.,
Brodine S.K., Strathdee S.A, “Mycobacterium bovis
Tuberculosis in Binational Communities,” Emerging
Infectious Diseases, June 2008, Volume 14 (6), pp. 909–916.
19/08/11 11:35 AM
Available from />htm.

Scolex


Sexual
reproduction

5

2

Intermediate host
ingests eggs.

▶

Definitive
host

Asexual
reproduction

LM

Definitive host eats
intermediate host,
ingesting cysts.

Larva

10 cm

3
Hydatid cyst


Eggs hatch, and larvae
migrate to liver or lungs.

not produce O2 and is called anoxygenic. The anoxygenic photoautotrophs are the green and purple bacteria. The green bacteria,
such as Chlorobium (klô-rŌ ʹ bē-um), use sulfur (S), sulfur compounds (such as hydrogen sulfide, H2S), or hydrogen gas (H2) to
reduce carbon dioxide and form organic compounds. Applying
the energy from light and the appropriate enzymes, these bacteria

NEW! Life Cycle Figures

Life Cycle figures break down complex
processes into more readily understandable
steps. Each Life Cycle figure is color-coded
to differentiate between steps that involve
sexual or asexual reproduction.

7497_Ch05_pp0111-0152.indd 142

Brood capsule
Scolex

4

Larvae develop
into hyadid cysts.

Figure 12.28 The life cycle of the tapeworm, Echinococcus, spp. Dogs are the most
common definitive host of E. granulosus. E. multilocularis infections in humans are rare. The
parasite can complete its life cycle only if the cysts are ingested by a definitive host that

eats the intermediate host.

Q

Why isn’t being in a human of benefit to Echinococcus?

oxidize sulfide (S2−) or sulfur (S) to sulfate (SO42−) or oxidize
hydrogen gas to water (H2O). The purple bacteria, such as
Chromatium (krō-mā ʹ tē-um), also use sulfur, sulfur compounds,
or hydrogen gas to reduce carbon dioxide. They are distinguished
from the green bacteria by their type of chlorophyll, location of
stored sulfur, and ribosomal RNA.

21/09/11 4:24 PM


Arrive prepared for lecture and lab
The Mastering online homework and tutoring system delivers self-paced
tutorials that provide you with individualized coaching set to your professor’s
course objectives. MasteringMicrobiology helps you arrive better
prepared for lecture and lab with reading questions, coaching activities,
tutorials and more. Research shows that Mastering’s immediate feedback
and tutorial assistance help you understand and master microbiology
concepts— meaning that you retain more knowledge and perform better in
subsequent courses.

▶

NEW! Lab
Technique Videos

Lab Technique Videos are 3-5 minute
videos, demonstrating specific lab
techniques. These videos cover
commonly performed procedures,
such as aseptic technique, Gram
staining, and preparation of smears.
The videos help you get prepared
for your wet lab and also allow
you to review the techniques on
your own time. Quizzes test your
comprehension of the steps involved
in each technique to make sure you
get the most out of the videos.


www.masteringmicrobiology.com

▶

NEW! MicroLab Tutors
These tutors help you get the most out of lab time. Each MicroLab Tutor begins with clinical backround and
a technique video. Select MicroLab Tutors, like the Gram Stain MicroLab Tutor, also contain an animation
illustrating the procedure at the molecular level, helping you visualize each process. Each tutorial’s questions
contain hints and feedback that include photomicrographs, video clips or animation clips and are designed to
make sure that you are prepared for lab by introducing and assessing your understanding of lab concepts and
techniques outside of formal lecture and lab time. Select Tutors will contain an animation illustrating the
procedure at the molecular level, as is the case in this sample for the Gram stain tutor.

What instructors are saying —
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lab time needed for explanation
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Texas A&M University

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both correct and incorrect procedures.”
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East Mississippi Community College


Unparalleled Online Resources for
Additional Student Practice and Assessment
All of the resources previously found on the Microbiology Place™
website are now accessible and assignable in MasteringMicrobiology®.
MasteringMicrobiology builds on these study tools and includes new
content and assessments, enabling more frequent student practice
and more meaningful course management.
▶
MicroFlix™ are 3D movie-quality animations
with self-paced tutorials and gradable 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®

animations that help them review relevant
concepts from general biology.

Foundation Figures ▶
Coaching Activities
Foundations Figures are reinforced
in MasteringMicrobiology® with
Coaching Activities that ensure
students master the toughest topics
before moving on in the chapter. The
results of the Coaching Activities
feed directly into the gradebook.


www.masteringmicrobiology.com

▶

Case Study
Coaching Activities
These activities in
MasteringMicrobiology
help students connect
microbiological theory to
real-world disease diagnosis
and treatment, are assignable,
and feed directly into the
MasteringMicrobiology
gradebook.


2-D Microbiology
Animations

▶

More than 120 multi-step
Microbiology Animations explain
and visually demonstrate core
concepts, providing an additional
opportunity for students to visualize
and understand core microbiology
concepts. They are accompanied by
gradable quizzes. References to the
Microbiology Animations appear
throughout the chapters of the book.

For Instructors
A TRUSTED PARTNER

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The Mastering platform was developed by scientists for
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MasteringMicrobiology can be successfully
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Integrated usage of MasteringMicrobiology
has demonstrated quantifiable differences
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Mastering questions are tied to the
specific Learning Outcomes in Tortora,
Funke, and Case as well as global science
Learning Outcomes and those provided by
the American Society of Microbiology Center for Undergraduate
Educators. These provide a powerful tool for tracking individual
student learning and assessing course objectives.


The Best Support for
Instructors and Students
NEW! Laboratory
Experiments in
Microbiology,
Tenth Edition

▶

by Ted R. Johnson
and Christine L. Case
978-0-321-79438-3 • 0-321-79438-9
Containing 57 thoroughly classtested exercises, this manual
provides engaging labs with
instruction on performing basic
microbiology techniques and

applications in diverse areas, including the biological sciences,
allied health sciences, agriculture, environmental science,
nutrition, pharmacy, and various pre-professional programs. The
Tenth Edition is easily customizable and features an updated art
program and a full-color design, integrating valuable micrographs
throughout each exercise. Additionally, many of the illustrations
have been re-rendered in a modern, realistic, three-dimensional
style to better visually engage students. Experiments have been
refined throughout the manual and the Tenth Edition includes
a new exercise using pGLO to demonstrate transformation in
bacteria and introduce students to this important technique.

Also available to help prepare
your students for lab:
Preparation Guide for Laboratory
Experiments in Microbiology,
Tenth Edition
by Ted R. Johnson and Christine L. Case
978-0-321-80910-0 • 0-321-80910-6

Techniques for Microbiology:
A Student Handbook
by John M. Lammert
978-0-13-224011-6 • 0-13-224011-4
Lammert’s Techniques in Microbiology
is highly visual and incorporates “voice
balloons” that keep you focused on the
relevant process. The techniques are those
that will be used frequently for studying
microbes in the laboratory, and include

those identified by the American Society
for Microbiology in its recommendations
for the Microbiology Laboratory Core
Curriculum (recommendations in which
the author participated).

ADDITIONAL SUPPLEMENTS
For Instructors
Instructor Resource DVD/CD-ROM
978-0-321-79309-6 • 0-321-79309-9
This cross-platform set of DVDs organizes instructor
media resources by chapter for easy reference and
presentation. The instructor media package includes:

• All figures from the book with and without
labels in both JPEG and PowerPoint® formats
• All figures from the book with the Label
Edit feature in PowerPoint 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 and BioFlix® Animations
• PowerPoint lecture outlines, including
figures from the book, tables from the
book, and links to multimedia
• Clicker Questions
• The Instructor Guide and Test Bank
as editable Microsoft® Word files

• Test Bank in TestGen® and Word formats.

For Students
Instructor Guide/Test Bank

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Microbiology
An Introduction
ELEVENTH EDITION

Gerard J. Tortora
Bergen Community College
Berdell R. Funke
North Dakota State University
Christine L. Case
Skyline College


Acquisitions Editor: Kelsey Volker
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Cover Photo Credit: Alfred Pasieka/Photo Researchers, Inc.
Credits and acknowledgments for material borrowed from other sources and reproduced, with
permission, in this textbook appear on the appropriate page within the text or after the Glossary.
Copyright © 2013, 2010, 2007 Pearson Education, Inc. All rights reserved. Manufactured in the
United States of America. This publication is protected by Copyright, and permission should be
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Library of Congress Cataloging-in-Publication Data
Tortora, Gerard J.
Microbiology : an introduction / Gerard J. Tortora, Berdell R. Funke, Christine L. Case.—11th ed.
       p. ; cm.
  Includes bibliographical references and index.
  ISBN-13: 978-0-321-73360-3 (student ed.)
  ISBN-10: 0-321-73360-6 (student ed.)
  ISBN-13: 978-0-321-79310-2 (exam copy)
  ISBN-10: 0-321-79310-2 (exam copy)
  I. Funke, Berdell R. II. Case, Christine L., 1948- III. Title.
  [DNLM: 1.  Microbiology.  QW 4]
 
  579—dc23
                                                            2011042916

ISBN 10: 0-321-73360-6; ISBN 13: 978-0-321-73360-3 (Student edition)
ISBN 10: 0-321-79310-2; ISBN 13: 978-0-321-79310-2 (Instructor’s Review Copy)
www.pearsonhighered.com

1 2 3 4 5 6 7 8 9 10—CRK—15 14 13 12 11


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 for Microbiology
(ASM), Human Anatomy and Physiology Society (HAPS), American
Association for the Advancement of Science (AAAS), National Education

Courtesy of Rev.
Association (NEA), New Jersey Educational Association (NJEA), and
Dr. James F. Tortora
the Metropolitan Association of College and University Biologists (MACUB). 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 40 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 (SIM) 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 Community/Tribal 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


PREFACE
Since the publication of the first edition nearly 30 years ago, well
over one million students have used Microbiology: An Introduction
at colleges and universities around the world, making it the
leading textbook for non-majors microbiology. The eleventh
edition continues to be a comprehensive beginning text, assuming 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
science, animal science, forestry, agriculture, home economics,
and the liberal arts.
The eleventh edition has retained the features that have made
this book so popular:
An appropriate balance between microbiological
fundamentals and applications, and between medical
applications and other applied areas of microbiology. Basic
microbiological principles are given greater emphasis than
are applications, and health-related applications are featured.
■ Straightforward presentation of complex topics. Each section of the text is written with the student in mind.
■ Clear, accurate, and pedagogically effective illustrations
and photos. Step-by-step diagrams that closely coordinate
with narrative descriptions aid student comprehension of
concepts.

■

Flexible organization. We have organized the book in what
we think is a useful fashion while recognizing that the material might be effectively presented in 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 guidelines for organizing the material in several other ways.

■

New to the Eleventh Edition
The visual introduction at the beginning of the book contains
more details on the eleventh edition.
The eleventh edition meets all students at their respective levels
of skill and understanding while addressing the biggest challenges
that instructors face. Updates to the new eleventh edition enhance
the book’s consistent pedagogy and clear explanations. Some of
the highlights of the eleventh edition follow:

Cutting-edge media integration. MasteringMicrobiology
(www.masteringmicrobiology.com) provides unprecedented,
cutting-edge assessment resources for instructors as well as
self-study tools for students. The 3-D MicroFlix and Microbiology Animations allow students to visualize key concepts;
new Foundation Figure questions allow students to master the
foundational material; new Case Studies stress real-world applications; and Lab Technique videos partner with the lab manual
to prepare students so that they get the most out of lab time.
■ New Clinical Cases that relate the study of microbiology to
real-world applications. The Clinical Cases allow students
to apply what they have learned to real-life scenarios. As

the student reads the chapter they can follow along with the
Clinical Case and answer critical thinking questions that directly relate to the material that they have just read.
■ Illustrations and photos that enhance student understanding. The Foundation Figures and Life Cycle figures have
been stunningly revised to foster student comprehension.
The Foundation Figures, which integrate text and visuals to
help students master the core concepts of microbiology, now
include a bulleted list of Key Concepts. All stepwise figures
(including Foundation Figures and Life Cycle figures) have
been made to be entirely self-explanatory so that the student
doesn’t have to rely on lengthy captions to follow them. The
new edition also includes over 100 new electron and light
micrographs of quality unmatched in the market.
■

Addition of a Name It! activity to the Study Questions at
the end of each chapter. This question provides clues about
the physical and biochemical nature of a microbe, signs and
symptoms of the disease the microbe causes, information
about treatment, etc., and then asks students to use their
critical thinking skills to identify the microbe.

■

Chapter-by-Chapter Revisions
Every chapter in this edition has been thoroughly revised, and
data in the text, tables, Clinical Focus boxes, and figures have
been updated through February 2011. The main changes to each
chapter are summarized below.
Chapter 1


A new section on H1N1 influenza (swine flu) has been added.
A new section on multi-drug-resistant tuberculosis has been

■
■

added.
Figure 1.3 is now a Foundation Figure.

■

iv




PREFACE

Chapter 2

A new table on chemical bonds has been added.
■ A new table compares DNA and RNA.
■

Chapter 5

The discussion of photophosphorylation has been revised.

■


Chapter 16

The section on inflammation has been revised.
The table on innate immunity responses has been revised.

■
■

Chapter 17

A discussion of TH17 T cells and the ineffectiveness of other

■

T cells to deal with certain infections has been greatly
expanded.

Chapter 6

A new Applications of Microbiology box addresses life in

■

extreme environments.

Chapter 8

MicroRNA and epigenetic control are now included.

■


Chapter 9

The discussions of gene silencing and forensic microbiology

■

have been revised.

Examples of veterinary uses of rDNA technology and

■

nanotechnology are included.
The Minimal Genome Project is introduced.

■

Chapter 10

The tree of life has been revised to include new information

■

on horizontal gene transfer between lineages.
■ A molecular clock is introduced.
■ Nucleic acid amplification tests are explained.
Chapter 11

The section on the nonproteobacteria gram-negative bacteria


■

has been reorganized.

The material on purple and green photosynthetic bacteria

■

has been extensively revised. A discussion of the deinococci
has been added.

Chapter 12

Newest changes to fungal and protozoan taxonomy are included.
■ The chapter now includes discussion of microsporidia,
■

emerging opportunistic pathogens.

Chapter 13

Discussions on influenza epidemics and crossing the species

■

barrier have been updated.

Chapter 14


Data in the epidemiology graphs have been updated through

■

2010.

A new section on the Human Microbiome Project has been

■

added.
■ A new section on health care–associated infections has been
included.
Chapter 15

A new Applications of Microbiology box addresses streptokinase.

■

Chapter 18

The discussion of the various types of vaccines has been

■

extensively updated and revised.
The discussion of adjuvants has been thoroughly updated
and revised.
■ A discussion of needle-free vaccines has been added.
■ The significance of spelling of the names of monoclonal antibodies is now explained.

■

Chapter 19

The discussion of HIV/AIDS has been extensively updated

■

and revised.

Chapter 20

Discussion of some of the newer antibiotics and antibiotic

■

types, such as the pleuromutilins, has been added.

A discussion of artemisinin-based malarial treatments is now

■

included.
The discussion of antibiotic-resistant superbugs has been
expanded.
■ An essay on the future of chemotherapeutic agents has been
added.
■

Chapter 21


An outbreak of Pseudomonas dermatitis is described in the
Clinical Case.

■

Chapter 22

The discussion of the meningococcal diseases and vaccines

■

for polio has been extensively revised.
Maps, graphs, and other art have been updated and revised.

■

Chapter 23

The first case of dengue fever acquired in the United States is
described in the Clinical Case.

■

Chapter 24

The discussion of the etiology and symptoms of the common

■


cold has been expanded.
The diagnosis of tuberculosis has been updated and expanded.
■ The discussion of influenza has been considerably expanded
and updated.
■

Chapter 25

The discussions of traveler’s diarrhea (E. coli gastroenteritis)

■

and hepatitis B infections have been revised extensively.
Discussion of Clostridium difficile–associated diarrhea is now
included.

■

v


PREFACE 

vi

Chapter 26

The discussion of the gonococcus now describes Opa proteins.
The discussion of neonatal herpes and genital warts has been


■
■

updated and revised.

Chapter 27

The figure depicting the sulfur cycle has been revised.

■

Chapter 28

Quality assurance microbiology is demonstrated in the Clinical

■

Case.


ACknowledgments
In preparing this textbook, we have benefited from the guidance
and advice of a large number of microbiology instructors across
the country. These reviewers have provided constructive criticism and valuable suggestions at various stages of the revision.
We gratefully acknowledge our debt to these individuals.
Michelle L. Badon, The University of Texas at Arlington
James K. Collins, University of Arizona
Robin L. Cotter, Phoenix College
Melissa A. Deadmond, Truckee Meadows Community College
Jennifer Freed, Rio Salado College

Edwin Gines-Candelaria, Miami Dade College
Fran Hardin, Ivy Tech Community College of Indiana
Dr. Mark Jaffe, Nova Southeastern University
Judy Kaufman, Monroe Community College
Ken Malachowsky, Florence-Darlington Technical College
John L. McKillip, Ball State University
Janie Milner, Santa Fe Community College
Virendra Nayyar, Windward Community College
Susan B. Roman, Georgia State University
Chris Sowers, Forsyth Technical Community College
Paula Steiert, St. John’s College of Nursing of Southwest Baptist
University
Donald L. Terpening, Ulster County Community College
John E. Whitlock, Hillsborough Community College
Brenda Zink, Northeastern Junior College

We also thank the staff at Benjamin Cummings for their dedication to excellence. Kelsey Volker, our acquisitions editor, successfully kept us all focused on where we wanted this revision to
go. Katie Cook, project editor, masterfully managed the book’s
schedule and progress, keeping communication lines open and
ensuring the highest quality at every stage. Sally Peyrefitte’s careful attention to continuity and detail in her copyedit of both text
and art served to keep concepts and information clear throughout. The developmental editor, Cindi Crimson Jones, was of
great assistance throughout the project.
  Michele Mangelli worked closely with editorial during the
early stages of this revision and masterfully guided the book
through the complex production process by managing the production team. Janet Vail expertly guided the text through the
production process and managed the day-to-day work flow. Elisheva Marcus and Marilyn Perry developed the stunning new
Foundation Figures and Life Cycle figures. Elisheva Marcus directed revisions to the art and photo program, provided concept
and style development, and worked closely with the team to ensure content accuracy and aesthetic standards. The talented staff
at Precision Graphics gracefully managed the high volume and


complex updates of our art and photo program. David Novak coordinated the many complex stages of the art and photo processing rendering. Our photo researcher, Maureen Spuhler, made
sure we had clear and striking images throughout the book. Gary
Hespenheide created the elegant interior design, and Yvo Riezebos did a wonderful job with the cover. The skilled team at Nesbitt Graphics moved this book through the composition process.
Karen Hollister prepared the index, and Betsy Dietrich carefully
proofread all of the pages. Stacey Weinberger guided the book
through the manufacturing process.
  Denise Wright of Southern Editorial impeccably handled the
instructor and student supplements. Liz Winer managed the
media program, working many miracles to produce the impressive array of resources in MasteringMicrobiology. Dorothy Cox
and Shannon Kong managed the print and media supplements
through the complex production stages.
  Neena Bali, Executive Marketing Manager, and the entire Pearson sales force do a stellar job presenting this book to instructors
and students and ensuring its unwavering status as the best-selling
microbiology textbook.
  We would 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 J. Tortora   Berdell R. Funke   Christine L. Case

vii


Brief Contents
PART ONE  Fundamentals of Microbiology
1 The Microbial World and You  1
2 Chemical Principles  25
3 Observing Microorganisms Through
a Microscope  53

4 Functional Anatomy of Prokaryotic
and Eukaryotic Cells  75
5 Microbial Metabolism  111
6 Microbial Growth  153
7 The Control of Microbial Growth  181
8 Microbial Genetics  207
9 Biotechnology and DNA Technology  244

PART TWO A Survey of the Microbial
World
10 Classification of Microorganisms  272
11 The Prokaryotes: Domains Bacteria
and Archaea  299
12 The Eukaryotes: Fungi, Algae, Protozoa,
and Helminths  330
13 Viruses, Viroids, and Prions  369

PART THREE Interaction between
Microbe and Host
14 Principles of Disease and Epidemiology  401
15 Microbial Mechanisms of Pathogenicity  429
16 Innate Immunity: Nonspecific Defenses
of the Host  451
17 Adaptive Immunity: Specific Defenses
of the Host  478
18 Practical Applications of Immunology  504
19 Disorders Associated with the
Immune System  527
20 Antimicrobial Drugs  558


viii

PART FOUR  Microorganisms
and Human Disease
21 Microbial Diseases of the Skin and Eyes  589
22 Microbial Diseases of the Nervous System  615
23 Microbial Diseases of the Cardiovascular
and Lymphatic Systems  643
24 Microbial Diseases of the Respiratory
System  680
25 Microbial Diseases of the Digestive System  711
26 Microbial Diseases of the Urinary
and Reproductive Systems  749

PART FIVE Environmental
and Applied Microbiology
27 Environmental Microbiology  772
28 Applied and Industrial Microbiology  799
Answers to Review and Multiple Choice
Study Questions  AN-1
Appendix A Metabolic Pathways  AP-1
Appendix B Exponents, Exponential Notation,
Logarithms, and Generation Time  AP-7
Appendix C Methods for Taking Clinical
Samples  AP-8
Appendix D Pronunciation of Scientific Names  AP-9
Appendix E Word Roots Used in Microbiology  AP-13
Appendix F
Glossary  G-1
Credits  C-1

Index  I-1

Classification of Prokaryotes According to
Bergey’s Manual  AP-16


Contents
PART ONE Fundamentals of Microbiology 

1

 The Microbial World
and You  1

Microbes in Our Lives  2
Naming and Classifying Microorganisms  2
Nomenclature • Types of Microorganisms • Classification
of Microorganisms 

Organic Compounds  36
Structure and Chemistry • Carbohydrates • Lipids • Proteins
• Nucleic Acids • Adenosine Triphosphate (ATP) 
Study Outline • Study Questions  48

3

Observing Microorganisms
Through a Microscope  53

A Brief History of Microbiology  6

The First Observations • The Debate over Spontaneous
Generation • The Golden Age of Microbiology • The Birth
of Modern Chemotherapy: Dreams of a “Magic Bullet”
• Modern Developments in Microbiology 

Units of Measurement  54

Microbes and Human Welfare  15
Recycling Vital Elements • Sewage Treatment: Using
Microbes to Recycle Water • Bioremediation: Using
Microbes to Clean Up Pollutants • Insect Pest Control
by Microorganisms • Modern Biotechnology and
Recombinant DNA Technology 

Preparation of Specimens for Light Microscopy  64
Preparing Smears for Staining • Simple Stains • Differential
Stains • Special Stains 

Microbes and Human Disease  16
Normal Microbiota • Biofilms • Infectious
Diseases • Emerging Infectious Diseases 
Study Outline • Study Questions  21

2

Chemical Principles  25

The Structure of Atoms  26
Chemical Elements • Electronic Configurations 
How Atoms Form Molecules: Chemical Bonds  27

Ionic Bonds • Covalent Bonds • Hydrogen Bonds
• Molecular Weight and Moles 
Chemical Reactions  31
Energy in Chemical Reactions • Synthesis Reactions
• Decomposition Reactions • Exchange Reactions
• The Reversibility of Chemical Reactions 
IMPORTANT BIOLOGICAL MOLECULES  33
Inorganic Compounds  33
Water • Acids, Bases, and Salts • Acid–Base Balance:
The Concept of pH 

Microscopy: The Instruments  54
Light Microscopy • Two-Photon Microscopy • Scanning
Acoustic Microscopy • Electron Microscopy • ScannedProbe Microscopy 

Study Outline • Study Questions  71

4

Functional Anatomy

of Prokaryotic and
Eukaryotic Cells  75

Comparing Prokaryotic and Eukaryotic Cells:
An Overview  76
THE PROKARYOTIC CELL  76
The Size, Shape, and Arrangement of Bacterial Cells  77
Structures External to the Cell Wall  78
Glycocalyx • Flagella • Axial Filaments • Fimbriae and Pili

The Cell Wall  84
Composition and Characteristics • Cell Walls and the
Gram Stain Mechanism • Atypical Cell Walls • Damage
to the Cell Wall 
Structures Internal to the Cell Wall  88
The Plasma (Cytoplasmic) Membrane • The Movement
of Materials across Membranes • Cytoplasm • The Nucleoid
• Ribosomes • Inclusions • Endospores 
THE EUKARYOTIC CELL  97
Flagella and Cilia  99
The Cell Wall and Glycocalyx  99
ix


x

CONTENTS

The Plasma (Cytoplasmic) Membrane  100

Biofilms  160

Cytoplasm  101

Culture Media  161
Chemically Defined Media • Complex Media • Anaerobic
Growth Media and Methods • Special Culture Techniques
• Selective and Differential Media • Enrichment Culture 

Ribosomes  101

Organelles  101
The Nucleus • Endoplasmic Reticulum • Golgi Complex
• Lysosomes • Vacuoles • Mitochondria • Chloroplasts
• Peroxisomes • Centrosome 
The Evolution of Eukaryotes  105
Study Outline • Study Questions  106

5 

Microbial Metabolism  111

Catabolic and Anabolic Reactions  112
Enzymes  113
Collision Theory • Enzymes and Chemical Reactions
• Enzyme Specificity and Efficiency • Naming Enzymes
• Enzyme Components • The Mechanism of Enzymatic
Action • Factors Influencing Enzymatic Activity • Feedback
Inhibition • Ribozymes 
Energy Production  119
Oxidation-Reduction Reactions • The Generation of ATP
• Metabolic Pathways of Energy Production 
Carbohydrate Catabolism  122
Glycolysis • Alternatives to Glycolysis • Cellular Respiration
• Fermentation 
Lipid and Protein Catabolism  133
Biochemical Tests and Bacterial Identification  135
Photosynthesis  138
The Light-Dependent Reactions: Photophosphorylation
• The Light-Independent Reactions: The Calvin-Benson Cycle
A Summary of Energy Production Mechanisms  139

Metabolic Diversity among Organisms  140
Photoautotrophs • Photoheterotrophs • Chemoautotrophs
• Chemoheterotrophs 
Metabolic Pathways of Energy Use  144
Polysaccharide Biosynthesis • Lipid Biosynthesis • Amino
Acid and Protein Biosynthesis • Purine and Pyrimidine
Biosynthesis 
The Integration of Metabolism  146
Study Outline • Study Questions  148

6 

Microbial Growth  153

The Requirements for Growth  154
Physical Requirements • Chemical Requirements 

Obtaining Pure Cultures  167
Preserving Bacterial Cultures  167
The Growth of Bacterial Cultures  168
Bacterial Division • Generation Time • Logarithmic
Representation of Bacterial Populations • Phases of Growth
• Direct Measurement of Microbial Growth • Estimating
Bacterial Numbers by Indirect Methods 
Study Outline • Study Questions  177

7

The Control of Microbial
Growth  181


The Terminology of Microbial Control  182
The Rate of Microbial Death  182
Actions of Microbial Control Agents  183
Alteration of Membrane Permeability • Damage to Proteins
and Nucleic Acids 
Physical Methods of Microbial Control  185
Heat • Filtration • Low Temperatures • High Pressure
• Desiccation • Osmotic Pressure • Radiation 
Chemical Methods of Microbial Control  190
Principles of Effective Disinfection • Evaluating a
Disinfectant • Types of Disinfectants 
Microbial Characteristics and Microbial Control  200
Study Outline • Study Questions  203

8 

Microbial Genetics  207

Structure and Function of the Genetic Material  208
Genotype and Phenotype • DNA and Chromosomes • The
Flow of Genetic Information • DNA Replication • RNA and
Protein Synthesis 
The Regulation of Bacterial Gene Expression  218
Pre-transcriptional Control • Post-transcriptional Control 
Mutation: Change in the Genetic Material  223
Types of Mutations • Mutagens • The Frequency of
Mutation • Identifying Mutants • Identifying Chemical
Carcinogens 
Genetic Transfer and Recombination  231

Transformation in Bacteria • Conjugation in Bacteria
• Transduction in Bacteria • Plasmids and Transposons 




CONTENTS

Genes and Evolution  239
Study Outline • Study Questions  239

9

Biotechnology and DNA
Technology  244

Introduction to Biotechnology  245
Recombinant DNA Technology • An Overview of
Recombinant DNA Procedures 
Tools of Biotechnology  247
Selection • Mutation • Restriction Enzymes • Vectors
• Polymerase Chain Reaction 
Techniques of Genetic Modification  251
Inserting Foreign DNA into Cells • Obtaining DNA
• Selecting a Clone • Making a Gene Product 
Applications of DNA Technology  257
Therapeutic Applications • Genome Projects • Scientific
Applications • Agricultural Applications 
Safety Issues and the Ethics of Using DNA Technology  266
Study Outline • Study Questions  268


PART TWO A Survey of the
Microbial World 

10

Classification of

Microorganisms  272

The Study of Phylogenetic Relationships  273
The Three Domains • A Phylogenetic Hierarchy 
Classification of Organisms  277
Scientific Nomenclature • The Taxonomic Hierarchy
• Classification of Prokaryotes • Classification of Eukaryotes
• Classification of Viruses 
Methods of Classifying and Identifying Microorganisms  281
Morphological Characteristics • Differential Staining
• Biochemical Tests • Serology • Phage Typing • Fatty Acid
Profiles • Flow Cytometry • DNA Base Composition • DNA
Fingerprinting • Nucleic Acid Amplification Tests (NAATs)
• Nucleic Acid Hybridization • Putting Classification
Methods Together 
Study Outline • Study Questions  295

11

The Prokaryotes: Domains
Bacteria and Archaea  299


The Prokaryotic Groups  300
DOMAIN BACTERIA  303

The Proteobacteria  303
The Alphaproteobacteria • The Betaproteobacteria • The
Gammaproteobacteria • The Deltaproteobacteria • The
Epsilonproteobacteria 
The Gram-Positive Bacteria  314
Firmicutes (Low G 1 C Gram-Positive Bacteria)
• Actinobacteria (High G 1 C Gram-Positive Bacteria) 
The Nonproteobacteria Gram-Negative Bacteria  320
Cyanobacteria (The Oxygenic Photosynthetic Bacteria)
• Chlamydiae • Planctomycetes • Bacteroidetes
Fusobacteria  322
Purple and Green Photosynthetic Bacteria (The Anoxygenic
Photosynthetic Bacteria) • Spirochaetes • Deinococci 
DOMAIN ARCHAEA  326
Diversity within the Archaea  326
MICROBIAL DIVERSITY  327
Discoveries Illustrating the Range of Diversity  327
Study Outline • Study Questions  328

12

 The Eukaryotes: Fungi,

Algae, Protozoa,
and Helminths  330

Fungi  331

Characteristics of Fungi • Medically Important Fungi
• Fungal Diseases • Economic Effects of Fungi 
Lichens  342
Algae  343
Characteristics of Algae • Selected Phyla of Algae • Roles of
Algae in Nature 
Protozoa  348
Characteristics of Protozoa • Medically Important Protozoa 
Slime Molds  353
Helminths  354
Characteristics of Helminths • Platyhelminths • Nematodes 
Arthropods as Vectors  363
Study Outline • Study Questions  365

13

Viruses, Viroids, and
Prions  369

General Characteristics of Viruses  370
Host Range • Viral Size 
Viral Structure  371
Nucleic Acid • Capsid and Envelope • General Morphology 

xi


xii

CONTENTS


Taxonomy of Viruses  374
Isolation, Cultivation, and Identification of Viruses  376
Growing Bacteriophages in the Laboratory • Growing
Animal Viruses in the Laboratory • Viral Identification 
Viral Multiplication  381
Multiplication of Bacteriophages • Multiplication of Animal
Viruses 
Viruses and Cancer  392
The Transformation of Normal Cells into Tumor Cells
• DNA Oncogenic Viruses • RNA Oncogenic Viruses 
Latent Viral Infections  394
Persistent Viral Infections  394
Prions  395

15

 Microbial Mechanisms of
Pathogenicity  429

How Microorganisms Enter a Host  430
Portals of Entry • The Preferred Portal of Entry • Numbers
of Invading Microbes • Adherence 
How Bacterial Pathogens Penetrate Host Defenses  433
Capsules • Cell Wall Components • Enzymes • Antigenic
Variation • Penetration into the Host Cell Cytoskeleton 
How Bacterial Pathogens Damage Host Cells  436
Using the Host’s Nutrients: Siderophores • Direct Damage
• The Production of Toxins • Plasmids, Lysogeny, and
Pathogenicity 


Study Outline • Study Questions  397

Pathogenic Properties of Viruses  443
Viral Mechanisms for Evading Host Defenses • Cytopathic
Effects of Viruses 

PART THREE Interaction between Microbe
and Host 

Pathogenic Properties of Fungi, Protozoa, Helminths, and
Algae  445
Fungi • Protozoa • Helminths • Algae 

Plant Viruses and Viroids  395

14

 Principles of Disease
and Epidemiology  401

Pathology, Infection, and Disease  402
Normal Microbiota  402
Relationships between the Normal Microbiota and the Host
• Opportunistic Microorganisms • Cooperation among
Microorganisms 

Portals of Exit  446
Study Outline • Study Questions  448


16

Innate Immunity:

Nonspecific Defenses
of the Host  451

The Concept of Immunity  452

The Etiology of Infectious Diseases  406
Koch’s Postulates • Exceptions to Koch’s Postulates 

FIRST LINE OF DEFENSE: SKIN AND MUCOUS
MEMBRANES  453

Classifying Infectious Diseases  408
Occurrence of a Disease • Severity or Duration of a Disease
• Extent of Host Involvement 

Physical Factors  453
Chemical Factors  455

Patterns of Disease  409
Predisposing Factors • Development of Disease 

SECOND LINE OF DEFENSE  456

The Spread of Infection  411
Reservoirs of Infection • Transmission of Disease 


The Lymphatic System  458

Nosocomial (Hospital-Acquired) Infections  414
Microorganisms in the Hospital • Compromised Host
• Chain of Transmission • Control of Nosocomial Infections 
Emerging Infectious Diseases  417
Epidemiology  419
Descriptive Epidemiology • Analytical Epidemiology
• Experimental Epidemiology • Case Reporting • The
Centers for Disease Control and Prevention (CDC) 
Study Outline • Study Questions  424

Normal Microbiota and Innate Immunity  455
Formed Elements in Blood  456
Phagocytes  460
Actions of Phagocytic Cells • The Mechanism of
Phagocytosis • Microbial Evasion of Phagocytosis 
Inflammation  463
Vasodilation and Increased Permeability of Blood Vessels
• Phagocyte Migration and Phagocytosis • Tissue Repair 
Fever  466
Antimicrobial Substances  466
The Complement System • Interferons • Iron-Binding
Proteins • Antimicrobial Peptides 
Study Outline • Study Questions  475




17


CONTENTS

Adaptive Immunity:

Specific Defenses of
the Host  478

The Adaptive Immune System  479
Dual Nature of the Adaptive Immune System  479
Humoral Immunity • Cellular Immunity 
Antigens and Antibodies  481
The Nature of Antigens • The Nature of Antibodies 
B Cells and Humoral Immunity  485
Clonal Selection of Antibody-Producing Cells
• The Diversity of Antibodies 
Antigen–Antibody Binding and Its Results  487
T Cells and Cellular Immunity  489
Classes of T Cells • T Helper Cells (CD41 T Cells)
• T Regulatory Cells • T Cytotoxic Cells (CD81 T Cells) 
Antigen-Presenting Cells (APCs)  494
Dendritic Cells • Macrophages 

19

Disorders Associated with
the Immune System  527

Hypersensitivity  528
Type I (Anaphylactic) Reactions • Type II (Cytotoxic)

Reactions • Type III (Immune Complex) Reactions
• Type IV (Delayed Cell-Mediated) Reactions 
Autoimmune Diseases  536
Cytotoxic Autoimmune Reactions • Immune Complex
Autoimmune Reactions • Cell-Mediated Autoimmune
Reactions 
Reactions Related to the Human Leukocyte Antigen (HLA)
Complex  538
Reactions to Transplantation • Immunosuppression 
The Immune System and Cancer  542
Immunotherapy for Cancer 
Immunodeficiencies  543
Congenital Immunodeficiencies • Acquired
Immunodeficiencies 

Immunological Memory  497

Acquired Immunodeficiency Syndrome (AIDS)  545
The Origin of AIDS • HIV Infection • Diagnostic Methods
• HIV Transmission • AIDS Worldwide • Preventing and
Treating AIDS • The AIDS Epidemic and the Importance of
Scientific Research 

Types of Adaptive Immunity  497

Study Outline • Study Questions  554

Extracellular Killing by the Immune System  495
Antibody-Dependent Cell-Mediated Cytotoxicity  495
Cytokines: Chemical Messengers of Immune Cells  495


Study Outline • Study Questions  501

18

Practical Applications
of Immunology  504

Vaccines  505
Principles and Effects of Vaccination • Types of Vaccines
and Their Characteristics • The Development of New
Vaccines • Adjuvants • Safety of Vaccines 
Diagnostic Immunology  511
Immunologic-Based Diagnostic Tests • Monoclonal
Antibodies • Precipitation Reactions • Agglutination
Reactions • Neutralization Reactions • ComplementFixation Reactions • Fluorescent-Antibody Techniques
• Enzyme-Linked Immunosorbent Assay (ELISA) • Western
Blotting (Immunoblotting) • The Future of Diagnostic and
Therapeutic Immunology 
Study Outline • Study Questions  524

20 

Antimicrobial Drugs  558

The History of Chemotherapy  559
Antibiotic Discovery Today 
The Spectrum of Antimicrobial Activity  560
The Action of Antimicrobial Drugs  561
Inhibiting Cell Wall Synthesis • Inhibiting Protein Synthesis

• Injuring the Plasma Membrane • Inhibiting Nucleic Acid
Synthesis • Inhibiting the Synthesis of Essential Metabolites 
A Survey of Commonly Used Antimicrobial Drugs  564
Antibacterial Antibiotics: Inhibitors of Cell Wall Synthesis
• Antimycobacterial Antibiotics • Inhibitors of Protein
Synthesis • Injury to the Plasma Membrane • Inhibitors
of Nucleic Acid (DNA/RNA) Synthesis • Competitive
Inhibitors of the Synthesis of Essential Metabolites
• Antifungal Drugs • Antiviral Drugs • Antiprotozoan
and Antihelminthic Drugs 
Tests to Guide Chemotherapy  577
The Diffusion Methods • Broth Dilution Tests 

xiii


xiv

CONTENTS

Resistance to Antimicrobial Drugs  579
Mechanisms of Resistance • Antibiotic Misuse • Cost and
Prevention of Resistance 
Antibiotic Safety  584

Disease Caused by Unidentified Agents  638
Chronic Fatigue Syndrome 
Study Outline • Study Questions  639

Effects of Combinations of Drugs  584

The Future of Chemotherapeutic Agents  584
Study Outline • Study Questions  585

PART FOUR Microorganisms
and Human Disease 

21

Microbial Diseases of the
Skin and Eyes  589

Structure and Function of the Skin  590
Mucous Membranes 
Normal Microbiota of the Skin  591
Microbial Diseases of the Skin  591
Bacterial Diseases of the Skin • Viral Diseases of the Skin
• Fungal Diseases of the Skin and Nails • Parasitic Infestation
of the Skin 
Microbial Diseases of the Eye  609
Inflammation of the Eye Membranes: Conjunctivitis
• Bacterial Diseases of the Eye • Other Infectious Diseases
of the Eye 
Study Outline • Study Questions  611

22

Microbial Diseases of the
Nervous System  615

Structure and Function of the Nervous System  616

Bacterial Diseases of the Nervous System  617
Bacterial Meningitis • Tetanus • Botulism • Leprosy 
Viral Diseases of the Nervous System  626
Poliomyelitis • Rabies • Arboviral Encephalitis 
Fungal Disease of the Nervous System  632
Cryptococcus neoformans Meningitis (Cryptococcosis) 
Protozoan Diseases of the Nervous System  633
African Trypanosomiasis • Amebic Meningoencephalitis 
Nervous System Diseases Caused by Prions  636
Bovine Spongiform Encephalopathy and Variant
Creutzfeldt-Jakob Disease 

23

Microbial Diseases of

the Cardiovascular and
Lymphatic Systems  643

Structure and Function of the Cardiovascular and Lymphatic
Systems  644
Bacterial Diseases of the Cardiovascular and Lymphatic
Systems  645
Sepsis and Septic Shock • Bacterial Infections of the Heart
• Rheumatic Fever • Tularemia • Brucellosis (Undulant Fever)
• Anthrax • Gangrene • Systemic Diseases Caused by Bites
and Scratches • Vector-Transmitted Diseases 
Viral Diseases of the Cardiovascular and Lymphatic
Systems  662
Burkitt’s Lymphoma • Infectious Mononucleosis • Other

Diseases and Epstein-Barr Virus • Cytomegalovirus
Infections • Chikungunya Fever • Classic Viral Hemorrhagic
Fevers • Emerging Viral Hemorrhagic Fevers 
Protozoan Diseases of the Cardiovascular and Lymphatic
Systems  666
Chagas’ Disease (American Trypanosomiasis)
• Toxoplasmosis • Malaria • Leishmaniasis • Babesiosis 
Helminthic Diseases of the Cardiovascular and Lymphatic
Systems  673
Schistosomiasis • Swimmer’s Itch 
Study Outline • Study Questions  676

24

Microbial Diseases of the
Respiratory System  680

Structure and Function of the Respiratory System  681
Normal Microbiota of the Respiratory System  682
MICROBIAL DISEASES OF THE UPPER RESPIRATORY
SYSTEM  682
Bacterial Diseases of the Upper Respiratory System  683
Streptococcal Pharyngitis (Strep Throat) • Scarlet Fever
• Diphtheria • Otitis Media 
Viral Disease of the Upper Respiratory System  685
The Common Cold 





CONTENTS

MICROBIAL DISEASES OF THE LOWER RESPIRATORY
SYSTEM  687
Bacterial Diseases of the Lower Respiratory System  687
Pertussis (Whooping Cough) • Tuberculosis • Bacterial
Pneumonias • Melioidosis 
Viral Diseases of the Lower Respiratory System  697
Viral Pneumonia • Respiratory Syncytial Virus (RSV)
• Influenza (Flu) 
Fungal Diseases of the Lower Respiratory System  702
Histoplasmosis • Coccidioidomycosis • Pneumocystis
Pneumonia • Blastomycosis (North American
Blastomycosis) • Other Fungi Involved in Respiratory
Disease 
Study Outline • Study Questions  707

25

Microbial Diseases of the
Digestive System  711

Structure and Function of the Digestive System  712
Normal Microbiota of the Digestive System  712
Bacterial Diseases of the Mouth  713
Dental Caries (Tooth Decay) • Periodontal Disease 
Bacterial Diseases of the Lower Digestive System  716
Staphylococcal Food Poisoning (Staphylococcal
Enterotoxicosis) • Shigellosis (Bacillary Dysentery)
• Salmonellosis (Salmonella Gastroenteritis) • Typhoid

Fever • Cholera • Noncholera Vibrios • Escherichia coli
Gastroenteritis • Campylobacter Gastroenteritis
• Helicobacter Peptic Ulcer Disease • Yersinia Gastroenteritis
• Clostridium perfringens Gastroenteritis • Clostridium
difficile–Associated Diarrhea • Bacillus cereus
Gastroenteritis 
Viral Diseases of the Digestive System  727
Mumps • Hepatitis • Viral Gastroenteritis 
Fungal Diseases of the Digestive System  735
Ergot Poisoning • Aflatoxin Poisoning 
Protozoan Diseases of the Digestive System  736
Giardiasis • Cryptosporidiosis • Cyclospora Diarrheal
Infection • Amebic Dysentery (Amebiasis) 
Helminthic Diseases of the Digestive System  738
Tapeworms • Hydatid Disease • Nematodes 
Study Outline • Study Questions  744

26

Microbial Diseases of the

Urinary and Reproductive
Systems  749

Structure and Function of the Urinary System  750
Structure and Function of the Reproductive Systems  750
Normal Microbiota of the Urinary and Reproductive
Systems  751
DISEASES OF THE URINARY SYSTEM  752
Bacterial Diseases of the Urinary System  752

Cystitis • Pyelonephritis • Leptospirosis 
DISEASES OF THE REPRODUCTIVE SYSTEMS  754
Bacterial Diseases of the Reproductive Systems  754
Gonorrhea • Nongonococcal Urethritis (NGU) • Pelvic
Inflammatory Disease (PID) • Syphilis • Lymphogranuloma
Venereum (LGV) • Chancroid (Soft Chancre) • Bacterial
Vaginosis 
Viral Diseases of the Reproductive Systems  763
Genital Herpes • Genital Warts • AIDS 
Fungal Disease of the Reproductive Systems  765
Candidiasis 
Protozoan Disease of the Reproductive Systems  766
Trichomoniasis • The TORCH Panel of Tests 
Study Outline • Study Questions  768

PART FIVE Environmental and Applied
Microbiology 

27

Environmental
Microbiology  772

Microbial Diversity and Habitats  773
Symbiosis 
Soil Microbiology and Biogeochemical Cycles  774
The Carbon Cycle • The Nitrogen Cycle • The Sulfur Cycle
• Life without Sunshine • The Phosphorus Cycle • The
Degradation of Synthetic Chemicals in Soil and Water 
Aquatic Microbiology and Sewage Treatment  782

Aquatic Microorganisms • The Role of Microorganisms in
Water Quality • Water Treatment • Sewage (Wastewater)
Treatment 
Study Outline • Study Questions  795

xv


xvi

CONTENTS

28

Applied and Industrial
Microbiology  799

Food Microbiology  800
Foods and Disease • Industrial Food Canning • Aseptic
Packaging • Radiation and Industrial Food Preservation
• High-Pressure Food Preservation • The Role of
Microorganisms in Food Production 
Industrial Microbiology  807
Fermentation Technology • Industrial Products
• Alternative Energy Sources Using Microorganisms
• Biofuels • Industrial Microbiology and the Future 
Study Outline • Study Questions  815

Answers to Review and Multiple Choice
Study Questions  AN-1

Appendix A

Metabolic Pathways  AP-1

Appendix B

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

Appendix C

Methods for Taking Clinical
Samples  AP-8

Appendix D

Pronunciation of Scientific
Names  AP-9

Appendix E

Word Roots Used in
Microbiology  AP-13

Appendix F

Classification of Prokaryotes
According to Bergey’s Manual  AP-16

Glossary  G-1

Credits  C-1
Index  I-1


FEATURES
FOUNDATION FIGURES
Figure 1.3
Figure 2.16
Figure 3.2
Figure 4.6
Figure 5.11
Figure 6.15
Figure 7.1
Figure 8.2
Figure 9.1
Figure 10.1
Figure 12.1
Figure 13.15
Figure 14.3
Figure 15.4
Figure 15.9
Figure 16.7
Figure 16.9
Figure 17.20
Figure 18.2
Figure 19.16
Figure 20.2
Figure 20.20

Disproving the Theory of Spontaneous

Generation  9
The Structure of DNA  46
Microscopes and Magnification  58
The Structure of a Prokaryotic Cell  79
An Overview of Respiration and Fermentation  123
Understanding the Bacterial Growth Curve  170
Understanding the Microbial Death Curve  184
The Flow of Genetic Information  210
A Typical Genetic Modification Procedure  246
The Three-Domain System  274
Exploring Pathogenic Eukaryotes  331
Replication of a DNA-Containing
Animal Virus  387
Koch’s Postulates: Understanding Disease  407
Mechanisms of Exotoxins and Endotoxins  437
Microbial Mechanisms of Pathogenicity  447
The Phases of Phagocytosis  461
Outcomes of Complement Activation  468
The Dual Nature of the Adaptive Immune
System  500
The Production of Monoclonal Antibodies  513
The Progression of HIV Infection  548
Major Action Modes of Antimicrobial Drugs  561
Bacterial Resistance to Antibiotics  580

Life Cycle Figures
Figure 11.11
Figure 11.22
Figure 12.7
Figure 12.8

Figure 12.9
Figure 12.10
Figure 12.13
Figure 12.16
Figure 12.20
Figure 12.22
Figure 12.23
Figure 12.26

The Life Cycle of Myxococcales  313
The Life Cycle of Chlamydias  323
The Life Cycle of Rhizopus, a Zygomycete  336
The Life Cycle of Encephalitozoon,
a Microsporidian  337
The Life Cycle of Talaromyces, an Ascomycete  338
A Generalized Life Cycle of a Basidiomycete  339
Green Algae  345
Oomycotes  347
The Life Cycle of Plasmodium vivax  352
The Generalized Life Cycle of a Cellular
Slime Mold  354
The Life Cycle of a Plasmodial Slime Mold  355
The Life Cycle of the Lung Fluke,
Paragonimus spp.  359

Figure 12.28
Figure 23.14
Figure 23.17
Figure 23.24
Figure 23.28

Figure 24.18
Figure 24.20
Figure 25.26

The Life Cycle of the Tapeworm,
Echinococcus spp.  361
The Life Cycle of the Tick Vector of Lyme
Disease  659
The Life Cycle of the Tick Vector (Dermacentor
spp.) of Rocky Mountain Spotted Fever  661
The Life Cycle of Toxoplasma gondii  669
Schistosomiasis  674
The Life Cycle of Coccidioides immitis  703
The Life Cycle of Pneumocystis jirovecii  705
The Life Cycle of Trichinella spiralis 743

Clinical Focus
Human Tuberculosis—Dallas, Texas  142
Infection Following Steroid Injection  198
Tracking West Nile Virus  220
Norovirus—Who Is Responsible for the Outbreak?  265
The Most Frequent Cause of Recreational Waterborne
Diarrhea  357
Influenza: Crossing the Species Barrier  374
Nosocomial Infections  423
A World Health Problem  510
A Delayed Rash  537
Antibiotics in Animal Feed Linked to Human Disease  583
Infections in the Gym  598
A Neurological Disease  631

A Sick Child  651
Outbreak  698
A Foodborne Infection  721
Survival of the Fittest  757

Applications of Microbiology
Designer Jeans: Made by Microbes?  3
Bioremediation—Bacteria Clean Up Pollution  32
What Is That Slime?  56
Why Microbiologists Study Termites  106
What Is Fermentation?  134
Life in the Extreme—Hydrothermal Vents  157
Mass Deaths of Marine Mammals Spur Veterinary
Microbiology  282
Bacteria and Insect Sex  308
Streptococcus: Harmful or Helpful?  434
Serum Collection  472
xvii