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Micro
biology
An Introduction

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Micro
biology
An Introduction
Twelfth Edition
Gerard J. Tortora
Bergen Community College
Berdell R. Funke
North Dakota State University
Christine L. Case
Skyline College

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Library of Congress Cataloging-in-Publication Data

Tortora, Gerard J., author.
Microbiology : an introduction / Gerard J. Tortora, Berdell R. Funke, Christine L. Case. -- Twelfth edition.
p. ; cm.
Includes bibliographical references and index.
ISBN 978-0-321-92915-0 (student edition)
ISBN 0-321-92915-2 (student edition)
ISBN 978-0-13-390557-1 (instructor’s review copy)
ISBN 0-13-390557-8 (instructor’s review copy)
I. Funke, Berdell R., author. II. Case, Christine L., 1948- , author. III. Title.
[DNLM: 1. Microbiology. QW 4]
QR41.2
579--dc23
2014038680
ISBN 10: 0-321-92915-2; ISBN 13: 978-0-321-92915-0 (Student edition)

ISBN 10: 0-13-390557-8; ISBN 13: 978-0-13-390557-1 (Instructor’s Review Copy)
www.pearsonhighered.com 1 2 3 4 5 6 7 8 9 10—V357—18 17 16 15 14

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
Courtesy of Rev.
Association for the Advancement of Science (AAAS), National Education
Dr. James F. Tortora Association (NEA), New Jersey Educational Association (NJEA), and
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 44 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.


Preface
Since the publication of the first edition nearly 30 years ago, well cutting-edge assessment resources for instructors as well as
over 1 million students have used Microbiology: An Introduction self-study tools for students. Big Picture Coaching Activi-
at colleges and universities around the world, making it the lead- ties are paired with the book’s new Big Picture: Tough Topics
ing textbook for non-majors microbiology. The twelfth edition and Big Picture: Disease features; Interactive Microbiology is
continues to be a comprehensive beginning text, assuming no a dynamic suite of interactive tutorials and animations that
previous study of biology or chemistry. The text is appropriate for teach key concepts in microbiology; and MicroBoosters are
students in a wide variety of programs, including the allied health brief video tutorials that cover key concepts that some
sciences, biological sciences environmental science, animal sci- students need to review or re-learn.
ence, forestry, agriculture, home economics, and the liberal arts. ● Big Picture “tough topic” features. These two-page
The twelfth edition has retained the features that have made spreads focus on the most challenging topics for students to
this book so popular: master: metabolism (Chapter 5), genetics (Chapter 8), and
● An appropriate balance between microbiological immunology (Chapter 16). Each spread breaks down these
fundamentals and applications, and between medical important concepts into manageable steps and gives students
applications and other applied areas of microbiology. a clear learning framework for the related chapters. Each
Basic microbiological principles are given greater includes a quick-reference (QR) code that allows students to
emphasis, and health-related applications are featured. link to related MicroFlix videos with their smartphones.
● Straightforward presentation of complex topics. Each
● Big Picture Disease features. These two-page spreads appear
section of the text is written with the student in mind. within each organ-system disease chapter (Chapters 21–26)
as well as Chapter 19 (Disorders of the Immune System).
● Clear, accurate, and pedagogically effective illustrations
Each spread focuses on a particular disease and applies it to a
and photos. Step-by-step diagrams that closely coordinate
related real-world challenge, many dealing with public health
with narrative descriptions aid student comprehension of
issues.
concepts.
● Reworked complement section in Chapter 16 (Innate
● Flexible organization. We have organized the book in
Immunity: Nonspecific Defenses of the Host). New art
what we think is a useful fashion while recognizing that the
and more straightforward discussions make this challenging
material might be effectively presented in other sequences. For
and critical material easier for students to understand
instructors who wish to use a different order, we have made each
and retain.
chapter as independent as possible and have included numerous
cross-references. The Instructor’s Guide provides detailed
● In the Clinic. This new feature, appearing at the start of
guidelines for organizing the material in several other ways. every chapter, includes critical thinking questions that
encourage students to think as health care professionals
would in various clinical scenarios and spark student interest
New to the Twelfth Edition in the forthcoming chapter content.
The twelfth edition focuses on big-picture concepts and themes ● ASM guidelines. The American Society of Microbiology
in microbiology, encouraging students to visualize and synthesize has released six underlying concepts and 22 related topics to
more difficult topics such as microbial metabolism, immunology, provide a framework for key microbiological topics deemed
and microbial genetics. to be of lasting importance beyond the classroom. The
The twelfth edition meets all students at their respective twelfth edition explains the themes and competencies at the
levels of skill and understanding while addressing the biggest beginning of the book and incorporates callouts when chapter
challenges that instructors face. Updates to the twelfth edition content matches one of these 22 topics. Doing so addresses
enhance the book’s consistent pedagogy and clear explanations. two key challenges: it helps students and instructors focus
Some of the highlights follow. on the enduring principles of the course, and it provides
● Cutting-edge media integration. MasteringMicrobiology another pedagogical tool for instructors to assess students’
(www.masteringmicrobiology.com) provides unprecedented, understanding and encourage critical thinking.
vii

viii Preface
Chapter-by-Chapter Revisions Chapter 13

● A discussion of the use of oncolytic viruses to treat cancer
Every chapter in this edition has been thoroughly revised, and has been added.
data in the text, tables, and figures have been updated. The main ● The discussion of viral enzymes has been revised.
changes to each chapter are summarized below.
Chapter 14
Chapter 1 ● The chapter has been updated to reflect the use of the term
● New sections on Middle East respiratory syndrome (MERS), healthcare-associated infection.
coronavirus, and severe acute respiratory syndrome (SARS)
Chapter 16
have been added.
● A new Big Picture feature, addressing immunity, has been
● A new table, Table 1.2, addresses representative discoveries of
added.
the Golden Age of Microbiology.
● A new figure and discussion of hematopoiesis have been
Chapter 2 added.
● The section on activation energy has been revised. ● Figure 16.14 has been revised.
Chapter 3 ● The discussions of the complement system and interferons
● Foundation Figure 3.2, Microscopes and Magnification, has have been extensively revised.
been revised. Chapter 17
Chapter 4 ● The introductory material has been revised.
● The discussion of facilitated diffusion has been revised. ● Several figures have been revised.
● The cell art has been revised. Chapter 18
Chapter 5 ● The tables showing vaccination schedules have been updated.
● A new Big Picture feature, addressing metabolism, has been ● A discussion of virus-like particle (VLP) vaccines has been
added. added.
● The discussion of enzyme specificity has been revised. ● Clinical Focus box has been rewritten and updated.
● Figure 5.25, showing photophosphorylation, has been revised. ● The discussions of vaccination technologies and monoclonal
● The discussion of chemoheterotrophs has been revised. antibodies have been updated.
Chapter 8 Chapter 19
● A new Big Picture feature, addressing genetics, has been added. ● A new Big Picture Disease feature, Human Microbiome and
● The central dogma of genetics is described. IBD, has been added.
● Mutation and gene transfers are now included in a new section. ● The discussion of HIV/AIDS has been updated with new,
Chapter 9
informative maps.
● Vectors are defined.
● The chemotherapy of AIDS section has been completely
revised, including new figures depicting the action of HIV
Chapter 10 therapies.
● Figure 10.9, showing the new EnteroPluri-test, is revised.
Chapter 20
Chapter 11 ● The discussion of antiviral drugs has been updated.
● The order Thiotrichales is now included. ● The discussion of antibiotics effective against dormant cells
● Discussion of the new genus Cronobacter has been added. has been expanded.
● Several of the figures have been replaced with improved
illustrations. Chapter 21
● The tables have been revised and simplified.
● A new Big Picture Disease feature, Fungal Keratitis, has been
● Nomenclature has been updated. added.
● A discussion of hand-foot-and-mouth disease is now
Chapter 12 included.
● The discussion of algal and protozoan taxonomy is updated.

ix
PREFACE
Chapter 22 ● A discussion of Kawasaki syndrome has been added.

● A new Big Picture Disease feature, Neglected Tropical ● The discussion of dengue and severe dengue is updated.
Diseases, has been added. Chapter 24
● The discussion of developments in testing for leprosy has ● A new Big Picture Disease feature, Pertussis, has been added.
been updated. ● The discussion of melioidosis has been updated.
Chapter 23 Chapter 25
● A new Big Picture Disease feature, Climate Change and ● A new Big Picture Disease feature, Cholera After Natural
Disease, has been added. Disasters, has been added.
● Several of the maps have been updated.
Chapter 26
● The discussion of sepsis and septic shock has been revised.
● The discussion of Lyme disease has been revised to include
● A new Big Picture Disease feature, STI Home Test Kits, has
the topic of immunity to reinfection. been added.

Acknowledgments
In preparing this textbook, we have benefited from the guidance Michele Mangelli worked closely with editorial during the
and advice of a large number of microbiology instructors across early stages of this revision and masterfully guided the book
the country. These reviewers have provided constructive criti- through the complex production process by managing the pro-
cism and valuable suggestions at various stages of the revision. duction team. Karen Gulliver expertly guided the text through the
We gratefully acknowledge our debt to these individuals. production process and managed the day-to-day work flow. Kelly
Murphy and Erin Strathmann worked closely in the development
Payam Benyamini, University of California, Los Angeles
of the new Big Picture features and received invaluable help and
Shima Chaudhary, South Texas College
Jean Cremins, Middlesex Community College instruction from Professor Judy Meier Penn, Shoreline Commu-
Michael J. Dul, Central Arizona College nity College; Dr. Mark Hollier, Georgia Perimeter College, Deca-
Axel Duwe, Diablo Valley College–Pleasant Hill Campus tur; and Dr. Warner Bair, Lone Star College, CyFair. Without their
Jennifer Freed, Rio Salado College input, these informative and compelling features could not have
Ellen Fynan, Worcester State University been conceived. Dr. Hollier also provided expert feedback and
Kamal M. Gandhi, United States University and National University revisions on the Immune System for this edition. Kelly Murphy
Gina Holland, Sacramento City College directed revisions to the art and photo program, provided con-
Suzanne Keller, Indian Hills Community College cept and style development, and worked closely with the team
Janette Gomos Klein, Hunter College to ensure content accuracy and aesthetic standards. The talented
Peter Kourtev, Central Michigan University staff at Precision Graphics gracefully managed the high volume
Carol R. Lauzon, California State University, East Bay
and complex updates of our art and photo program. Jean Lake
Mark R. Liles, Auburn University
coordinated the many complex stages of the art and photo pro-
Mary G. Miller, Baton Rouge Community College
Paul Mink, Lansing Community College cessing rendering. Our photo researcher, Kristin Piljay, made sure
Fernando P. Monroy, Northern Arizona University we had clear and striking images throughout the book. Gary Hes-
Rita B. Moyes, Texas A&M University penheide created the elegant interior design and cover. The skilled
Marcia Pierce, Eastern Kentucky University team at Cenveo Publisher Services moved this book through the
Ben Rowley, University of Central Arkansas composition process. Sallie Steele prepared the index, and Betsy
Heather Seitz, Johnson County Community College Dietrich carefully proofread all of the pages. Stacey Weinberger
Karen Sellins, Front Range Community College guided the book through the manufacturing process.
Elizabeth Sharpe-Aparicio, Blinn College Joe Mochnick managed the media program and produced the
Henry Siu, Miami Dade College–North Campus impressive array of resources in MasteringMicrobiology. Doro-
Michelle Stettner, Meridian Community College thy Cox and Kyle Doctor managed the print and media supple-
Jennifer R. Walker, University of Georgia
ments through the complex production stages.
Patricia G. Wilber, Central New Mexico Community College
Neena Bali and Lauren Harp, Executive Product Marketing
We also thank the staff at Pearson Education for their dedication to Managers, and the entire Pearson sales force do a stellar job pre-
excellence. Kelsey Churchman, senior acquisitions editor, success- senting this book to instructors and students and ensuring its
fully kept us all focused on where we wanted this revision to go. unwavering status as the best-selling microbiology textbook.
Jessica Picone, project manager, masterfully managed the book’s We would like to acknowledge our spouses and families,
schedule and progress, keeping communication lines open and en- who have provided invaluable support throughout the writing
suring the highest quality at every stage. Chriscelle Palaganas, pro- process.
gram manager, provided overall help and support to the team. Sally Finally, we have an enduring appreciation for our students,
Peyrefitte’s careful attention to continuity and detail in her copyedit whose comments and suggestions provide insight and remind us
of both text and art served to keep concepts and information clear of their needs. This text is for them.
throughout. The developmental editors, Erin Strathmann and
Laura Cheu, were of great assistance throughout the project. Gerard J. Tortora Berdell R. Funke Christine L. Case

Brief Contents
PART ONE Fundamentals of Microbiology PART FOUR Microorganisms

1 The Microbial World and You 1 and Human Disease
2 Chemical Principles 24 21 Microbial Diseases of the Skin and Eyes 579
3 Observing Microorganisms Through 22 Microbial Diseases of the Nervous System 607
a Microscope 51 23 Microbial Diseases of the Cardiovascular
4 Functional Anatomy of Prokaryotic and Lymphatic Systems 637
and Eukaryotic Cells 72 24 Microbial Diseases of the Respiratory System 675
5 Microbial Metabolism 107 25 Microbial Diseases of the Digestive System 707
6 Microbial Growth 149 26 Microbial Diseases of the Urinary
7 The Control of Microbial Growth 176 and Reproductive Systems 746
8 Microbial Genetics 201
PART FIVE Environmental
9 Biotechnology and DNA Technology 238
and Applied Microbiology
PART TWO A Survey of the Microbial World 27 Environmental Microbiology 771
28 Applied and Industrial Microbiology 794
10 Classification of Microorganisms 264
11 The Prokaryotes: Domains Bacteria and Archaea 290
Answers to Knowledge and Comprehension
12 The Eukaryotes: Fungi, Algae, Protozoa, Questions AN-1
and Helminths 319
Appendix A Metabolic Pathways AP-1
13 Viruses, Viroids, and Prions 358
Appendix B Exponents, Exponential Notation,
PART THREE Interaction Logarithms, and Generation Time AP-3
between Microbe and Host Appendix C Methods for Taking Clinical
14 Principles of Disease and Epidemiology 389 Samples AP-5
15 Microbial Mechanisms of Pathogenicity 417 Appendix D Pronunciation of Scientific Names AP-7
16 Innate Immunity: Nonspecific Defenses Appendix E Word Roots Used in Microbiology AP-9
of the Host 439
Appendix F Classification of Prokaryotes According
17 Adaptive Immunity: Specific Defenses of the Host 468
to Bergey’s Manual AP-13
18 Practical Applications of Immunology 492
Glossary G-1
19 Disorders Associated with the Immune System 515
Credits C-1
20 Antimicrobial Drugs 548
Index I-1
xi


Contents
PART ONE Fundamentals of Microbiology

3 Observing Microorganisms
1 The Microbial World Through a Microscope 51

and You 1 Units of Measurement 52
Microscopy: The Instruments 52
Microbes in Our Lives 2 Light Microscopy • Two-Photon Microscopy • Scanning Acoustic
Naming and Classifying Microorganisms 2 Microscopy • Electron Microscopy • Scanned-Probe Microscopy
Nomenclature • Types of Microorganisms • Classification Preparation of Specimens for Light Microscopy 62
of Microorganisms Preparing Smears for Staining • Simple Stains • Differential
A Brief History of Microbiology 6 Stains • Special Stains
The First Observations • The Debate over Spontaneous Study Outline • Study Questions 69
Generation • The Golden Age of Microbiology • The Birth
of Modern Chemotherapy: Dreams of a “Magic Bullet”
4
• Modern Developments in Microbiology
unctional Anatomy of Prokaryotic
F
Microbes and Human Welfare 13
Recycling Vital Elements • Sewage Treatment: Using Microbes and Eukaryotic Cells 72
to Recycle Water • Bioremediation: Using Microbes to Clean Up
Pollutants • Insect Pest Control by Microorganisms • Modern Comparing Prokaryotic and Eukaryotic Cells: An Overview 73
Biotechnology and Recombinant DNA Technology THE PROKARYOTIC CELL 73
Microbes and Human Disease 15 The Size, Shape, and Arrangement of Bacterial Cells 73
Normal Microbiota • Biofilms • Infectious Diseases Structures External to the Cell Wall 75
• Emerging Infectious Diseases Glycocalyx • Flagella • Axial Filaments • Fimbriae and Pili
Study Outline • Study Questions 20 The Cell Wall 80
2 Chemical Principles 24
Composition and Characteristics • Cell Walls and the Gram Stain
Mechanism • Atypical Cell Walls • Damage to the Cell Wall
Structures Internal to the Cell Wall 85
The Plasma (Cytoplasmic) Membrane • The Movement
The Structure of Atoms 25
Chemical Elements • Electronic Configurations of Materials across Membranes • Cytoplasm • The Nucleoid
• Ribosomes • Inclusions • Endospores
How Atoms Form Molecules: Chemical Bonds 27
THE EUKARYOTIC CELL 94
Ionic Bonds • Covalent Bonds • Hydrogen Bonds • Molecular
Weight and Moles Flagella and Cilia 96
Chemical Reactions 30 The Cell Wall and Glycocalyx 96
Energy in Chemical Reactions • Synthesis Reactions The Plasma (Cytoplasmic) Membrane 97
• Decomposition Reactions • Exchange Reactions Cytoplasm 98
• The Reversibility of Chemical Reactions Ribosomes 98
Important Biological Molecules 31 Organelles 98
Inorganic Compounds 32 The Nucleus • Endoplasmic Reticulum • Golgi Complex
Water • Acids, Bases, and Salts • Acid–Base Balance: • Lysosomes • Vacuoles • Mitochondria • Chloroplasts
The Concept of pH • Peroxisomes • Centrosome
Organic Compounds 34 The Evolution of Eukaryotes 102
Structure and Chemistry • Carbohydrates • Lipids • Proteins Study Outline • Study Questions 103
• Nucleic Acids • Adenosine Triphosphate (ATP)
Study Outline • Study Questions 47
xiii

xiv CONTENTS
5 Microbial Metabolism 107 7 The Control of Microbial

Growth 176
Catabolic and Anabolic Reactions 110 The Terminology of Microbial Control 177
Enzymes 111 The Rate of Microbial Death 178
Collision Theory • Enzymes and Chemical Reactions Actions of Microbial Control Agents 178
• Enzyme Specificity and Efficiency • Naming Enzymes Alteration of Membrane Permeability • Damage to Proteins
• Enzyme Components • Factors Influencing Enzymatic and Nucleic Acids
Activity • Feedback Inhibition • Ribozymes
Physical Methods of Microbial Control 180
Energy Production 117
Heat • Filtration • Low Temperatures • High Pressure
Oxidation-Reduction Reactions • The Generation of ATP • Desiccation • Osmotic Pressure • Radiation
• Metabolic Pathways of Energy Production
Chemical Methods of Microbial Control 185
Carbohydrate Catabolism 119
Principles of Effective Disinfection • Evaluating a Disinfectant
Glycolysis • Additional Pathways to Glycolysis • Cellular • Types of Disinfectants
Respiration • Fermentation
Lipid and Protein Catabolism 131 Microbial Characteristics and Microbial Control 194
Biochemical Tests and Bacterial Identification 131 Study Outline • Study Questions 197
Photosynthesis 133
The Light-Dependent Reactions: Photophosphorylation
• The Light-Independent Reactions: The Calvin-Benson Cycle
A Summary of Energy Production Mechanisms 135
8 Microbial Genetics 201
Structure and Function of the Genetic Material 204
Metabolic Diversity among Organisms 136 Genotype and Phenotype • DNA and Chromosomes • The Flow
Photoautotrophs • Photoheterotrophs • Chemoautotrophs of Genetic Information • DNA Replication • RNA and Protein
• Chemoheterotrophs Synthesis
Metabolic Pathways of Energy Use 140 The Regulation of Bacterial Gene Expression 214
Polysaccharide Biosynthesis • Lipid Biosynthesis • Amino Pre-transcriptional Control • Post-transcriptional Control
Acid and Protein Biosynthesis • Purine and Pyrimidine Changes in the Genetic Material 218
Biosynthesis Mutation • Types of Mutations • Mutagens • The Frequency
The Integration of Metabolism 142 of Mutation • Identifying Mutants • Identifying
Study Outline • Study Questions 144 Chemical Carcinogens
6
Genetic Transfer and Recombination 225
Transformation in Bacteria • Conjugation in Bacteria
Microbial Growth 149 • Transduction in Bacteria • Plasmids and Transposons
Genes and Evolution 233
The Requirements for Growth 150
Physical Requirements • Chemical Requirements
Biofilms 156
Culture Media 157
Chemically Defined Media • Complex Media • Anaerobic
Growth Media and Methods • Special Culture
Techniques • Selective and Differential Media • Enrichment
9 iotechnology and DNA
B
Technology 238
Introduction to Biotechnology 239
Culture
Recombinant DNA Technology • An Overview of Recombinant
Obtaining Pure Cultures 162
DNA Procedures
Preserving Bacterial Cultures 163
Tools of Biotechnology 241
The Growth of Bacterial Cultures 163 Selection • Mutation • Restriction Enzymes • Vectors
Bacterial Division • Generation Time • Logarithmic • Polymerase Chain Reaction
Representation of Bacterial Populations • Phases of Growth Techniques of Genetic Modification 244
• Direct Measurement of Microbial Growth • Estimating
Bacterial Numbers by Indirect Methods Inserting Foreign DNA into Cells • Obtaining DNA • Selecting
a Clone • Making a Gene Product
Applications of DNA Technology 250
Therapeutic Applications • Genome Projects • Scientific
Applications • Agricultural Applications

CONTENTS xv
Safety Issues and the Ethics of Using DNA Technology 258 Algae 332
Study Outline • Study Questions 260 Characteristics of Algae • Selected Phyla of Algae • Roles of Algae
in Nature
Protozoa 337
PART TWO A Survey of the Microbial World Characteristics of Protozoa • Medically Important Protozoa
10 lassification of
C Slime Molds 342
Helminths 343
Microorganisms 264 Characteristics of Helminths • Platyhelminths • Nematodes
The Study of Phylogenetic Relationships 265 Arthropods as Vectors 351
The Three Domains • A Phylogenetic Tree Study Outline • Study Questions 353
13
Classification of Organisms 269
Scientific Nomenclature • The Taxonomic Hierarchy
• Classification of Prokaryotes • Classification of Eukaryotes Viruses, Viroids, and Prions 358
• Classification of Viruses
General Characteristics of Viruses 359
Methods of Classifying and Identifying Microorganisms 272
Host Range • Viral Size
Morphological Characteristics • Differential Staining
• Biochemical Tests • Serology • Phage Typing • Fatty Acid Viral Structure 360
Profiles • Flow Cytometry • DNA Base Composition • DNA Nucleic Acid • Capsid and Envelope • General Morphology
Fingerprinting • Nucleic Acid Amplification Tests (NAATs)
• Nucleic Acid Hybridization • Putting Classification Methods
Taxonomy of Viruses 362
Together Isolation, Cultivation, and Identification of Viruses 363
Study Outline • Study Questions 286 Growing Bacteriophages in the Laboratory • Growing Animal
Viruses in the Laboratory • Viral Identification
Viral Multiplication 369
11 he Prokaryotes: Domains
T
Bacteria and Archaea 290
Multiplication of Bacteriophages • Multiplication of Animal
Viruses
Viruses and Cancer 380
The Transformation of Normal Cells into Tumor Cells • DNA
The Prokaryotic Groups 291
Oncogenic Viruses • RNA Oncogenic Viruses • Viruses
DOMAIN BACTERIA 292 to Treat Cancer
Gram-Negative Bacteria 292 Latent Viral Infections 382
Proteobacteria • The Nonproteobacteria Gram-Negative
Persistent Viral Infections 382
Bacteria
Prions 383
The Gram-Positive Bacteria 308
Plant Viruses and Viroids 383
Firmicutes (Low G + C Gram-Positive Bacteria)
• Actinobacteria (High G + C Gram-Positive Bacteria) Study Outline • Study Questions 385
DOMAIN ARCHAEA 314
Diversity within the Archaea 314 PART THREE Interaction
MICROBIAL DIVERSITY 315 between Microbe and Host
14
Discoveries Illustrating the Range of Diversity 315
rinciples of Disease
P
and Epidemiology 389
12 he Eukaryotes: Fungi, Algae,

T
Protozoa, and Helminths 319
Pathology, Infection, and Disease 390
Normal Microbiota 390
Relationships between the Normal Microbiota and the Host
• Opportunistic Microorganisms • Cooperation among
Fungi 320
Microorganisms
Characteristics of Fungi • Medically Important Fungi • Fungal
Diseases • Economic Effects of Fungi The Etiology of Infectious Diseases 394
Lichens 331 Koch’s Postulates • Exceptions to Koch’s Postulates

xvi CONTENTS
Classifying Infectious Diseases 395 SECOND LINE OF DEFENSE 446

Occurrence of a Disease • Severity or Duration of a Disease Formed Elements in Blood 446
• Extent of Host Involvement
The Lymphatic System 448
Patterns of Disease 397 Phagocytes 449
Predisposing Factors • Development of Disease
Actions of Phagocytic Cells • The Mechanism of Phagocytosis
The Spread of Infection 398 • Microbial Evasion of Phagocytosis
Reservoirs of Infection • Transmission of Disease Inflammation 452
Healthcare-Associated Infections 402 Vasodilation and Increased Permeability of Blood Vessels
Microorganisms in the Hospital • Compromised Host • Phagocyte Migration and Phagocytosis • Tissue Repair
• Chain of Transmission • Control of Healthcare-Associated Fever 455
Infections
Antimicrobial Substances 456
Emerging Infectious Diseases 405 The Complement System • Interferons • Iron-Binding Proteins
Epidemiology 407 • Antimicrobial Peptides
Descriptive Epidemiology • Analytical Epidemiology Study Outline • Study Questions 464
• Experimental Epidemiology • Case Reporting • The Centers
for Disease Control and Prevention (CDC)
17
Study Outline • Study Questions 412 daptive Immunity: Specific
A
Defenses of the Host 468
15 icrobial Mechanisms
M
of Pathogenicity 417
The Adaptive Immune System 469
Dual Nature of the Adaptive Immune System 469
Overview of Humoral Immunity • Overview of Cellular
How Microorganisms Enter a Host 418
Immunity
Portals of Entry • The Preferred Portal of Entry • Numbers
of Invading Microbes • Adherence Cytokines: Chemical Messengers of Immune Cells 470
How Bacterial Pathogens Penetrate Host Defenses 421 Antigens and Antibodies 471
Capsules • Cell Wall Components • Enzymes • Antigenic Antigens • Antibodies
Variation • Penetration into the Host Cell Cytoskeleton
Humoral Immunity Response Process 475
How Bacterial Pathogens Damage Host Cells 424 Clonal Selection of Antibody-Producing Cells • The Diversity
Using the Host’s Nutrients: Siderophores • Direct Damage of Antibodies
• Production of Toxins • Plasmids, Lysogeny, and Pathogenicity Antigen–Antibody Binding and Its Results 477
Pathogenic Properties of Viruses 430 Cellular Immunity Response Process 479
Viral Mechanisms for Evading Host Defenses • Cytopathic Antigen-Presenting Cells (APCs) • Classes of T Cells
Effects of Viruses
Extracellular Killing by the Immune System 484
Pathogenic Properties of Fungi, Protozoa, Helminths,
and Algae 432 Antibody-Dependent Cell-Mediated Cytotoxicity 484
Fungi • Protozoa • Helminths • Algae Immunological Memory 485
Portals of Exit 433 Types of Adaptive Immunity 486
Study Outline • Study Questions 435 Study Outline • Study Questions 489
16 I nnate Immunity: Nonspecific

Defenses of the Host 439 18 ractical Applications
P
of Immunology 492
The Concept of Immunity 442 Vaccines 493
FIRST LINE OF DEFENSE: SKIN AND MUCOUS Principles and Effects of Vaccination • Types of Vaccines
MEMBRANES 442 and Their Characteristics • The Development of New Vaccines
• Vaccination Technologies • Adjuvants • Safety of Vaccines
Physical Factors 442
Diagnostic Immunology 500
Chemical Factors 444
Immunologic-Based Diagnostic Tests • Monoclonal Antibodies
Normal Microbiota and Innate Immunity 445 • Precipitation Reactions • Agglutination Reactions
• Neutralization Reactions • Complement-Fixation Reactions

CONTENTS xvii
• Fluorescent-Antibody Techniques • Enzyme-Linked Tests to Guide Chemotherapy 567

Immunosorbent Assay (ELISA) • Western Blotting The Diffusion Methods • Broth Dilution Tests
(Immunoblotting) • The Future of Diagnostic and Therapeutic
Immunology Resistance to Antimicrobial Drugs 569
Mechanisms of Resistance • Antibiotic Misuse • Cost and
Study Outline • Study Questions 512 Prevention of Resistance
Antibiotic Safety 574
19
Effects of Combinations of Drugs 574
isorders Associated with
D
Future of Chemotherapeutic Agents 574
the Immune System 515 Study Outline • Study Questions 576
Hypersensitivity 516
Allergies and the Microbiome • Type I (Anaphylactic) Reactions PART FOUR Microorganisms
• Preventing Anaphylactic Reactions • Type II (Cytotoxic)
Reactions • Type III (Immune Complex) Reactions • Type IV and Human Disease
21
(Delayed Cell-Mediated) Reactions
Autoimmune Diseases 526
icrobial Diseases of
M
Cytotoxic Autoimmune Reactions • Immune Complex the Skin and Eyes 579
Autoimmune Reactions • Cell-Mediated Autoimmune Reactions
Structure and Function of the Skin 580
Reactions Related to the Human Leukocyte Antigen (HLA)
Mucous Membranes
Complex 528
Reactions to Transplantation • Immunosuppression Normal Microbiota of the Skin 580
The Immune System and Cancer 532 Microbial Diseases of the Skin 581
Immunotherapy for Cancer Bacterial Diseases of the Skin • Viral Diseases of the Skin
• Fungal Diseases of the Skin and Nails • Parasitic Infestation
Immunodeficiencies 533 of the Skin
Congenital Immunodeficiencies • Acquired Immunodeficiencies
Microbial Diseases of the Eye 599
Acquired Immunodeficiency Syndrome (AIDS) 534 Inflammation of the Eye Membranes: Conjunctivitis • Bacterial
The Origin of AIDS • HIV Infection • Diagnostic Methods Diseases of the Eye • Other Infectious Diseases of the Eye
• HIV Transmission • AIDS Worldwide • Preventing Study Outline • Study Questions 603
and Treating AIDS • The AIDS Epidemic and the Importance
of Scientific Research
22 icrobial Diseases of
M
the Nervous System 607
20 Antimicrobial Drugs 548

Structure and Function of the Nervous System 608
Bacterial Diseases of the Nervous System 609
Bacterial Meningitis • Tetanus • Botulism • Leprosy
The History of Chemotherapy 549
Antibiotic Use and Discovery Today Viral Diseases of the Nervous System 618
Poliomyelitis • Rabies • Arboviral Encephalitis
Spectrum of Antimicrobial Activity 550
The Action of Antimicrobial Drugs 551 Fungal Disease of the Nervous System 626
Inhibiting Cell Wall Synthesis • Inhibiting Protein Synthesis Cryptococcus neoformans Meningitis (Cryptococcosis)
• Injuring the Plasma Membrane • Inhibiting Nucleic Acid Protozoan Diseases of the Nervous System 627
Synthesis • Inhibiting the Synthesis of Essential Metabolites African Trypanosomiasis • Amebic Meningoencephalitis
Common Antimicrobial Drugs 554 Nervous System Diseases Caused by Prions 630
Antibacterial Antibiotics: Inhibitors of Cell Wall Synthesis Bovine Spongiform Encephalopathy and Variant
• Antimycobacterial Antibiotics • Inhibitors of Protein Synthesis Creutzfeldt-Jakob Disease
• Injury to the Plasma Membrane • Nucleic Acid Synthesis
Inhibitors • Competitive Inhibition of Essential Metabolites Disease Caused by Unidentified Agents 632
• Antifungal Drugs • Antiviral Drugs • Antiprotozoan Chronic Fatigue Syndrome
and Antihelminthic Drugs Study Outline • Study Questions 633

xviii CONTENTS
23 icrobial Diseases
M
of the Cardiovascular
and Lymphatic Systems 637
Fungal Diseases of the Lower Respiratory System 698
Histoplasmosis • Coccidioidomycosis • Pneumocystis Pneumonia
• Blastomycosis (North American Blastomycosis) • Other Fungi
Involved in Respiratory Disease
Structure and Function of the Cardiovascular and Lymphatic
Systems 638
Bacterial Diseases of the Cardiovascular and Lymphatic
Systems 639
Sepsis and Septic Shock • Bacterial Infections of the Heart
• Rheumatic Fever • Tularemia • Brucellosis (Undulant Fever)
M
the Digestive System 707
Structure and Function of the Digestive System 708
• Anthrax • Gangrene • Systemic Diseases Caused by Bites
and Scratches • Vector-Transmitted Diseases Normal Microbiota of the Digestive System 708
Viral Diseases of the Cardiovascular and Lymphatic Bacterial Diseases of the Mouth 709
Systems 655 Dental Caries (Tooth Decay) • Periodontal Disease
Burkitt’s Lymphoma • Infectious Mononucleosis • Other Diseases Bacterial Diseases of the Lower Digestive System 712
and Epstein-Barr Virus • Cytomegalovirus Infections Staphylococcal Food Poisoning (Staphylococcal Enterotoxicosis)
• Chikungunya Fever • Classic Viral Hemorrhagic Fevers • Shigellosis (Bacillary Dysentery) • Salmonellosis (Salmonella
• Emerging Viral Hemorrhagic Fevers Gastroenteritis) • Typhoid Fever • Cholera • Noncholera
Protozoan Diseases of the Cardiovascular and Lymphatic Vibrios • Escherichia coli Gastroenteritis • Campylobacter
Systems 661 Gastroenteritis • Helicobacter Peptic Ulcer Disease • Yersinia
Gastroenteritis • Clostridium perfringens Gastroenteritis
Chagas’ Disease (American Trypanosomiasis) • Toxoplasmosis
• Clostridium difficile–Associated Diarrhea • Bacillus cereus
• Malaria • Leishmaniasis • Babesiosis
Gastroenteritis
Helminthic Disease of the Cardiovascular and Lymphatic
Viral Diseases of the Digestive System 724
Systems 668
Mumps • Hepatitis • Viral Gastroenteritis
Schistosomiasis
Fungal Diseases of the Digestive System 732
Disease of Unknown Etiology 670
Kawasaki Syndrome Protozoan Diseases of the Digestive System 733
Giardiasis • Cryptosporidiosis • Cyclospora Diarrheal Infection
Study Outline • Study Questions 671 • Amebic Dysentery (Amebiasis)
Helminthic Diseases of the Digestive System 735
M Tapeworms • Hydatid Disease • Nematodes
the Respiratory System 675
26
Structure and Function of the Respiratory System 676
icrobial Diseases
M
Normal Microbiota of the Respiratory System 677
MICROBIAL DISEASES OF THE UPPER RESPIRATORY
of the Urinary
SYSTEM 677 and Reproductive
Bacterial Diseases of the Upper Respiratory System 678 Systems 746
Streptococcal Pharyngitis (Strep Throat) • Scarlet Fever
• Diphtheria • Otitis Media Structure and Function of the Urinary System 747
Viral Disease of the Upper Respiratory System 680 Structure and Function of the Reproductive Systems 747
The Common Cold Normal Microbiota of the Urinary and Reproductive
MICROBIAL DISEASES OF THE LOWER RESPIRATORY Systems 748
SYSTEM 681 DISEASES OF THE URINARY SYSTEM 749
Bacterial Diseases of the Lower Respiratory System 681 Bacterial Diseases of the Urinary System 749
Pertussis (Whooping Cough) • Tuberculosis • Bacterial Cystitis • Pyelonephritis • Leptospirosis
Pneumonias • Melioidosis DISEASES OF THE REPRODUCTIVE SYSTEMS 751
Viral Diseases of the Lower Respiratory System 694
Viral Pneumonia • Respiratory Syncytial Virus (RSV)
• Influenza (Flu)

CONTENTS xix
Bacterial Diseases of the Reproductive Systems 751

Gonorrhea • Nongonococcal Urethritis (NGU) • Pelvic
Inflammatory Disease (PID) • Syphilis • Lymphogranuloma
Venereum (LGV) • Chancroid (Soft Chancre) • Bacterial
28 pplied and Industrial
A
Microbiology 794
Vaginosis Food Microbiology 795
Foods and Disease • Industrial Food Canning • Aseptic
Viral Diseases of the Reproductive Systems 762 Packaging • Radiation and Industrial Food Preservation
Genital Herpes • Genital Warts • AIDS • High-Pressure Food Preservation • The Role of Microorganisms
Fungal Disease of the Reproductive Systems 764 in Food Production
Candidiasis Industrial Microbiology 801
Protozoan Disease of the Reproductive Systems 765 Fermentation Technology • Industrial Products • Alternative
Trichomoniasis • The TORCH Panel of Tests Energy Sources Using Microorganisms • Biofuels
• Industrial Microbiology and the Future
Study Outline • Study Questions 767 Study Outline • Study Questions 808
PART FIVE Environmental
and Applied Microbiology Answers to Knowledge and Comprehension Questions AN-1
27
Appendix A Metabolic Pathways AP-1
nvironmental
E
Appendix B Exponents, Exponential Notation,
Microbiology 771 Logarithms, and Generation Time AP-3
Microbial Diversity and Habitats 772 Appendix C Methods for Taking Clinical Samples AP-5
Symbiosis
Appendix D Pronunciation of Scientific Names AP-7
Soil Microbiology and Biogeochemical Cycles 772
The Carbon Cycle • The Nitrogen Cycle • The Sulfur Cycle Appendix E Word Roots Used in Microbiology AP-9
• Life without Sunshine • The Phosphorus Cycle
• The Degradation of Synthetic Chemicals in Soil and Water
Appendix F Classification of Prokaryotes According
to Bergey’s Manual AP-13
Aquatic Microbiology and Sewage Treatment 780
Aquatic Microorganisms • The Role of Microorganisms in Water Glossary G-1
Quality • Water Treatment • Sewage (Wastewater) Treatment
Credits C-1
Index I-1

Features
Big Picture Tough Topics Life Cycle Figures

Metabolism 108–109 Figure 11.11 Myxococcales 302
Genetics 202–203 Figure 11.15 Chlamydias 305
Immunity 440–441 Figure 12.7 The Life Cycle of Rhizopus, a Zygomycete 325
Figure 12.8 The Life Cycle of Encephalitozoon,
Big Picture Diseases a Microsporidian 326
Figure 12.9 The Life Cycle of Talaromyces, an Ascomycete 327
Human Microbiome and IBD 518–519
Figure 12.10 A Generalized Life Cycle of a Basidiomycete 328
Fungal Keratitis 600–601
Figure 12.13 Green Algae 334
Neglected Tropical Diseases 622–623
Figure 12.16 Oomycotes 336
Climate Change and Disease 658–659
Figure 12.20 The Life Cycle of Plasmodium vivax 341
Pertussis 682–683
Figure 12.22 The Generalized Life Cycle of a Cellular Slime
Cholera After Natural Disasters 720–721
Mold 344
STI Home Test Kits 752–753
Figure 12.23 The Life Cycle of a Plasmodial Slime Mold 345
Figure 12.26 The Life Cycle of the Lung Fluke,
Foundation Figures Paragonimus spp. 346
Figure 1.3 Disproving the Theory of Spontaneous Figure 12.28 The Life Cycle of the Tapeworm,
Generation 8 Echinococcus spp. 349
Figure 2.16 The Structure of DNA 44 Figure 23.13 The Life Cycle of the Tick Vector of Lyme
Figure 3.2 Microscopes and Magnification 55 Disease 652
Figure 4.6 The Structure of a Prokaryotic Cell 76 Figure 23.16 The Life Cycle of the Tick Vector (Dermacentor spp.)
of Rocky Mountain Spotted Fever 654
Figure 5.11 An Overview of Respiration and Fermentation 120
Figure 23.23 The Life Cycle of Toxoplasma gondii 663
Figure 6.15 Understanding the Bacterial Growth Curve 166
Figure 23.27 Schistosomiasis 669
Figure 7.1 Understanding the Microbial Death Curve 179
Figure 24.17 The Life Cycle of Coccidioides immitis 699
Figure 8.2 The Flow of Genetic Information 206
Figure 24.19 The Life Cycle of Pneumocystis jirovecii 700
Figure 9.1 A Typical Genetic Modification Procedure 240
Figure 25.25 The Life Cycle of Trichinella spiralis 740
Figure 10.1 Three-Domain System 266
Figure 12.1 Exploring Pathogenic Eukaryotes 320
Clinical Focus
Figure 13.15 Replication of a DNA-Containing Animal
Virus 375 Human Tuberculosis—Dallas, Texas 139
Figure 14.3 Koch’s Postulates: Understanding Disease 395 Infection Following Anesthesia Injection 193
Figure 15.4 Mechanisms of Exotoxins and Endotoxins 425 Tracking West Nile Virus 215
Figure 15.9 Microbial Mechanisms of Pathogenicity 434 Norovirus—Who Is Responsible for the Outbreak? 259
Figure 16.8 The Phases of Phagocytosis 451 The Most Frequent Cause of Recreational Waterborne
Diarrhea 347
Figure 16.12 Outcomes of Complement Activation 459
Influenza: Crossing the Species Barrier 364
Figure 17.20 The Dual Nature of the Adaptive Immune
System 488 Healthcare-Associated Infections 411
Figure 18.2 The Production of Monoclonal Antibodies 502 A World Health Problem 498
Figure 19.16 The Progression of HIV Infection 538 A Delayed Rash 527
Figure 20.2 Major Action Modes of Antimicrobial Drugs 551 Antibiotics in Animal Feed Linked to Human Disease 573
Figure 20.20 Bacterial Resistance to Antibiotics 570 Infections in the Gym 588
xx

xxi
Features
A Neurological Disease 625 21.3 Patchy Redness and Pimple-Like Conditions 587
A Sick Child 645 21.4 Microbial Diseases of the Eye 599
Outbreak 694 22.1 Meningitis and Encephalitis 615
A Foodborne Infection 717 22.2 Types of Arboviral Encephalitis 628
Survival of the Fittest 756 22.3 Microbial Diseases with Neurological Symptoms
or Paralysis 632
Applications of Microbiology 23.1 Human-Reservoir Infections 643
23.2 Infections from Animal Reservoirs Transmitted by Direct
Designer Jeans: Made by Microbes? 3
Contact 649
Bioremediation—Bacteria Clean Up Pollution 31
23.3 Infections Transmitted by Vectors 650
What is that Slime? 54
23.4 Viral Hemorrhagic Fevers 662
Why Microbiologists Study Termites 94
23.5 Infections Transmitted by Soil and Water 668
Life in the Extreme 153
24.1 Microbial Diseases of the Upper Respiratory
Mass Deaths of Marine Mammals Spur Veterinary System 681
Microbiology 275
24.2 Common Bacterial Pneumonias 691
Bacteria and Insect Sex 297
24.3 Microbial Diseases of the Lower Respiratory
Streptococcus: Harmful or Helpful? 422 System 702
Serum Collection 462 25.1 Bacterial Diseases of the Mouth 712
Interleukin-12: The Next “Magic Bullet”? 471 25.2 Bacterial Diseases of the Lower Digestive System 726
Protection against Bioterrorism 648 25.3 Characteristics of Viral Hepatitis 728
A Safe Blood Supply 730 25.4 Viral Diseases of the Digestive System 733
Biosensors: Bacteria That Detect Pollutants and Pathogens 783 25.5 Fungal, Protozoan, and Helminthic Diseases of the Lower
From Plant Disease to Shampoo and Salad Dressing 801 Digestive System 737
26.1 Bacterial Diseases of the Urinary System 750
Diseases in Focus 26.2 Characteristics of the Most Common Types of Vaginitis
and Vaginosis 764
21.1 Macular Rashes 584
26.3 Microbial Diseases of the Reproductive Systems 766
21.2 Vesicular and Pustular Rashes 586

ASM Recommended Curriculum Guidelines
for Undergraduate Microbiology
The American Society for Microbiology (ASM) endorses a Metabolic Pathways:
concept-based curriculum for introductory microbiology, empha- ● Bacteria and Archaea exhibit extensive, and often unique,
sizing skills and concepts that remain important long after students
metabolic diversity (e.g. nitrogen fixation, methane
exit the course. The ASM Curriculum Guidelines for Undergraduate
production, anoxygenic photosynthesis).
Microbiology Education provide a framework for key microbio-
logical topics and agree with scientific literacy reports from the
● Interactions of microorganisms among themselves and with
American Association for the Advancement of Science and How- their environment are determined by their metabolic abilities
ard Hughes Medical Institute. This textbook references part one (e.g., quorum sensing, oxygen consumption, nitrogen
of curriculum guidelines throughout chapters. When a discussion transformations).
touches on one of the concepts, readers ● Survival and growth of any microorganism in a given
will see the ASM icon, along with a ASM: environment depends on its metabolic characteristics.
summary of the relevant statement. ● Growth of microorganisms can be controlled by physical,
chemical, mechanical, or biological means.
ASM Guideline Concepts
Information Flow and Genetics:
and Statements ● Genetic variations can impact microbial functions (e.g. in
Evolution: biofilm formation, pathogenicity, and drug resistance).
● Cells, organelles (e.g., mitochondria and chloroplasts), and all ● Although the central dogma is universal in all cells, the
major metabolic pathways evolved from early prokaryotic cells. processes of replication, transcription, and translation differ
● Mutations and horizontal gene transfer, with the immense in Bacteria, Archaea, and Eukaryotes.
variety of microenvironments, have selected for a huge ● Regulation of gene expression is influenced by external and
diversity of microorganisms. internal molecular cues and/or signals.
● Human impact on the environment influences the evolution ● Synthesis of viral genetic material and proteins is dependent
of microorganisms (e.g., emerging diseases and the selection on host cells.
of antibiotic resistance). ● Cell genomes can be manipulated to alter cell function.
● Traditional concept of species is not readily applicable to
microbes due to asexual reproduction and the frequent Microbial Systems:
occurrence of horizontal gene transfer. ● Microorganisms are ubiquitous and live in diverse and
● Evolutionary relatedness of organisms is best reflected in dynamic ecosystems.
phylogenetic trees. ● Most bacteria in nature live in biofilm communities.
● Microorganisms and their environment interact with and
Cell Structure and Function: modify each other.
● Structure and function of microorganisms have been ● Microorganisms, cellular and viral, can interact with both
revealed by the use of microscopy (including brightfield, human and nonhuman hosts in beneficial, neutral, or
phase contrast, fluorescent, and electron). detrimental ways.
● Bacteria have unique cell structures that can be targets for
antibiotics, immunity, and phage infection. Impact of Microorganisms:
● Bacteria and Archaea have specialized structures (e.g. flagella, ● Microbes are essential for life as we know it and the processes
endospores, and pili) that often confer critical capabilities. that support life (e.g., in biogeochemical cycles and plant
● While microscopic eukaryotes (for example, fungi, protozoa, and/or animal microbiota).
and algae) carry out some of the same processes as bacteria, ● Microorganisms provide essential models that give us
many of the cellular properties are fundamentally different. fundamental knowledge about life processes.
● Replication cycles of viruses (lytic and lysogenic) differ ● Humans utilize and harness microorganisms and their products.
among viruses and are determined by their unique structures ● Because the true diversity of microbial life is largely unknown,
and genomes. its effects and potential benefits have not been fully explored.
xxii

In the Clinic
As the nurse practitioner in a rural
hospital, you are reviewing a microscope
slide of a skin scraping from a 12-year-
old girl. The slide shows branched,
intertwined nucleated hyphae. The girl
has dry, scaly, itchy patches on her arms.
What is causing her skin problem?
Hint: Read about types of microorganisms (pages 3–5).
Note: Answers to In the Clinic questions are found online at MasteringMicrobiology.
1T
The Microbial World and You
he overall theme of this textbook is the relationship between microbes—

very small organisms that usually require a microscope to be seen—and
our lives. This relationship involves not only the familiar harmful effects
of certain microorganisms, such as disease and food spoilage, but also their
many beneficial effects. In this chapter we introduce you to some of the many
ways microbes affect our lives. We begin by discussing how organisms are
named and classified, followed by a short history of microbiology that reveals
how much we have learned in just a few hundred years. Then we discuss the
incredible diversity of microorganisms and their ecological importance, noting
how they maintain balance in the environment
by recycling chemical elements such as carbon ASM: Microorganisms provide essential
and nitrogen among the soil, organisms, and the models that give us fundamental
knowledge about life processes.
atmosphere. We also examine how microbes are
used in commercial and industrial applications
to produce foods, chemicals, and drugs (such as antibiotics); and to treat sewage,
control pests, and clean up pollutants. We will discuss microbes as the cause of
such diseases as avian (bird) flu, West Nile encephalitis, mad cow disease, diarrhea,
hemorrhagic fever, and AIDS, and we examine the growing public health problem
of antibiotic-resistant bacteria.
Staphylococcus aureus (STAF-i-lō-kok’kus OR-ē-us) bacteria on human nasal
epithelial cells are shown in the photograph. These bacteria live harmlessly on skin
or inside the nose. Misuse of antibiotics allows the survival of bacteria with antibiotic-
resistance genes, such as methicillin-resistant S. aureus (MRSA). As illustrated in the
Clinical Case, an infection caused by these bacteria is resistant to antibiotic treatment.
1
Staphylococcus aureus bacteria on human nasal epithelial cells.
M01_TORT9150_12_CH01_pp001-023.indd 1 28/10/14 5:39 PM

2 Part one Fundamentals of Microbiology
Microbes in Our Lives for medicine and the related health sciences. For example,
hospital workers must be able to protect patients from common
Learning Objective microbes that are normally harmless but pose a threat to the sick
1-1 List several ways in which microbes affect our lives. and injured.
Today we understand that microorganisms are found almost
For many people, the words germ and microbe bring to mind a
everywhere. Yet not long ago, before the invention of the micro-
group of tiny creatures that do not quite fit into any of the cat-
scope, microbes were unknown to scientists. Thousands of peo-
egories in that old question, “Is it animal, vegetable, or mineral?”
ple died in devastating epidemics, the causes and transmission
Microbes, also called microorganisms, are minute living things
of which were not understood. Entire families died because vac-
that individually are usually too small to be seen with the un-
cinations and antibiotics were not available to fight infections.
aided eye. The group includes bacteria, fungi (yeasts and molds),
We can get an idea of how our current concepts of microbi-
protozoa, and microscopic algae. It also includes viruses, those
ology developed by looking at a few historic milestones in mi-
noncellular entities sometimes regarded as straddling the border
crobiology that have changed our lives. First, however, we will
between life and nonlife (Chapters 11, 12, and 13, respectively).
look at the major groups of microbes and how they are named
We tend to associate these small organisms only with un-
and classified.
comfortable infections, with common inconveniences such as
spoiled food, or with major diseases such as AIDS. However, the Check Your Understanding
majority of microorganisms actually help maintain the balance ✓ Describe some of the destructive and beneficial actions of
of life in our environment. Marine and freshwater microorgan- microbes. 1-1*
isms form the basis of the food chain in oceans, lakes, and rivers.
Soil microbes help break down wastes and incorporate nitrogen
gas from the air into organic compounds, thereby recycling
Naming and Classifying
chemical elements among soil, water, living organisms, and air. Microorganisms
Certain microbes play important roles in photosynthesis, a food-
and oxygen-generating process that is critical to life on Earth. Learning Objectives
Humans and many other animals depend on the microbes in 1-2 Recognize the system of scientific nomenclature that uses
their intestines for digestion and the synthesis of some vitamins two names: a genus and a specific epithet.
that their bodies require, including some B vitamins for metab- 1-3 Differentiate the major characteristics of each group of
olism and vitamin K for blood clotting. microorganisms.
Microorganisms also have many commercial applications. 1-4 List the three domains.
They are used in the synthesis of such chemical products as
vitamins, organic acids, enzymes, alcohols, and many drugs.
For example, microbes are used to produce acetone and buta- Nomenclature
nol, and the vitamins B2 (riboflavin) and B12 (cobalamin) are The system of nomenclature (naming) for organisms in use
made biochemically. The process by which microbes produce today was established in 1735 by Carolus Linnaeus. Scientific
acetone and butanol was discovered in 1914 by Chaim Weiz- names are latinized because Latin was the language tradition-
mann, a Russian-born chemist working in England. With the ally used by scholars. Scientific nomenclature assigns each or-
outbreak of World War I in August of that year, the produc- ganism two names—the genus (plural: genera) is the first name
tion of acetone became very important for making cordite (a and is always capitalized; the specific epithet (species name)
smokeless form of gunpowder used in munitions). Weizmann’s follows and is not capitalized. The organism is referred to by
discovery played a significant role in determining the outcome both the genus and the specific epithet, and both names are
of the war. underlined or italicized. By custom, after a scientific name has
The food industry also uses microbes in producing, for been mentioned once, it can be abbreviated with the initial of
example, vinegar, sauerkraut, pickles, soy sauce, cheese, yogurt, the genus followed by the specific epithet.
bread, and alcoholic beverages. In addition, enzymes from mi- Scientific names can, among other things, describe an or-
crobes can now be manipulated to cause the microbes to produce ganism, honor a researcher, or identify the habitat of a species.
substances they normally don’t synthesize, including cellulose, For example, consider Staphylococcus aureus, a bacterium com-
digestive aids, and drain cleaner, plus important therapeutic monly found on human skin. Staphylo- describes the clustered
substances such as insulin. Microbial enzymes may even have arrangement of the cells; -coccus indicates that they are shaped
helped produce your favorite pair of jeans (see the Applications like spheres. The specific epithet, aureus, is Latin for golden,
of Microbiology box).
Though only a minority of microorganisms are pathogenic *The numbers following Check Your Understanding questions refer to the corre-
(disease-producing), practical knowledge of microbes is necessary sponding Learning Objectives.

Chapter 1 The Microbial World and You 3
Applications of Microbiology
Designer Jeans: Made by Microbes?

Denim blue jeans have been popular ever Bleaching
since Levi Strauss and Jacob Davis first made Peroxide is a safer bleaching agent than
them for California gold miners in 1873. chlorine and can be easily removed from fabric
Now, companies that manufacture blue and wastewater by enzymes. Researchers at
jeans are turning to microbiology to develop Novo Nordisk Biotech cloned a mushroom
environmentally sound production methods that peroxidase gene in yeast and grew the yeasts
minimize toxic wastes and the associated costs. in washing machine conditions. The yeast that
survived the washing machine were selected as E. coli bacteria produce indigo
Soft, Faded Jeans from tryptophan.
the peroxidase producers.
A softer, faded denim is made with enzymes
called cellulases from Trichoderma fungus. They Indigo
digest some of the cellulose in the cotton. Chemical synthesis of indigo requires a high pH a biodegradable alternative to conventional
Unlike many chemical reactions, enzymes and produces waste that explodes on contact with plastic, which is made from petroleum.
usually operate at safe temperatures and pH. air. However, a California biotechnology company,
Moreover, enzymes are proteins, so they are Genencor, has developed a method to produce
readily degraded for removal from wastewater. indigo by using bacteria. Researchers identified a
gene from a soil bacterium, Pseudomonas putida,
Fabric
that converts the bacterial by-product indole
Cotton production requires large tracts of land,
to indigo. This gene was put into Escherichia coli
pesticides, and fertilizer, and the crop yield
bacteria, which then turned blue.
depends on the weather. However, bacteria
can produce both cotton and polyester with Bioplastic
less environmental impact. Gluconacetobacter Microbes can even make plastic zippers and
xylinus bacteria make cellulose by attaching packaging material for the jeans. Over 25
glucose units to simple chains in the outer bacteria make polyhydroxyalkanoate (PHA)
membrane of the bacterial cell wall. The inclusion granules as a food reserve. PHAs are
cellulose microfibrils are extruded through similar to common plastics, and because they are Indigo-producing E. coli 0.3 μ m
pores in the outer membrane, and bundles of made by bacteria, they are also readily bacteria.
microfibrils then twist into ribbons. degraded by many bacteria. PHAs could provide TEM
the color of many colonies of this bacterium. The genus of the

bacterium Escherichia coli (eshʹer-IK-ē-ah KŌ-lī, or KŌ-lē) is Clinical Case: A Simple Spider Bite?
named for a scientist, Theodor Escherich, whereas its specific
epithet, coli, reminds us that E. coli live in the colon, or large Andrea is a normally healthy 22-year-old college student who
intestine. Table 1.1 contains more examples. lives at home with her mother and younger sister, a high
school gymnast. She is trying to work on a paper for her
Check Your Understanding psychology class but is having a hard time because a red,
✓ Distinguish a genus from a specific epithet. 1-2 swollen sore on her right wrist is making typing difficult. “Why
won’t this spider bite heal?” she wonders. “It’s been there for
days!” She makes an appointment with her doctor so she can
Types of Microorganisms
show him the painful lesion. Although Andrea does not have
Here is an overview of the main types of microorganisms. (The a fever, she does have an elevated white blood cell count that
classification and identification of microorganisms are discussed indicates a bacterial infection. Andrea’s doctor suspects that
in Chapter 10.) this isn’t a spider bite at all, but a staph infection. He prescribes
a β-lactam antibiotic, cephalosporin. Learn more about the
Bacteria development of Andrea’s illness on the following pages.
Bacteria (singular: bacterium) are relatively simple, single-celled What is staph? Read on to find out.
(unicellular) organisms. Because their genetic material is not
enclosed in a special nuclear membrane, bacterial cells are called
prokaryotes (prō-KAR-e-ōts), from Greek words meaning pre- 3 15 17 18 19
nucleus. Prokaryotes include both bacteria and archaea. ▲

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DANCE ON STILTS AT THE GIRLS’ UNYAGO, NIUCHI
Newala, too, suffers from the distance of its water-supply—at least

the Newala of to-day does; there was once another Newala in a lovely
valley at the foot of the plateau. I visited it and found scarcely a trace
of houses, only a Christian cemetery, with the graves of several
missionaries and their converts, remaining as a monument of its
former glories. But the surroundings are wonderfully beautiful. A
thick grove of splendid mango-trees closes in the weather-worn
crosses and headstones; behind them, combining the useful and the
agreeable, is a whole plantation of lemon-trees covered with ripe
fruit; not the small African kind, but a much larger and also juicier
imported variety, which drops into the hands of the passing traveller,
without calling for any exertion on his part. Old Newala is now under
the jurisdiction of the native pastor, Daudi, at Chingulungulu, who,
as I am on very friendly terms with him, allows me, as a matter of
course, the use of this lemon-grove during my stay at Newala.
FEET MUTILATED BY THE RAVAGES OF THE “JIGGER”
(Sarcopsylla penetrans)
The water-supply of New Newala is in the bottom of the valley,

some 1,600 feet lower down. The way is not only long and fatiguing,
but the water, when we get it, is thoroughly bad. We are suffering not
only from this, but from the fact that the arrangements at Newala are
nothing short of luxurious. We have a separate kitchen—a hut built
against the boma palisade on the right of the baraza, the interior of
which is not visible from our usual position. Our two cooks were not
long in finding this out, and they consequently do—or rather neglect
to do—what they please. In any case they do not seem to be very
particular about the boiling of our drinking-water—at least I can
attribute to no other cause certain attacks of a dysenteric nature,
from which both Knudsen and I have suffered for some time. If a
man like Omari has to be left unwatched for a moment, he is capable
of anything. Besides this complaint, we are inconvenienced by the
state of our nails, which have become as hard as glass, and crack on
the slightest provocation, and I have the additional infliction of
pimples all over me. As if all this were not enough, we have also, for
the last week been waging war against the jigger, who has found his
Eldorado in the hot sand of the Makonde plateau. Our men are seen
all day long—whenever their chronic colds and the dysentery likewise
raging among them permit—occupied in removing this scourge of
Africa from their feet and trying to prevent the disastrous
consequences of its presence. It is quite common to see natives of
this place with one or two toes missing; many have lost all their toes,
or even the whole front part of the foot, so that a well-formed leg
ends in a shapeless stump. These ravages are caused by the female of
Sarcopsylla penetrans, which bores its way under the skin and there
develops an egg-sac the size of a pea. In all books on the subject, it is
stated that one’s attention is called to the presence of this parasite by
an intolerable itching. This agrees very well with my experience, so
far as the softer parts of the sole, the spaces between and under the
toes, and the side of the foot are concerned, but if the creature
penetrates through the harder parts of the heel or ball of the foot, it
may escape even the most careful search till it has reached maturity.
Then there is no time to be lost, if the horrible ulceration, of which
we see cases by the dozen every day, is to be prevented. It is much
easier, by the way, to discover the insect on the white skin of a
European than on that of a native, on which the dark speck scarcely
shows. The four or five jiggers which, in spite of the fact that I
constantly wore high laced boots, chose my feet to settle in, were
taken out for me by the all-accomplished Knudsen, after which I
thought it advisable to wash out the cavities with corrosive
sublimate. The natives have a different sort of disinfectant—they fill
the hole with scraped roots. In a tiny Makua village on the slope of
the plateau south of Newala, we saw an old woman who had filled all
the spaces under her toe-nails with powdered roots by way of
prophylactic treatment. What will be the result, if any, who can say?
The rest of the many trifling ills which trouble our existence are
really more comic than serious. In the absence of anything else to
smoke, Knudsen and I at last opened a box of cigars procured from
the Indian store-keeper at Lindi, and tried them, with the most
distressing results. Whether they contain opium or some other
narcotic, neither of us can say, but after the tenth puff we were both
“off,” three-quarters stupefied and unspeakably wretched. Slowly we
recovered—and what happened next? Half-an-hour later we were
once more smoking these poisonous concoctions—so insatiable is the
craving for tobacco in the tropics.
Even my present attacks of fever scarcely deserve to be taken
seriously. I have had no less than three here at Newala, all of which
have run their course in an incredibly short time. In the early
afternoon, I am busy with my old natives, asking questions and
making notes. The strong midday coffee has stimulated my spirits to
an extraordinary degree, the brain is active and vigorous, and work
progresses rapidly, while a pleasant warmth pervades the whole
body. Suddenly this gives place to a violent chill, forcing me to put on
my overcoat, though it is only half-past three and the afternoon sun
is at its hottest. Now the brain no longer works with such acuteness
and logical precision; more especially does it fail me in trying to
establish the syntax of the difficult Makua language on which I have
ventured, as if I had not enough to do without it. Under the
circumstances it seems advisable to take my temperature, and I do
so, to save trouble, without leaving my seat, and while going on with
my work. On examination, I find it to be 101·48°. My tutors are
abruptly dismissed and my bed set up in the baraza; a few minutes
later I am in it and treating myself internally with hot water and
lemon-juice.
Three hours later, the thermometer marks nearly 104°, and I make
them carry me back into the tent, bed and all, as I am now perspiring
heavily, and exposure to the cold wind just beginning to blow might
mean a fatal chill. I lie still for a little while, and then find, to my
great relief, that the temperature is not rising, but rather falling. This
is about 7.30 p.m. At 8 p.m. I find, to my unbounded astonishment,
that it has fallen below 98·6°, and I feel perfectly well. I read for an
hour or two, and could very well enjoy a smoke, if I had the
wherewithal—Indian cigars being out of the question.
Having no medical training, I am at a loss to account for this state
of things. It is impossible that these transitory attacks of high fever
should be malarial; it seems more probable that they are due to a
kind of sunstroke. On consulting my note-book, I become more and
more inclined to think this is the case, for these attacks regularly
follow extreme fatigue and long exposure to strong sunshine. They at
least have the advantage of being only short interruptions to my
work, as on the following morning I am always quite fresh and fit.
My treasure of a cook is suffering from an enormous hydrocele which
makes it difficult for him to get up, and Moritz is obliged to keep in
the dark on account of his inflamed eyes. Knudsen’s cook, a raw boy
from somewhere in the bush, knows still less of cooking than Omari;
consequently Nils Knudsen himself has been promoted to the vacant
post. Finding that we had come to the end of our supplies, he began
by sending to Chingulungulu for the four sucking-pigs which we had
bought from Matola and temporarily left in his charge; and when
they came up, neatly packed in a large crate, he callously slaughtered
the biggest of them. The first joint we were thoughtless enough to
entrust for roasting to Knudsen’s mshenzi cook, and it was
consequently uneatable; but we made the rest of the animal into a
jelly which we ate with great relish after weeks of underfeeding,
consuming incredible helpings of it at both midday and evening
meals. The only drawback is a certain want of variety in the tinned
vegetables. Dr. Jäger, to whom the Geographical Commission
entrusted the provisioning of the expeditions—mine as well as his
own—because he had more time on his hands than the rest of us,
seems to have laid in a huge stock of Teltow turnips,[46] an article of
food which is all very well for occasional use, but which quickly palls
when set before one every day; and we seem to have no other tins
left. There is no help for it—we must put up with the turnips; but I
am certain that, once I am home again, I shall not touch them for ten
years to come.
Amid all these minor evils, which, after all, go to make up the
genuine flavour of Africa, there is at least one cheering touch:
Knudsen has, with the dexterity of a skilled mechanic, repaired my 9
× 12 cm. camera, at least so far that I can use it with a little care.
How, in the absence of finger-nails, he was able to accomplish such a
ticklish piece of work, having no tool but a clumsy screw-driver for
taking to pieces and putting together again the complicated
mechanism of the instantaneous shutter, is still a mystery to me; but
he did it successfully. The loss of his finger-nails shows him in a light
contrasting curiously enough with the intelligence evinced by the
above operation; though, after all, it is scarcely surprising after his
ten years’ residence in the bush. One day, at Lindi, he had occasion
to wash a dog, which must have been in need of very thorough
cleansing, for the bottle handed to our friend for the purpose had an
extremely strong smell. Having performed his task in the most
conscientious manner, he perceived with some surprise that the dog
did not appear much the better for it, and was further surprised by
finding his own nails ulcerating away in the course of the next few
days. “How was I to know that carbolic acid has to be diluted?” he
mutters indignantly, from time to time, with a troubled gaze at his
mutilated finger-tips.
Since we came to Newala we have been making excursions in all
directions through the surrounding country, in accordance with old
habit, and also because the akida Sefu did not get together the tribal
elders from whom I wanted information so speedily as he had
promised. There is, however, no harm done, as, even if seen only
from the outside, the country and people are interesting enough.
The Makonde plateau is like a large rectangular table rounded off
at the corners. Measured from the Indian Ocean to Newala, it is
about seventy-five miles long, and between the Rovuma and the
Lukuledi it averages fifty miles in breadth, so that its superficial area
is about two-thirds of that of the kingdom of Saxony. The surface,
however, is not level, but uniformly inclined from its south-western
edge to the ocean. From the upper edge, on which Newala lies, the
eye ranges for many miles east and north-east, without encountering
any obstacle, over the Makonde bush. It is a green sea, from which
here and there thick clouds of smoke rise, to show that it, too, is
inhabited by men who carry on their tillage like so many other
primitive peoples, by cutting down and burning the bush, and
manuring with the ashes. Even in the radiant light of a tropical day
such a fire is a grand sight.
Much less effective is the impression produced just now by the
great western plain as seen from the edge of the plateau. As often as
time permits, I stroll along this edge, sometimes in one direction,
sometimes in another, in the hope of finding the air clear enough to
let me enjoy the view; but I have always been disappointed.
Wherever one looks, clouds of smoke rise from the burning bush,
and the air is full of smoke and vapour. It is a pity, for under more
favourable circumstances the panorama of the whole country up to
the distant Majeje hills must be truly magnificent. It is of little use
taking photographs now, and an outline sketch gives a very poor idea
of the scenery. In one of these excursions I went out of my way to
make a personal attempt on the Makonde bush. The present edge of
the plateau is the result of a far-reaching process of destruction
through erosion and denudation. The Makonde strata are
everywhere cut into by ravines, which, though short, are hundreds of
yards in depth. In consequence of the loose stratification of these
beds, not only are the walls of these ravines nearly vertical, but their
upper end is closed by an equally steep escarpment, so that the
western edge of the Makonde plateau is hemmed in by a series of
deep, basin-like valleys. In order to get from one side of such a ravine
to the other, I cut my way through the bush with a dozen of my men.
It was a very open part, with more grass than scrub, but even so the
short stretch of less than two hundred yards was very hard work; at
the end of it the men’s calicoes were in rags and they themselves
bleeding from hundreds of scratches, while even our strong khaki
suits had not escaped scatheless.
NATIVE PATH THROUGH THE MAKONDE BUSH, NEAR

MAHUTA
I see increasing reason to believe that the view formed some time
back as to the origin of the Makonde bush is the correct one. I have
no doubt that it is not a natural product, but the result of human
occupation. Those parts of the high country where man—as a very
slight amount of practice enables the eye to perceive at once—has not
yet penetrated with axe and hoe, are still occupied by a splendid
timber forest quite able to sustain a comparison with our mixed
forests in Germany. But wherever man has once built his hut or tilled
his field, this horrible bush springs up. Every phase of this process
may be seen in the course of a couple of hours’ walk along the main
road. From the bush to right or left, one hears the sound of the axe—
not from one spot only, but from several directions at once. A few
steps further on, we can see what is taking place. The brush has been
cut down and piled up in heaps to the height of a yard or more,
between which the trunks of the large trees stand up like the last
pillars of a magnificent ruined building. These, too, present a
melancholy spectacle: the destructive Makonde have ringed them—
cut a broad strip of bark all round to ensure their dying off—and also
piled up pyramids of brush round them. Father and son, mother and
son-in-law, are chopping away perseveringly in the background—too
busy, almost, to look round at the white stranger, who usually excites
so much interest. If you pass by the same place a week later, the piles
of brushwood have disappeared and a thick layer of ashes has taken
the place of the green forest. The large trees stretch their
smouldering trunks and branches in dumb accusation to heaven—if
they have not already fallen and been more or less reduced to ashes,
perhaps only showing as a white stripe on the dark ground.
This work of destruction is carried out by the Makonde alike on the
virgin forest and on the bush which has sprung up on sites already
cultivated and deserted. In the second case they are saved the trouble
of burning the large trees, these being entirely absent in the
secondary bush.
After burning this piece of forest ground and loosening it with the
hoe, the native sows his corn and plants his vegetables. All over the
country, he goes in for bed-culture, which requires, and, in fact,
receives, the most careful attention. Weeds are nowhere tolerated in
the south of German East Africa. The crops may fail on the plains,
where droughts are frequent, but never on the plateau with its
abundant rains and heavy dews. Its fortunate inhabitants even have
the satisfaction of seeing the proud Wayao and Wamakua working
for them as labourers, driven by hunger to serve where they were
accustomed to rule.
But the light, sandy soil is soon exhausted, and would yield no
harvest the second year if cultivated twice running. This fact has
been familiar to the native for ages; consequently he provides in
time, and, while his crop is growing, prepares the next plot with axe
and firebrand. Next year he plants this with his various crops and
lets the first piece lie fallow. For a short time it remains waste and
desolate; then nature steps in to repair the destruction wrought by
man; a thousand new growths spring out of the exhausted soil, and
even the old stumps put forth fresh shoots. Next year the new growth
is up to one’s knees, and in a few years more it is that terrible,
impenetrable bush, which maintains its position till the black
occupier of the land has made the round of all the available sites and
come back to his starting point.
The Makonde are, body and soul, so to speak, one with this bush.
According to my Yao informants, indeed, their name means nothing
else but “bush people.” Their own tradition says that they have been
settled up here for a very long time, but to my surprise they laid great
stress on an original immigration. Their old homes were in the
south-east, near Mikindani and the mouth of the Rovuma, whence
their peaceful forefathers were driven by the continual raids of the
Sakalavas from Madagascar and the warlike Shirazis[47] of the coast,
to take refuge on the almost inaccessible plateau. I have studied
African ethnology for twenty years, but the fact that changes of
population in this apparently quiet and peaceable corner of the earth
could have been occasioned by outside enterprises taking place on
the high seas, was completely new to me. It is, no doubt, however,
correct.
The charming tribal legend of the Makonde—besides informing us
of other interesting matters—explains why they have to live in the
thickest of the bush and a long way from the edge of the plateau,
instead of making their permanent homes beside the purling brooks
and springs of the low country.
“The place where the tribe originated is Mahuta, on the southern
side of the plateau towards the Rovuma, where of old time there was
nothing but thick bush. Out of this bush came a man who never
washed himself or shaved his head, and who ate and drank but little.
He went out and made a human figure from the wood of a tree
growing in the open country, which he took home to his abode in the
bush and there set it upright. In the night this image came to life and
was a woman. The man and woman went down together to the
Rovuma to wash themselves. Here the woman gave birth to a still-
born child. They left that place and passed over the high land into the
valley of the Mbemkuru, where the woman had another child, which
was also born dead. Then they returned to the high bush country of
Mahuta, where the third child was born, which lived and grew up. In
course of time, the couple had many more children, and called
themselves Wamatanda. These were the ancestral stock of the
Makonde, also called Wamakonde,[48] i.e., aborigines. Their
forefather, the man from the bush, gave his children the command to
bury their dead upright, in memory of the mother of their race who
was cut out of wood and awoke to life when standing upright. He also
warned them against settling in the valleys and near large streams,
for sickness and death dwelt there. They were to make it a rule to
have their huts at least an hour’s walk from the nearest watering-
place; then their children would thrive and escape illness.”
The explanation of the name Makonde given by my informants is
somewhat different from that contained in the above legend, which I
extract from a little book (small, but packed with information), by
Pater Adams, entitled Lindi und sein Hinterland. Otherwise, my
results agree exactly with the statements of the legend. Washing?
Hapana—there is no such thing. Why should they do so? As it is, the
supply of water scarcely suffices for cooking and drinking; other
people do not wash, so why should the Makonde distinguish himself
by such needless eccentricity? As for shaving the head, the short,
woolly crop scarcely needs it,[49] so the second ancestral precept is
likewise easy enough to follow. Beyond this, however, there is
nothing ridiculous in the ancestor’s advice. I have obtained from
various local artists a fairly large number of figures carved in wood,
ranging from fifteen to twenty-three inches in height, and
representing women belonging to the great group of the Mavia,
Makonde, and Matambwe tribes. The carving is remarkably well
done and renders the female type with great accuracy, especially the
keloid ornamentation, to be described later on. As to the object and
meaning of their works the sculptors either could or (more probably)
would tell me nothing, and I was forced to content myself with the
scanty information vouchsafed by one man, who said that the figures
were merely intended to represent the nembo—the artificial
deformations of pelele, ear-discs, and keloids. The legend recorded
by Pater Adams places these figures in a new light. They must surely
be more than mere dolls; and we may even venture to assume that
they are—though the majority of present-day Makonde are probably
unaware of the fact—representations of the tribal ancestress.
The references in the legend to the descent from Mahuta to the
Rovuma, and to a journey across the highlands into the Mbekuru
valley, undoubtedly indicate the previous history of the tribe, the
travels of the ancestral pair typifying the migrations of their
descendants. The descent to the neighbouring Rovuma valley, with
its extraordinary fertility and great abundance of game, is intelligible
at a glance—but the crossing of the Lukuledi depression, the ascent
to the Rondo Plateau and the descent to the Mbemkuru, also lie
within the bounds of probability, for all these districts have exactly
the same character as the extreme south. Now, however, comes a
point of especial interest for our bacteriological age. The primitive
Makonde did not enjoy their lives in the marshy river-valleys.
Disease raged among them, and many died. It was only after they
had returned to their original home near Mahuta, that the health
conditions of these people improved. We are very apt to think of the
African as a stupid person whose ignorance of nature is only equalled
by his fear of it, and who looks on all mishaps as caused by evil
spirits and malignant natural powers. It is much more correct to
assume in this case that the people very early learnt to distinguish
districts infested with malaria from those where it is absent.
This knowledge is crystallized in the
ancestral warning against settling in the
valleys and near the great waters, the
dwelling-places of disease and death. At the
same time, for security against the hostile
Mavia south of the Rovuma, it was enacted
that every settlement must be not less than a
certain distance from the southern edge of the
plateau. Such in fact is their mode of life at the
present day. It is not such a bad one, and
certainly they are both safer and more
comfortable than the Makua, the recent
intruders from the south, who have made USUAL METHOD OF
good their footing on the western edge of the CLOSING HUT-DOOR
plateau, extending over a fairly wide belt of
country. Neither Makua nor Makonde show in their dwellings
anything of the size and comeliness of the Yao houses in the plain,
especially at Masasi, Chingulungulu and Zuza’s. Jumbe Chauro, a
Makonde hamlet not far from Newala, on the road to Mahuta, is the
most important settlement of the tribe I have yet seen, and has fairly
spacious huts. But how slovenly is their construction compared with
the palatial residences of the elephant-hunters living in the plain.
The roofs are still more untidy than in the general run of huts during
the dry season, the walls show here and there the scanty beginnings
or the lamentable remains of the mud plastering, and the interior is a
veritable dog-kennel; dirt, dust and disorder everywhere. A few huts
only show any attempt at division into rooms, and this consists
merely of very roughly-made bamboo partitions. In one point alone
have I noticed any indication of progress—in the method of fastening
the door. Houses all over the south are secured in a simple but
ingenious manner. The door consists of a set of stout pieces of wood
or bamboo, tied with bark-string to two cross-pieces, and moving in
two grooves round one of the door-posts, so as to open inwards. If
the owner wishes to leave home, he takes two logs as thick as a man’s
upper arm and about a yard long. One of these is placed obliquely
against the middle of the door from the inside, so as to form an angle
of from 60° to 75° with the ground. He then places the second piece
horizontally across the first, pressing it downward with all his might.
It is kept in place by two strong posts planted in the ground a few
inches inside the door. This fastening is absolutely safe, but of course
cannot be applied to both doors at once, otherwise how could the
owner leave or enter his house? I have not yet succeeded in finding
out how the back door is fastened.
MAKONDE LOCK AND KEY AT JUMBE CHAURO

This is the general way of closing a house. The Makonde at Jumbe
Chauro, however, have a much more complicated, solid and original
one. Here, too, the door is as already described, except that there is
only one post on the inside, standing by itself about six inches from
one side of the doorway. Opposite this post is a hole in the wall just
large enough to admit a man’s arm. The door is closed inside by a
large wooden bolt passing through a hole in this post and pressing
with its free end against the door. The other end has three holes into
which fit three pegs running in vertical grooves inside the post. The
door is opened with a wooden key about a foot long, somewhat
curved and sloped off at the butt; the other end has three pegs
corresponding to the holes, in the bolt, so that, when it is thrust
through the hole in the wall and inserted into the rectangular
opening in the post, the pegs can be lifted and the bolt drawn out.[50]
MODE OF INSERTING THE KEY
With no small pride first one householder and then a second

showed me on the spot the action of this greatest invention of the
Makonde Highlands. To both with an admiring exclamation of
“Vizuri sana!” (“Very fine!”). I expressed the wish to take back these
marvels with me to Ulaya, to show the Wazungu what clever fellows
the Makonde are. Scarcely five minutes after my return to camp at
Newala, the two men came up sweating under the weight of two
heavy logs which they laid down at my feet, handing over at the same
time the keys of the fallen fortress. Arguing, logically enough, that if
the key was wanted, the lock would be wanted with it, they had taken
their axes and chopped down the posts—as it never occurred to them
to dig them out of the ground and so bring them intact. Thus I have
two badly damaged specimens, and the owners, instead of praise,
come in for a blowing-up.
The Makua huts in the environs of Newala are especially
miserable; their more than slovenly construction reminds one of the
temporary erections of the Makua at Hatia’s, though the people here
have not been concerned in a war. It must therefore be due to
congenital idleness, or else to the absence of a powerful chief. Even
the baraza at Mlipa’s, a short hour’s walk south-east of Newala,
shares in this general neglect. While public buildings in this country
are usually looked after more or less carefully, this is in evident
danger of being blown over by the first strong easterly gale. The only
attractive object in this whole district is the grave of the late chief
Mlipa. I visited it in the morning, while the sun was still trying with
partial success to break through the rolling mists, and the circular
grove of tall euphorbias, which, with a broken pot, is all that marks
the old king’s resting-place, impressed one with a touch of pathos.
Even my very materially-minded carriers seemed to feel something
of the sort, for instead of their usual ribald songs, they chanted
solemnly, as we marched on through the dense green of the Makonde
bush:—
“We shall arrive with the great master; we stand in a row and have
no fear about getting our food and our money from the Serkali (the
Government). We are not afraid; we are going along with the great
master, the lion; we are going down to the coast and back.”
With regard to the characteristic features of the various tribes here
on the western edge of the plateau, I can arrive at no other
conclusion than the one already come to in the plain, viz., that it is
impossible for anyone but a trained anthropologist to assign any
given individual at once to his proper tribe. In fact, I think that even
an anthropological specialist, after the most careful examination,
might find it a difficult task to decide. The whole congeries of peoples
collected in the region bounded on the west by the great Central
African rift, Tanganyika and Nyasa, and on the east by the Indian
Ocean, are closely related to each other—some of their languages are
only distinguished from one another as dialects of the same speech,
and no doubt all the tribes present the same shape of skull and
structure of skeleton. Thus, surely, there can be no very striking
differences in outward appearance.
Even did such exist, I should have no time
to concern myself with them, for day after day,
I have to see or hear, as the case may be—in
any case to grasp and record—an
extraordinary number of ethnographic
phenomena. I am almost disposed to think it
fortunate that some departments of inquiry, at
least, are barred by external circumstances.
Chief among these is the subject of iron-
working. We are apt to think of Africa as a
country where iron ore is everywhere, so to
speak, to be picked up by the roadside, and
where it would be quite surprising if the
inhabitants had not learnt to smelt the
material ready to their hand. In fact, the
knowledge of this art ranges all over the
continent, from the Kabyles in the north to the
Kafirs in the south. Here between the Rovuma
and the Lukuledi the conditions are not so
favourable. According to the statements of the
Makonde, neither ironstone nor any other
form of iron ore is known to them. They have
not therefore advanced to the art of smelting
the metal, but have hitherto bought all their
THE ANCESTRESS OF
THE MAKONDE
iron implements from neighbouring tribes.
Even in the plain the inhabitants are not much
better off. Only one man now living is said to
understand the art of smelting iron. This old fundi lives close to
Huwe, that isolated, steep-sided block of granite which rises out of
the green solitude between Masasi and Chingulungulu, and whose
jagged and splintered top meets the traveller’s eye everywhere. While
still at Masasi I wished to see this man at work, but was told that,
frightened by the rising, he had retired across the Rovuma, though
he would soon return. All subsequent inquiries as to whether the
fundi had come back met with the genuine African answer, “Bado”
(“Not yet”).
BRAZIER
Some consolation was afforded me by a brassfounder, whom I

came across in the bush near Akundonde’s. This man is the favourite
of women, and therefore no doubt of the gods; he welds the glittering
brass rods purchased at the coast into those massive, heavy rings
which, on the wrists and ankles of the local fair ones, continually give
me fresh food for admiration. Like every decent master-craftsman he
had all his tools with him, consisting of a pair of bellows, three
crucibles and a hammer—nothing more, apparently. He was quite
willing to show his skill, and in a twinkling had fixed his bellows on
the ground. They are simply two goat-skins, taken off whole, the four
legs being closed by knots, while the upper opening, intended to
admit the air, is kept stretched by two pieces of wood. At the lower
end of the skin a smaller opening is left into which a wooden tube is
stuck. The fundi has quickly borrowed a heap of wood-embers from
the nearest hut; he then fixes the free ends of the two tubes into an
earthen pipe, and clamps them to the ground by means of a bent
piece of wood. Now he fills one of his small clay crucibles, the dross
on which shows that they have been long in use, with the yellow
material, places it in the midst of the embers, which, at present are
only faintly glimmering, and begins his work. In quick alternation
the smith’s two hands move up and down with the open ends of the
bellows; as he raises his hand he holds the slit wide open, so as to let
the air enter the skin bag unhindered. In pressing it down he closes
the bag, and the air puffs through the bamboo tube and clay pipe into
the fire, which quickly burns up. The smith, however, does not keep
on with this work, but beckons to another man, who relieves him at
the bellows, while he takes some more tools out of a large skin pouch
carried on his back. I look on in wonder as, with a smooth round
stick about the thickness of a finger, he bores a few vertical holes into
the clean sand of the soil. This should not be difficult, yet the man
seems to be taking great pains over it. Then he fastens down to the
ground, with a couple of wooden clamps, a neat little trough made by
splitting a joint of bamboo in half, so that the ends are closed by the
two knots. At last the yellow metal has attained the right consistency,
and the fundi lifts the crucible from the fire by means of two sticks
split at the end to serve as tongs. A short swift turn to the left—a
tilting of the crucible—and the molten brass, hissing and giving forth
clouds of smoke, flows first into the bamboo mould and then into the
holes in the ground.
The technique of this backwoods craftsman may not be very far
advanced, but it cannot be denied that he knows how to obtain an
adequate result by the simplest means. The ladies of highest rank in
this country—that is to say, those who can afford it, wear two kinds
of these massive brass rings, one cylindrical, the other semicircular
in section. The latter are cast in the most ingenious way in the
bamboo mould, the former in the circular hole in the sand. It is quite
a simple matter for the fundi to fit these bars to the limbs of his fair
customers; with a few light strokes of his hammer he bends the
pliable brass round arm or ankle without further inconvenience to
the wearer.
SHAPING THE POT
SMOOTHING WITH MAIZE-COB
CUTTING THE EDGE

FINISHING THE BOTTOM
LAST SMOOTHING BEFORE

BURNING
FIRING THE BRUSH-PILE

LIGHTING THE FARTHER SIDE OF
THE PILE
TURNING THE RED-HOT VESSEL
NYASA WOMAN MAKING POTS AT MASASI

Pottery is an art which must always and everywhere excite the
interest of the student, just because it is so intimately connected with
the development of human culture, and because its relics are one of
the principal factors in the reconstruction of our own condition in
prehistoric times. I shall always remember with pleasure the two or
three afternoons at Masasi when Salim Matola’s mother, a slightly-
built, graceful, pleasant-looking woman, explained to me with
touching patience, by means of concrete illustrations, the ceramic art
of her people. The only implements for this primitive process were a
lump of clay in her left hand, and in the right a calabash containing
the following valuables: the fragment of a maize-cob stripped of all
its grains, a smooth, oval pebble, about the size of a pigeon’s egg, a
few chips of gourd-shell, a bamboo splinter about the length of one’s
hand, a small shell, and a bunch of some herb resembling spinach.
Nothing more. The woman scraped with the
shell a round, shallow hole in the soft, fine
sand of the soil, and, when an active young
girl had filled the calabash with water for her,
she began to knead the clay. As if by magic it
gradually assumed the shape of a rough but
already well-shaped vessel, which only wanted
a little touching up with the instruments
before mentioned. I looked out with the
MAKUA WOMAN closest attention for any indication of the use
MAKING A POT. of the potter’s wheel, in however rudimentary
SHOWS THE a form, but no—hapana (there is none). The
BEGINNINGS OF THE embryo pot stood firmly in its little
POTTER’S WHEEL
depression, and the woman walked round it in
a stooping posture, whether she was removing
small stones or similar foreign bodies with the maize-cob, smoothing
the inner or outer surface with the splinter of bamboo, or later, after
letting it dry for a day, pricking in the ornamentation with a pointed
bit of gourd-shell, or working out the bottom, or cutting the edge
with a sharp bamboo knife, or giving the last touches to the finished
vessel. This occupation of the women is infinitely toilsome, but it is
without doubt an accurate reproduction of the process in use among
our ancestors of the Neolithic and Bronze ages.
There is no doubt that the invention of pottery, an item in human
progress whose importance cannot be over-estimated, is due to
women. Rough, coarse and unfeeling, the men of the horde range
over the countryside. When the united cunning of the hunters has
succeeded in killing the game; not one of them thinks of carrying
home the spoil. A bright fire, kindled by a vigorous wielding of the
drill, is crackling beside them; the animal has been cleaned and cut
up secundum artem, and, after a slight singeing, will soon disappear
under their sharp teeth; no one all this time giving a single thought
to wife or child.
To what shifts, on the other hand, the primitive wife, and still more
the primitive mother, was put! Not even prehistoric stomachs could
endure an unvarying diet of raw food. Something or other suggested
the beneficial effect of hot water on the majority of approved but
indigestible dishes. Perhaps a neighbour had tried holding the hard
roots or tubers over the fire in a calabash filled with water—or maybe
an ostrich-egg-shell, or a hastily improvised vessel of bark. They
became much softer and more palatable than they had previously
been; but, unfortunately, the vessel could not stand the fire and got
charred on the outside. That can be remedied, thought our
ancestress, and plastered a layer of wet clay round a similar vessel.
This is an improvement; the cooking utensil remains uninjured, but
the heat of the fire has shrunk it, so that it is loose in its shell. The
next step is to detach it, so, with a firm grip and a jerk, shell and
kernel are separated, and pottery is invented. Perhaps, however, the
discovery which led to an intelligent use of the burnt-clay shell, was
made in a slightly different way. Ostrich-eggs and calabashes are not
to be found in every part of the world, but everywhere mankind has
arrived at the art of making baskets out of pliant materials, such as
bark, bast, strips of palm-leaf, supple twigs, etc. Our inventor has no
water-tight vessel provided by nature. “Never mind, let us line the
basket with clay.” This answers the purpose, but alas! the basket gets
burnt over the blazing fire, the woman watches the process of
cooking with increasing uneasiness, fearing a leak, but no leak
appears. The food, done to a turn, is eaten with peculiar relish; and
the cooking-vessel is examined, half in curiosity, half in satisfaction
at the result. The plastic clay is now hard as stone, and at the same
time looks exceedingly well, for the neat plaiting of the burnt basket
is traced all over it in a pretty pattern. Thus, simultaneously with
pottery, its ornamentation was invented.
Primitive woman has another claim to respect. It was the man,
roving abroad, who invented the art of producing fire at will, but the
woman, unable to imitate him in this, has been a Vestal from the
earliest times. Nothing gives so much trouble as the keeping alight of
the smouldering brand, and, above all, when all the men are absent
from the camp. Heavy rain-clouds gather, already the first large
drops are falling, the first gusts of the storm rage over the plain. The
little flame, a greater anxiety to the woman than her own children,
flickers unsteadily in the blast. What is to be done? A sudden thought
occurs to her, and in an instant she has constructed a primitive hut
out of strips of bark, to protect the flame against rain and wind.
This, or something very like it, was the way in which the principle
of the house was discovered; and even the most hardened misogynist
cannot fairly refuse a woman the credit of it. The protection of the
hearth-fire from the weather is the germ from which the human
dwelling was evolved. Men had little, if any share, in this forward
step, and that only at a late stage. Even at the present day, the
plastering of the housewall with clay and the manufacture of pottery
are exclusively the women’s business. These are two very significant
survivals. Our European kitchen-garden, too, is originally a woman’s
invention, and the hoe, the primitive instrument of agriculture, is,
characteristically enough, still used in this department. But the
noblest achievement which we owe to the other sex is unquestionably
the art of cookery. Roasting alone—the oldest process—is one for
which men took the hint (a very obvious one) from nature. It must
have been suggested by the scorched carcase of some animal
overtaken by the destructive forest-fires. But boiling—the process of
improving organic substances by the help of water heated to boiling-
point—is a much later discovery. It is so recent that it has not even
yet penetrated to all parts of the world. The Polynesians understand
how to steam food, that is, to cook it, neatly wrapped in leaves, in a
hole in the earth between hot stones, the air being excluded, and
(sometimes) a few drops of water sprinkled on the stones; but they
do not understand boiling.
To come back from this digression, we find that the slender Nyasa
woman has, after once more carefully examining the finished pot,
put it aside in the shade to dry. On the following day she sends me
word by her son, Salim Matola, who is always on hand, that she is
going to do the burning, and, on coming out of my house, I find her
already hard at work. She has spread on the ground a layer of very
dry sticks, about as thick as one’s thumb, has laid the pot (now of a
yellowish-grey colour) on them, and is piling brushwood round it.
My faithful Pesa mbili, the mnyampara, who has been standing by,
most obligingly, with a lighted stick, now hands it to her. Both of
them, blowing steadily, light the pile on the lee side, and, when the
flame begins to catch, on the weather side also. Soon the whole is in a
blaze, but the dry fuel is quickly consumed and the fire dies down, so
that we see the red-hot vessel rising from the ashes. The woman
turns it continually with a long stick, sometimes one way and
sometimes another, so that it may be evenly heated all over. In
twenty minutes she rolls it out of the ash-heap, takes up the bundle
of spinach, which has been lying for two days in a jar of water, and
sprinkles the red-hot clay with it. The places where the drops fall are
marked by black spots on the uniform reddish-brown surface. With a
sigh of relief, and with visible satisfaction, the woman rises to an
erect position; she is standing just in a line between me and the fire,
from which a cloud of smoke is just rising: I press the ball of my
camera, the shutter clicks—the apotheosis is achieved! Like a
priestess, representative of her inventive sex, the graceful woman
stands: at her feet the hearth-fire she has given us beside her the
invention she has devised for us, in the background the home she has
built for us.
At Newala, also, I have had the manufacture of pottery carried on
in my presence. Technically the process is better than that already
described, for here we find the beginnings of the potter’s wheel,
which does not seem to exist in the plains; at least I have seen
nothing of the sort. The artist, a frightfully stupid Makua woman, did
not make a depression in the ground to receive the pot she was about
to shape, but used instead a large potsherd. Otherwise, she went to
work in much the same way as Salim’s mother, except that she saved
herself the trouble of walking round and round her work by squatting
at her ease and letting the pot and potsherd rotate round her; this is
surely the first step towards a machine. But it does not follow that
the pot was improved by the process. It is true that it was beautifully
rounded and presented a very creditable appearance when finished,
but the numerous large and small vessels which I have seen, and, in
part, collected, in the “less advanced” districts, are no less so. We
moderns imagine that instruments of precision are necessary to
produce excellent results. Go to the prehistoric collections of our
museums and look at the pots, urns and bowls of our ancestors in the
dim ages of the past, and you will at once perceive your error.
MAKING LONGITUDINAL CUT IN
BARK
DRAWING THE BARK OFF THE LOG
REMOVING THE OUTER BARK

BEATING THE BARK
WORKING THE BARK-CLOTH AFTER BEATING, TO MAKE IT

SOFT
MANUFACTURE OF BARK-CLOTH AT NEWALA

To-day, nearly the whole population of German East Africa is
clothed in imported calico. This was not always the case; even now in
some parts of the north dressed skins are still the prevailing wear,
and in the north-western districts—east and north of Lake
Tanganyika—lies a zone where bark-cloth has not yet been
superseded. Probably not many generations have passed since such
bark fabrics and kilts of skins were the only clothing even in the
south. Even to-day, large quantities of this bright-red or drab
material are still to be found; but if we wish to see it, we must look in
the granaries and on the drying stages inside the native huts, where
it serves less ambitious uses as wrappings for those seeds and fruits
which require to be packed with special care. The salt produced at
Masasi, too, is packed for transport to a distance in large sheets of
bark-cloth. Wherever I found it in any degree possible, I studied the
process of making this cloth. The native requisitioned for the
purpose arrived, carrying a log between two and three yards long and
as thick as his thigh, and nothing else except a curiously-shaped
mallet and the usual long, sharp and pointed knife which all men and
boys wear in a belt at their backs without a sheath—horribile dictu!
[51]
Silently he squats down before me, and with two rapid cuts has
drawn a couple of circles round the log some two yards apart, and
slits the bark lengthwise between them with the point of his knife.
With evident care, he then scrapes off the outer rind all round the
log, so that in a quarter of an hour the inner red layer of the bark
shows up brightly-coloured between the two untouched ends. With
some trouble and much caution, he now loosens the bark at one end,
and opens the cylinder. He then stands up, takes hold of the free
edge with both hands, and turning it inside out, slowly but steadily
pulls it off in one piece. Now comes the troublesome work of
scraping all superfluous particles of outer bark from the outside of
the long, narrow piece of material, while the inner side is carefully
scrutinised for defective spots. At last it is ready for beating. Having
signalled to a friend, who immediately places a bowl of water beside
him, the artificer damps his sheet of bark all over, seizes his mallet,
lays one end of the stuff on the smoothest spot of the log, and
hammers away slowly but continuously. “Very simple!” I think to
myself. “Why, I could do that, too!”—but I am forced to change my
opinions a little later on; for the beating is quite an art, if the fabric is
not to be beaten to pieces. To prevent the breaking of the fibres, the
stuff is several times folded across, so as to interpose several
thicknesses between the mallet and the block. At last the required
state is reached, and the fundi seizes the sheet, still folded, by both
ends, and wrings it out, or calls an assistant to take one end while he
holds the other. The cloth produced in this way is not nearly so fine
and uniform in texture as the famous Uganda bark-cloth, but it is
quite soft, and, above all, cheap.
Now, too, I examine the mallet. My craftsman has been using the
simpler but better form of this implement, a conical block of some
hard wood, its base—the striking surface—being scored across and
across with more or less deeply-cut grooves, and the handle stuck
into a hole in the middle. The other and earlier form of mallet is
shaped in the same way, but the head is fastened by an ingenious
network of bark strips into the split bamboo serving as a handle. The
observation so often made, that ancient customs persist longest in
connection with religious ceremonies and in the life of children, here
finds confirmation. As we shall soon see, bark-cloth is still worn
during the unyago,[52] having been prepared with special solemn
ceremonies; and many a mother, if she has no other garment handy,
will still put her little one into a kilt of bark-cloth, which, after all,
looks better, besides being more in keeping with its African
surroundings, than the ridiculous bit of print from Ulaya.
MAKUA WOMEN

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Microbiology: An Introduction 12th

Edition – Ebook PDF Version

is now more mobile-friendly, allowing

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Cover Photo Credit: Staphylococcus, Sebastian Kaulitzki/Alamy

Library of Congress Cataloging-in-Publication Data

ISBN 10: 0-321-92915-2; ISBN 13: 978-0-321-92915-0 (Student edition)

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Gerard J. Tortora Jerry Tortora is a professor of biology and teaches

Christine L. Case Chris Case is a registered microbiologist and a

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Chapter-by-Chapter Revisions Chapter 13

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Chapter 22 ● A discussion of Kawasaki syndrome has been added.

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PART ONE Fundamentals of Microbiology PART FOUR Microorganisms

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PART ONE Fundamentals of Microbiology

1 The Microbial World Through a Microscope 51

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5 Microbial Metabolism 107 7 The Control of Microbial

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12  he Eukaryotes: Fungi, Algae,

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Classifying Infectious Diseases 395 SECOND LINE OF DEFENSE 446

16 I nnate Immunity: Nonspecific

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• Fluorescent-Antibody Techniques • Enzyme-Linked Tests to Guide Chemotherapy 567

20 Antimicrobial Drugs 548

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Bacterial Diseases of the Reproductive Systems 751

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Big Picture Tough Topics Life Cycle Figures

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Note: Answers to In the Clinic questions are found online at MasteringMicrobiology.

he overall theme of this textbook is the relationship between microbes—

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Designer Jeans: Made by Microbes?

the color of many colonies of this bacterium. The genus of the

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Newala, too, suffers from the distance of its water-supply—at least

The water-supply of New Newala is in the bottom of the valley,

NATIVE PATH THROUGH THE MAKONDE BUSH, NEAR

MAKONDE LOCK AND KEY AT JUMBE CHAURO

1 The Microbial World Through a Microscope 51

5 Microbial Metabolism 107 7 The Control of Microbial

12 he Eukaryotes: Fungi, Algae,

16 I nnate Immunity: Nonspecific

20 Antimicrobial Drugs 548