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Fifth Edition

Microbiology
With Diseases by Taxonomy

Robert W. Bauman, Ph.D.

Amarillo College

Co n t r i b u t i o n s By:

Todd P. Primm, Ph.D.

Sam Houston State University

Elizabeth Machunis-Masuoka, Ph.D.

University of Virginia

C l i n i c a l C o n s u lta n t s :

Cecily D. Cosby, Ph.D., FNP-C, PA-C

Jean E. Montgomery, MSN, RN
Senior Acquisitions Editor: Kelsey Churchman
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Library of Congress Cataloging-in-Publication Data

Bauman, Robert W., author.
   Microbiology. With diseases by taxonomy/Robert W. Bauman; contributions by Todd
P. Primm, Elizabeth Machunis-Masuoka; clinical consultants, Cecily D. Cosby,
Jean E. Montgomery. — Fifth edition.
   p.; cm.
   With diseases by taxonomy
   ISBN 978-0-13-401919-2 (Student edition) — ISBN 0-13-401919-9 (Student edition) —
ISBN 0-13-429875-6 (Instructor’s review copy) — ISBN 978-0-13-429875-7
(Instructor’s review copy)
   I. Primm, Todd P., contributor. II. Machunis-Masuoka, Elizabeth, contributor. III. Title.
IV. Title: With diseases by taxonomy.
   [DNLM: 1. Microbiological Phenomena. 2. Microbiological Processes. QW 4]
  QR41.2
  616.9’041—dc23
2015029578

ISBN 10: 0-13-401919-9 (Student edition)

ISBN 13: 978-0-13-401919-2 (Student edition)
ISBN 10: 0-13-429875-6 (Instructor’s Review Copy)
ISBN 13: 978-0-13-429875-7 (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 Author
ROBERT W. BAUMAN is a professor of biology and past chairman of the Department of
Biological Sciences at Amarillo College in Amarillo, Texas. He has taught microbiology, human anatomy and
physiology, and botany. In 2004, the students of Amarillo College selected Dr. Bauman as the recipient of the
John F. Mead Faculty Excellence Award and he has been nominated for the one-time award every year since.
He received an M.A. degree in botany from the University of Texas at Austin and a Ph.D. in biology from
Stanford University. His research interests have included the morphology and ecology of freshwater algae,
the cell biology of marine algae (particularly the deposition of cell walls and intercellular communication),
environmentally triggered chromogenesis in butterflies, and terrestrial oil pollution remediation by naturally
occurring bacteria. He is a member of the American Society of Microbiology (ASM) where he has held
national offices, Texas Community College Teachers Association (TCCTA) where he serves in a statewide
position of leadership, American Association for the Advancement of Science (AAAS), Human Anatomy
and Physiology Society (HAPS), and The Lepidopterists’ Society. When he is not writing books, he enjoys
spending time with his family: gardening, hiking, camping, rock climbing, backpacking, cycling, skiing, and
reading by a crackling fire in the winter and in a gently swaying hammock in the summer.

TODD P. PRIMM (contributor) is an associate professor at Sam Houston State University,

where he teaches pre-nursing microbiology. He also serves as Director of the Professional and Academic
Center for Excellence, which focuses on improving teaching and learning on campus. In 2010, he was
Distinguished Alumnus of the Graduate School of Biomedical Sciences of Baylor College of Medicine,
where he earned a Ph.D. in Biochemistry in 1997. He received a B.S. from Texas A&M University in
1992. He is very active in the American Society for Microbiology and received the Texas Branch 2015
Faculty Teaching Award. He was chair of the organizing committee for the 2013 ASM Conference
for Undergraduate Educators, participated in the 2012 Research Residency of the ASM/NSF Biology
Scholars Program, and currently serves on the editorial board for the Journal of Microbiology and Biology
Education. He is also an affiliate staff member with the international organization Cru. He loves teaching
and mentoring students and spending time with his wonderful wife of 23 years and four children.

About the Clinical Consultants

CECILY D. COSBY is nationally certified as both
a family nurse practitioner and physician assistant. She is a
professor of nursing, currently teaching at Samuel Merritt
University in Oakland, California, and has been in clinical
practice since 1980. She received her Ph.D. and M.S. from the
University of California, San Francisco; her BSN from California
State University, Long Beach; and her P.A. certificate from the
Stanford Primary Care program. She is the Director of Samuel
Merritt University’s Doctor of Nursing Practice Program.
JEAN E. MONTGOMERY is a registered nurse
formerly teaching in the associate degree nursing program at
Austin Community College in Texas. She received her MSN
from the University of Texas Health Science Center at San
Antonio, Texas.
ix
x CHAPTER 1 A Brief History of Microbiology

Preface
The reemergence of whooping cough, mumps, and measles and the emergence of snail fever, spot-
ted fever rickettsiosis, Middle East respiratory syndrome, and other diseases; the cases of strep
throat, MRSA, and tuberculosis; the progress of cutting-edge research into microbial genetics; the
challenge of increasingly drug-resistant pathogens; the continual discovery of microorganisms
previously unknown—these are just a few examples of why exploring microbiology has never
been more exciting, or more important. Welcome!
I have taught microbiology to undergraduates for over 27 years and witnessed firsthand how
students struggle with the same topics and concepts year after year. To address these challenging
topics, I have created 14 new Video Tutors: three in addition to those already incorporated into the
first 18 chapters of the text and 11 that cover the Disease in Depth features. The Video Tutors and
Disease in Depth features walk students through key concepts in microbiology, bringing the art of
the textbook to life and important concepts into view. In creating this textbook, my aim was to help
students see complex topics of microbiology—especially metabolism, genetics, and immunology—
in a way that they can understand, while at the same time presenting a thorough and accurate
overview of microbiology. I also wished to highlight the many positive effects of microorganisms
on our lives, along with the medically important microorganisms that cause disease.

New to This Edition

In approaching the fifth edition, my goal was to build upon the strengths and success of the previ-
ous editions by updating it with the latest scientific and educational research and data available
and by incorporating the many terrific suggestions I have received from colleagues and students
alike. The feedback from instructors who adopted previous editions has been immensely gratifying
and is much appreciated. The Microbe at a Glance features have been widely praised by instructors
and students, so I, along with art editor Kelly Murphy, developed 11 new Disease in Depth features,
most as two-page spreads, that use compelling art and photos to provide a detailed, visually un-
surpassed overview of a specific disease. Each Disease in Depth feature includes an Investigate It!
question with a QR code directing students to a Video Tutor that explores the topic and encourages
further, independent research. These activities are assignable in MasteringMicrobiology®. Another
goal for this edition was to provide additional instruction on important foundational concepts and
processes. To that end, I developed and narrated three new core concept Video Tutors, accessible
via QR codes in the textbook and assignable in MasteringMicrobiology.
The result is, once again, a collaborative effort of educators, students, editors, and top scientific
illustrators: a textbook that, I hope, continues to improve upon conventional explanations and
illustrations in substantive and effective ways.
In this new edition:

• NEW Disease in Depth features highlight important and representative diseases for each body
system, extending the visual impact of the art program as well as the highly praised M ­ icrobe at a
Glance features. Each of these 11 visual features contains infographics, provides i­ n-depth cover-
age of the selected disease, and includes a QR code and Investigate It! question that d ­ irects stu-
dents to a Video Tutor exploring the topic and prompting further inquiry and critical thinking.

x
 Preface xi

New assignable Disease in Depth coaching activities in Mastering­Microbiology® ­encourage

­students to apply and test their understanding of key concepts.

• NEW Video Tutors developed and narrated by the author walk students through key con-
cepts. New to this edition are Video Tutors on glycolysis, protein translation, and antigen
processing. These Video Tutors bring the textbook art to life and help students visualize and
understand tough topics and important processes. Thirty-two video tutorials are accessible
via QR codes in the textbook and are accompanied by multiple-choice questions, assignable
in MasteringMicrobiology®.

• NEW Tell Me Why critical thinking questions end every main section within each chapter.
These questions strengthen the pedagogy and organization of each chapter and consistently
provide stop-and-think opportunities for students as they read.

• NEW Expanded coverage of helminths is provided in new Highlight features, and an

­emphasis on virulence factors is included the Disease in Depth features.

• The genetics chapters (Chapters 7–8) have been reviewed and revised by genetics special-
ists. These now reflect the most current understanding of this rapidly evolving field, includ-
ing new discussion of next-generation DNA sequencing.

• Over 330 NEW and revised micrographs, photos, and figures enhance student under-
standing of the text and boxed features.

• NEW and EXPANDED MasteringMicrobiology includes new Interactive Microbiology ­

animations and tutorials; new MicroBooster remedial video tutorials; new Disease in
Depth ­coaching activities; new Video Tutors with assessments; new MicroCareers and
Clinical Case Study coaching activities; and a plethora of microbiology lab resources. NEW
­Interactive ­Microbiology is a dynamic suite of interactive tutorials and animations that
teach key ­concepts in the context of a clinical setting. Students actively engage with each
topic and learn from manipulating variables, predicting outcomes, and answering forma-
tive and ­summative assessments. Topics include Operons; Complement; Biofilms and Quo-
rum ­Sensing; Antibiotic Resistance, Mechanisms; Antibiotic Resistance, Selection; Aerobic
­Respiration in Prokaryotes; and Human Microbiota. NEW MicroBoosters are a suite of brief
video tutorials that cover key concepts that students often need to review, including Study
Skills, Math, Basic Chemistry, Cell Biology, Basic Biology and more! The Micro Lab resources
include MicroLab Tutors, which use lab technique videos, 3-D molecular animations, and
step-by-step tutorials to help students make connections between lecture and lab; Lab
­Technique Videos and pre-lab quizzes to ensure that students come prepared for lab time;
and Lab Practical and post-lab quizzes to reinforce what students have learned.

MasteringMicrobiology offers students access to Dynamic Study Modules to help them acquire,
retain, and recall information faster and more efficiently than ever before with textbook-specific
explanations and art. Dynamic Study Modules are available for use as a self-study tool or as
assignments. Instructors also now have the option to give Adaptive Follow-Up assignments that
provide student-specific additional coaching and practice. These question sets continuously adapt
to each student’s needs, making efficient use of homework time.
MasteringMicrobiology also includes Learning Catalytics—a “bring your own device” student
engagement, assessment, and classroom intelligence system. With Learning Catalytics, instructors
can assess students in real time using open-ended tasks to probe student understanding using
Pearson’s library of questions or designing their own.
The following section provides a detailed outline of this edition’s chapter-by-chapter revisions.
xii CHAPTER 1 A Brief History of Microbiology
Chapter-by-Chapter Revisions
CHAPTER 1 A Brief History of Microbiology
• Added three Tell Me Why critical thinking questions to text
• Added three new photos (chapter opener, Fig. 1.6b, Highlight box
on MERS)
• Updated map showing countries having transmission of variant
Creutzfeldt-Jakob disease (vJCD)
• Added CDC-preferred term “healthcare-associated infection (HAI)”
(formerly nosocomial infection)
• Added introductory coverage of normal microbiota and of agar in
micro labs
• Clarified the use of controls in Pasteur’s experiment to ­disprove
spontaneous generation
• Clarified industrial use of microbes in making yogurt and pest
control
• Introduced the success of gene therapy to treat several ­inherited
immune deficiencies
• Updated box: “The New Normal”: The Challenge of Emerging and
Reemerging Diseases to include Middle East respiratory syndrome
(MERS), Ebola, chikungunya, and measles
• Added to list of current problems in microbiology: biofilms, tests for
infections, and persistent antimicrobial-drug resistance
• Added three critical thinking questions to Emerging ­Disease Case
Study: Variant Creutzfeldt-Jacob Disease
• New end-of-chapter, short-answer question on healthcare-
associated (nosocomial) infections
• Added fill-in Concept Map over types of microbes and some of
their major characteristics
Chapter 2 The Chemistry of Microbiology
• Added five Tell Me Why critical thinking questions to text
• Eleven figures revised for better pedagogy (Figs. 2.2, 2.3, 2.6,
2.11, 2.15, 2.17, 2.19, 2.21, 2.22, 2.23; amino group in Table 2.3)
• New Learning Outcomes concerning terms regarding ­elements,
valence electrons and chemical bonding, organic compounds,
contrasting ionic and covalent bonds, and lipids
• New figure legend question for enhanced pedagogy (Fig. 2.3)
• Expanded coverage of term “nucleoside” because nucleoside
analogs treat many diseases
• Added fill-in Concept Map over nucleotide structure and function
Chapter 3 Cell Structure and Function
• Added 12 Tell Me Why critical thinking questions to text
• Two new photos (Figs. 3.5b, 3.8a)
• Revised and enhanced artwork in 14 figures for enhanced
pedagogy (Figs. 3.4, 3.8b, 3.9, 3.12, 3.14, 3.15, 3.17, 3.18, 3.19, 3.20,
3.21, 3.22, 3.24, 3.35)
• Added one new figure (structure of glucose versus NAG and NAM)
(Fig. 3.13)
• Enhanced discussion of flagella and cilia structure and ­function,
comparison and contrast between the outer and cytoplasmic
membranes of Gram-negative cells, and m ­ ovement across cell
membranes
xii
Chapter 4 Microscopy, Staining, and Classification
• Added four Tell Me Why critical thinking questions to text
• Revised two figures for enhanced pedagogy (Figs. 4.4, 4.6)
• Revised Learning Outcome regarding simple stains, which now
include Gomori methenamine silver stain and h ­ ematoxylin and
eosin stains
• Added fill-in-the-blank Concept Map about Gram stain and cell
wall structure to end-of-chapter review
• Revised coverage of history of taxonomy
• Expanded discussion of resolution, immersion oil, mordants,
definition of microbial species, and role of George Fox in the
discovery of the archaea and three domains of life
• Revised section on microbial taxonomy to more fully ­address
genomic techniques in taxonomy
• At request of reviewers and instructors, removed ­detailed figures
for dark field, phase, and scanning electron ­microscopy so as to
reduce complexity and chapter length
• Added three critical thinking questions and a new photo to
Emerging Disease Case Study: Necrotizing Fasciitis
Chapter 5 Microbial Metabolism
• Added six Tell Me Why critical thinking questions to text
• Added two new figure questions (Figs. 5.4, 5.13)
• Added one new end-of-chapter fill-in-the-blank question
• Revised 14 figures for greater clarity and better pedagogy (Figs. 5.5,
5.6, 5.10, 5.11, 5.12, 5.13, 5.14, 5.16, 5.17, 5.18, 5.19, 5.26, 5.30; end-of-
chapter critical thinking question 1)
• Clarified and expanded discussion of enzymatic activation through
allosteric sites and competitive and noncompetitive inhibition of
enzyme activity
• Added fill-in Concept Map over aerobic respiration
Chapter 6 Microbial Nutrition and Growth
• Added three Tell Me Why critical thinking questions to text
• Revised five figures for greater clarity and better pedagogy
(Figs. 6.7, 6.8, 6.9, 6.17, 6.20)
• Added two new photos (Figs. 6.13, 6.24b)
• Expanded discussion of singlet oxygen and superoxide ­radicals as
oxidizing agents
• Clarified the method of counting microbes using a cell counter
• Added fill-in Concept Map over culture media
Chapter 7 Microbial Genetics
• Added four Tell Me Why critical thinking questions to text
• Upgraded 20 figures for greater clarity, accuracy, ease of reading,
and better pedagogy (Figs. 7.1, 7.5, 7.6, 7.7, 7.9, 7.10, 7.11, 7.13, 7.20,
7.21, 7.22, 7.23, 7.26, 7.27, 7.28, 7.30, 7.34, 7.35, 7.36, 7.37)
• Updated text to discuss the smallest cellular genome at 112,091 bp
(candidatus Nasuia deltocephalinicola)
• Included recent discovery that chloroplast chromosomes are linear
rather than circular
• Increased discussion of use of RNA as enzymes (ribozymes)

• Expanded table comparing and contrasting DNA replication,
transcription, and translation
• Discussed codon and tRNA for 21st amino acid, selenocysteine
• Enhanced and clarified discussion of lac and trp operons and of the
action of cAMP and CAP as activators
• Expanded and reorganized discussion of DNA repair systems
• Clarified and updated information on the events in ­conjugation,
particularly with Hfr cells
• Expanded coverage of nucleotides and pyrophosphate (diphosphate)
• Added critical thinking questions to Emerging Disease Case Study:
Vibrio vulnificus Infection
• Revised the chapter to better explain differences between ­archaeal,
bacterial, and eukaryotic genetics
• Added fill-in Concept Map over point mutations
Chapter 8 Recombinant DNA Technology
• Added five Tell Me Why critical thinking questions to text
• Added six Learning Outcomes concerning uses of ­synthetic nucleic
acids, PCR, fluorescent in situ hybridization (FISH), functional
genomics, Sanger sequencing, and next-­generation sequencing
• Added one new figure (Fig. 8.10)
• Modified Fig. 8.7 for better pedagogy
• Deleted figures for Southern blots and Sanger automated DNA
sequencing as these techniques are historical and ­less-commonly
used today
• Added discussion of real-time PCR (RT-PCR), Sanger sequencing
methods, next-generation DNA sequencing (NGS), including
pyrosequencing and fluorescent methods, functional genomics,
microbiomes, and biomedical animal models
• New Highlight boxes: How Do You Fix a Mosquito? on controlling
dengue and The Human Microbiome Project
Chapter 9 Controlling Microbial Growth in the
Environment
• Added four Tell Me Why critical thinking questions to text
• Revised five figures for better accuracy, currency, and ­pedagogy
(Figs. 9.2, 9.7, 9.13, 9.15, 9.16)
• Two new photos (Fig. 9.9, Beneficial Microbes)
• Updated techniques for deactivation of prions, coverage of
thimerosal in vaccines, and activity of AOAC International in
developing disinfection standards
• Added three critical thinking questions to Emerging Disease Case
Study: Acanthamoeba Keratitis
• Added critical thinking question concerning salmonellosis
pandemic from smoked salmon
• Added fill-in Concept Map over moist heat applications to control
microbes
Chapter 10 Controlling Microbial Growth in the
Body: Antimicrobial Drugs
• Added four Tell Me Why critical thinking questions to text
• Updated and revised tables of antimicrobials to include all
new antimicrobials mentioned in disease chapters, including
carbapenems and capreomycin (antibacterials); enfuvirtide (newly
approved anti-HIV-1); ciclopirox (antifungal); and bithionol
(anthelmintic); updated sources of drugs, modes of action, clinical
considerations, and methods of resistance
• Updated adverse effects of aminoglycosides
• Updated the mechanism of resistance against quinolone
­antibacterial drugs
• Removed amantadine as a treatment for influenza A
Chapter-by-Chapter Revisions xiii
• Revised seven figures for greater clarity, accuracy, ease of reading,
and better pedagogy (Figs. 10.2, 10.3, 10.6, 10.8, 10.13, 10.15; map of
worldwide, community-associated MRSA)
• Three new photos (Highlight, Fig. 10.10, Clinical Case Study)
• Added three critical thinking questions to Emerging Disease Case
Study: Community-Associated MRSA and updated map with
newly published data
Chapter 11 Characterizing and Classifying
Prokaryotes
• Added four Tell Me Why critical thinking questions to text
• Six new Learning Outcomes (for proteobacteria, including newly
discovered zetaproteobacteria)
• Thirteen new photos (Figs. 11.1, 11.2a, 11.5, 11.7, 11.11a, 11.16, 11.17,
11.19, 11.21, 11.22, 11.23, 11.24b, 11.27b)
• Ten revised figures for better pedagogy (Figs. 11.1, 11.3, 11.4, 11.6,
11.10, 11.14, 11.17, 11.21, 11.26, 11.27)
• Clarified and expanded coverage of (1) “snapping ­division,”
which is a distinctive characteristic of corynebacteria, i­ ncluding C.
diphtheriae, (2) floc formation and its use in s­ ewage treatment, and
(3) methicillin-resistant strains of Staphylococcus aureus
• Updated with new discoveries in bacterial and archaeal
­­systematics: six classes of proteobacteria rather than four and five
phyla of archaea (rather than two)
• Removed box on Botox and box on the possible link between
­cyanobacteria and brain disease to make room for new material
• Three new critical thinking questions over pertussis as a
­reemerging disease
• Added fill-in Concept Map over domain Archaea
Chapter 12 Characterizing and Classifying
Eukaryotes
• Added six Tell Me Why critical thinking questions to text
• Eight new photos (Figs. 12.11, 12.12a and b, 12.13c, 12.14, 12.20,
12.25, 12.27)
• Seven revised figures for more accurate and lucid pedagogy
(Figs. 12.1, 12.3, 12.7, 12.8, 12.17, 12.23; map for aspergillosis)
• As reviewers requested, shortened chapter by ­eliminating detailed
discussion and artwork of ciliate (Paramecium) ­conjugation
and of sexual reproduction by zygomycetes, ­ascomycetes, and
basidiomycetes
• Updated algal, fungal, protozoan, water mold, and slime mold
taxonomy
• Clarified and expanded coverage of (1) meiosis, (2) alveoli in
protists, and (3) use of radiation as an energy source for some fungi
• Added new critical thinking questions: three about the emerging
disease aspergillosis and two at end of chapter about genomics in
relationship to metabolism in various environments
• Added fill-in Concept Map over eukaryotic microorganisms
Chapter 13 Characterizing and Classifying Viruses,
Viroids, and Prions
• Added four Tell Me Why critical thinking questions to text
• Four new photos (Figs. 13.1b, 13.21, 13.24; bacteriophage box)
• Upgraded eight figures for better pedagogy and currency (Figs.
13.5, 13.8, 13.12, 13.13, 13.14, 13.16, 13.18, 13.22)
• One new figure showing prion templating (Fig. 13.23)
• Two new Learning Outcomes concerning (1) structures of viruses
and (2) control of prions
• Updated viral nomenclature to correspond to changes ­approved by
the International Committee on Taxonomy of V ­ iruses (ICTV) in 2014
xiv Chapter-by-Chapter Revisions
• Added discussion on the benefits and costs to a virus of ­having an
envelope versus being naked
• Clarified and expanded text concerning lytic cycle of phage
replication; use of phage typing; replication of animal ­viruses,
particularly ssDNA viruses; link between viruses and human
cancers; viroids; and prions
• Updated techniques for deactivation of prions and treatment of
prion disease
• Updated Emerging Disease Case Study: Chikungunya; added three
critical thinking questions to the discussion
Chapter 14 Infection, Infectious Diseases, and
Epidemiology
• Added eight Tell Me Why critical thinking questions to text
• Changed eight figures for better pedagogy, timeliness, or clarity
(Figs. 14.3, 14.4, 14.5, 14.9, 14.10, 14.14, 14.16, 14.20)
• Revised and updated coverage of (1) number of human cells in a
body and the number of cellular microbiota, (2) microbiome, and
(3) symbioses (added terms symbiont and amensalism)
• Updated to replace term nosocomial with healthcare-associated (in all
chapters)
• Updated epidemiology charts, tables, and graphs
• Updated list of nationally notifiable infectious diseases
• Three new critical thinking questions added to the d­ iscussion of
Hantavirus as an emerging disease
• Added fill-in Concept Map over transmission of diseases
Chapter 15 Innate Immunity
• Added two Tell Me Why critical thinking questions to text
• Modified nine figures for enhanced clarity and better ­pedagogy
(Figs. 15.4, 15.6, 15.7, 15.8, 15.9, 15.11, 15.12, 15.13, 15.14)
• Three new photos (Figs. 15.1, 15.5b)
• Updated and expanded coverage of the action of ­antimicrobial
peptides (defensins), Toll-like receptor 10 (TLR10), complement
activation, complement cascade, and membrane attack complexes
• Expanded and clarified discussion of inflammatory mediators
Chapter 16 Specific Defense: Adaptive Immunity
• Added three Tell Me Why critical thinking questions to text
• Revised and clarified (1) function and structure of tonsils, (2) flow
of lymph, and (3) mucosa-associated lymphoid tissue
• Reordered the discussion of topics in adaptive immunity to better
align with the way events occur; for example, MHC and antigen
processing are discussed before T cells and ­cell-mediated immunity,
which are discussed before B cells and antibody-mediated immunity
• Removed discussion of T-independent antibody immunity as it was
too advanced for beginning students
• Revised three pieces of art for enhanced pedagogy (Figs. 16.2, 16.3,
16.10)
• Added three critical thinking questions and updated ­incidence map
for the discussion of microsporidiosis
• Added fill-in Concept Map over antibodies
Chapter 17 Immunization and Immune Testing
• Added a Tell Me Why critical thinking question to text
• Updated to newly revised CDC 2015 vaccination schedule for
children, adolescents, and adults
• Updated table of vaccine-preventable diseases in the United States
• Enhanced discussion of development of attenuated viral vaccines
• Added two points to chapter summary about recombinant gene
technology and vaccine production and about vaccine safety
• Revised five figures for better pedagogy (Figs. 17.2, 17.3, 17.6,
17.11, 17.14)
Chapter 18 Hypersensitivities, Autoimmune Diseases,
and Immune Deficiencies
• Added three Tell Me Why critical thinking questions to text
• Revised one figure for greater clarity and accuracy (Fig. 18.7)
• Expanded coverage of type III hypersensitivity, the relationship
between hypersensitivities and autoimmune disorders
• Removed figure and text for a very rare disease, immune throm­
bocytopenic purpura, to make room for new material in
Chapter 19
Chapter 19 Pathogenic Gram-Positive Bacteria
• Added nine Tell Me Why critical thinking questions to text
• Added three Disease in Depth visual presentations of disease:
necrotizing fasciitis, listeriosis, and tuberculosis
• Twenty-five new photos (Figs. 19.1, 19.12, 19.17, 19.19, 19.20, 19.21)
• Seven revisions to figures for consistency, currency, ­accuracy, and
better pedagogy (Figs. 19.5, 19.23; Disease in Depth: Necrottizing
Fasciitis, Listeriosis, and Tuberculosis; Microbe at a Glance:
Streptococcus and Clostridium)
• Updated all diagnoses and incidence data
• Revised two Learning Outcomes for better pedagogy (19.10, 19.13)
• Revised Chapter Summary for better pedagogy (for ­Staphylococcus;
Streptococcus; Enterococcus, Bacillus; Clostridium; Listeria; Mycoplasma;
Corynebacterium; Mycobacterium)
• Updated definitions for multi-drug-resistant (MDR) and
­extensively drug-resistant (XDR) tuberculosis
• Updated treatment regimen for inhalation anthrax, ­bioterrorist
anthrax, botulism, tetanus, listeriosis, m
­ ycoplasmal p ­ neumonia,
nongonococcal urethritis, and tuberculosis
• Updated and enhanced discussion of mycolic acids, role of
Streptococcus mutans in tooth decay, and anthrax vaccine
• Added a figure question regarding snapping division in
corynebacteria
• Added three critical thinking questions and updated ­incidence
maps for the discussion of Buruli ulcer
• Added Clinical Case Study regarding tuberculosis
Chapter 20 Pathogenic Gram-Negative Cocci
and Bacilli
• Added three Tell Me Why critical thinking questions to text
• Added one Disease in Depth visual presentation of disease on
urinary tract infections
• Updated all diagnoses and incidence data, including maps
• Updated to replace term nosocomial with healthcare-associated
• Revised Chapter Summary for better pedagogy (Pathogenic, Gram-
Negative, Facultatively Anaerobic Bacilli; ­Pathogenic, Gram-Negative,
Aerobic Bacilli; Pathogenic, Gram-­Negative, Anaerobic Bacilli)
• Updated treatment regimen for gonorrhea, meningococcus
meningitis, bubonic plague, bartonellosis, brucellosis, and
Legionnaires’ disease
• Added one new figure (Fig. 20.1) and figure question on the
potential effects of lipid A
• Revised nine figures for better pedagogy (Microbe at a Glance:
Neisseria gonorrhoeae; Figs. 20.2, 20.3, 20.14, 20.18, 20.19, 20.22,
20.23, 20.28)
• Added three critical thinking questions and updated ­incidence
maps for the discussion of melioidosis

Chapter 21 Rickettsias, Chlamydias, Spirochetes,
and Vibrios
• Added three Tell Me Why critical thinking questions to text
• New Disease in Depth: Spotted Fever Rickettsiosis
• Updated all diagnoses and incidence data
• Modified/updated nine figures (Figs. 21.1, 21.2, 21.3, 21.5, 21.8,
21.12, 21.13, 21.17, 21.20)
• Two new photos (Figs. 21.11, 21.19)
• Updated treatment regimen for rickettsial spotted fever (Rocky
Mountain spotted fever, RMSF), murine typhus, scrub typhus,
human monocytic ehrlichiosis, ­anaplasmosis (formerly called
human granulocytic ehrlichiosis), ­lymphogranuloma venereum,
trachoma, cholera, and gastric ulcers
• Updated and expanded coverage of epidemic typhus, murine
typhus, scrub typhus, spotted fever rickettsioses (RMSF),
ehrlichiosis, anaplasmosis, lymphogranuloma ­venereum, urethritis,
yaws, Borrelia, and cholera
Chapter 22 Pathogenic Fungi
• Added five Tell Me Why critical thinking questions to text
• Added new Disease in Depth: Candidiasis
• Updated all diagnoses and incidence data
• New Learning Outcomes: antifungal vaccines, mycetomas
• Added two new photos for enhanced pedagogy (Figs. 22.12, 22.20)
• Updated treatment regimen for ­paracoccidioidomycosis,
Pneumocystis pneumonia, candidiasis, aspergillosis, ­Malassezia
infections, mycetoma, and sporotrichosis
• Enhanced discussion of dearth of antifungal vaccines
• Added three critical thinking questions and updated ­incidence
maps for the discussion of blastomycosis
• Added fill-in Concept Map over systemic mycoses
Chapter 23 Parasitic Protozoa, Helminths,
and Arthropod Vectors
• Added four Tell Me Why critical thinking questions to text
• Added two new Disease in Depth spreads: Giardiasis and Malaria
• Rearranged the chapter to cover vectors first; expanded ­coverage of
vectors
• New Learning Outcomes: parasitology, definitive versus
intermediate hosts, biological versus mechanical vectors, ­ascariasis,
hookworm infestations, pinworms, anisakiasis
• Updated all diagnoses and incidence data
• Updated treatment regimen for Acanthamoeba keratitis,
­leishmaniasis, trichomoniasis, malaria, Cryptosporidium ­enteritis,
and infestation with Fasciola
• Added mention of emerging human pathogen of malaria:
Plasmodium knowlesi
• Updated stages in life cycle of Toxoplasma
• Simplified discussion of life cycles of Trypanosoma cruzi and of T.
brucei
• Added roundworm Anisakis and its disease anisakiasis at teachers’
requests
• Twenty-four new, more engaging photos (Figs. 23.2, 23.10, 23.12,
23.13, 23.18; Disease in Depth: Giardiasis; Disease in Depth:
Malaria; Emerging Disease Case Study: Babesiosis)
Chapter-by-Chapter Revisions xv
• Eight revised, updated, enhanced, and pedagogically more effective
figures (Figs. 23.1, 23.3, 23.5, 23.6, 23.9, 23.14, 23.17, 23.24)
• Added three critical thinking questions and updated ­incidence
maps for the discussions of babesiosis and of schistosomiasis
• Added fill-in Concept Map over intestinal protozoan parasites
Chapter 24 Pathogenic DNA Viruses
• Added five Tell Me Why critical thinking questions to text
• Updated all diagnoses and incidence data
• Updated treatment regimen for shingles, history of smallpox
vaccination, and the effect of adenovirus 36 on obesity
• Four new photos (Figs. 24.3, 24.15, 24.16c, 24.22)
• Reformatted one figure for better pedagogy (Fig. 24.21)
• Added three critical thinking questions and updated ­incidence
maps for the discussion of monkeypox
• New Disease in Depth: Papillomas with three new photos and three
new figures
Chapter 25 Pathogenic RNA Viruses
• Added six Tell Me Why critical thinking questions to text
• Updated all diagnoses and incidence data
• Updated treatment regimen for colds, hepatitis E, hepatitis C, AIDS,
measles, respiratory syncytial virus infection, and Lassa hemorrhagic
fever
• Updated, revised, and expanded discussion of coronavirus
respiratory syndromes, Nipah virus encephalitis, hepatitis E virus,
and respiratory syncytial viral disease
• Clarified definition of zoonosis
• Added Learning Outcome about mumps
• Sixteen figures revised, updated, or enhanced for better ­pedagogy
(Figs. 25.2, 25.9, 25.10, 25.11, 25.12, 25.14, 25.17, 25.18, 25.19, 25.21,
25.23, 25.24, 25.26, 25.28, 25.29, 25.36)
• Thirteen new photos (chapter opener; Figs. 25.1, 25.7, 25.16b,
25.22b, 25.27, 25.30, 25.32; Highlight box on bats and Nipah virus)
• New Microbe at a Glance box on measles virus
• Two new Emerging Disease Case Study boxes on norovirus
gastroenteritis and tick-borne encephalitis
• Two new Disease in Depth features on Ebola hemorrhagic fever
and influenza
• Added three critical thinking questions to the box on ­influenza
H1N1
Chapter 26 Industrial and Environmental
Microbiology
• Added four Tell Me Why critical thinking questions to text
• Added Learning Outcome on eutrophication
• Three figures revised, updated, or enhanced for better ­pedagogy
(Figs. 26.6, 26.8, 26.15)
• Revised and clarified water contamination and water pollution
• Updated list of bioterrorist threats to include the additions
to category C
• New Emerging Disease Case Study regarding primary ­amebic
meningoencephalitis (Naegleria fowleri infection)
Reviewers for the Fifth Edition
I wish to thank the hundreds of instructors and students who participated in reviews, class
tests, and focus groups for earlier editions of the textbook. Your comments have informed this
book from beginning to end, and I am deeply grateful. For the fifth edition, I extend my deepest
appreciation to the following reviewers.

Denice D. King
Book Reviewers Cleveland State Community College
Christopher Thompson
Loyola University, Maryland
Dena Berg Todd Martin Marie N. Yearling
Tarrant County College Metropolitan Community College, Laramie County Community College
Carroll Bottoms Blue River
Collin College Jennifer Metzler
Nick Butkevitch Ball State University Video Tutor Reviewers
Schoolcraft College Mary Miller Jason Adams
Kari Cargill Baton Rouge Community College College of Dupage
Montana State University Alicia Musser
Abiodun Adibi
Richard J. Cristiano Lansing Community College
Houston Community College Hampton University
Gregory Nasello
Northwest—Spring Branch campus Lewis and Clark Community College Melody J. Bernot
Ann Evancoe Dana Newton Ball State University
Hudson Valley Community College College of the Albemarle Denise Foley
Tod Fairbanks Johanna Porter-Kelley Santiago Canyon College
Palm Beach State College Winston-Salem State University Emily Getty
Teresa G. Fischer Jennifer Reaves Ivy Tech Community College
Indian River State College Jackson State Community College Mary Ann Arnold Hedrick
Sandra M. Fox-Moon Jackie Reynolds Wytheville Community College
Anne Arundel Community College and Richland College
University of Maryland Cristi Hunnes
Steven Scott Rocky Mountain College
Eric Gillock Merritt College
Fort Hays State University Sudeep Majumdar
Amy Siegesmund
Raymond Harris Temple College
Pacific Lutheran University
Prince George’s Community College Tony A. Slieman Bhavya Mathur
Jennifer Hatchel University of South Dakota Chattahoochee Technical College
College of Coastal Georgia Lori Smith Daniel Brian Nichols
Barbara R. Heard American River College Seton Hall University
Atlantic Cape Community College Vetaley Stashenko Kevin Sorensen
Nazanin Hebel Palm Beach State College, Belle Glade Snow College
Houston Community College—Northwest Jennifer Swartz Sandra L. Specht
Amy Helms Pikes Peak Community College Sinclair Community College
Collin College
David T. Jenkins
University of Alabama at Birmingham

xvi
Acknowledgments
As has been the case with all previous editions, I am ever more
cognizant that this book is a team effort. I am deeply grateful
once again to Kelsey Churchman of Pearson Science and to the
team she gathered to produce the fifth edition. Kelsey, dedicated
project manager Lauren Beebe, and invaluable program
manager Chriscelle Palaganas helped develop the vision for
this fifth edition, generating ideas to make it more effective and
compelling. As project manager, Lauren also had the unenviable
task of coordinating everything and keeping me on track—thank
you, Lauren, for being understanding, patient, and lenient,
especially when I misplaced a deadline. Kari Hopperstead was
invaluable in developmental editing. I am grateful.
Thank you to Barbara Yien, project editor of the first two
editions, for years of support and for introducing me to
chocolate truffles. I am excited for your growing family and
new responsibilities! I am grateful to Frank Ruggirello for his
unflagging encouragement and support of my work and this
book; enjoy your new adventures! I am also indebted to Daryl
Fox, whose early support for this book never wavered.
Anita Wagner Hueftle—the eagle-eyed—edited the manuscript
thoroughly and meticulously, suggesting important changes
for clarity, accuracy, and consistency. The incomparable Kelly
Murphy did a magnificently superb job as art development
editor, helping to conceptualize new illustrations and
suggesting ways to improve the art overall—thank you,
Kelly for taking the original art of my friend Ken Probst
and enhancing this book’s amazingly beautiful biological
illustrations. My thanks to Lachina for rendering the art in this
edition. Andrea Stefanowicz and Lumina Datamatics expertly
guided the project through production. Andrea, thank you
for meticulously improving the text. Maureen “Mo” Spuhler
remains the most amazing photo researcher. I am in your debt,
“Molybdenum.” Rich Robison and Brent Selinger supplied
many of the text’s wonderful and unique micrographs. Emily
Friel created the beautiful interior design and the stunning
cover.
Thanks to Michéle Shuster and Amy Helms for their work on
the media and print supplements for this edition. Special thanks
are due to Lauren Beebe and Andrea Stefanowicz for managing
the supplements, to Kyle Doctor in production for his work on
the Instructor’s Resource DVD, and to Joe Mochnick for his
management of the extraordinary array of media resources for
students and instructors, especially MasteringMicrobiology®.
Thanks also to Jordan Roeder, RN, and Nan Kemp and for
their administrative, editorial, and research assistance. I am
grateful to Neena Bali and now Lauren Harp in Marketing; they
lead the amazing Pearson sales representatives to do a terrific
job of keeping in touch with the professors and students who
provide so many wonderful suggestions for this textbook. As
always, I am humbled, inspired, and encouraged by the sales
representatives; your role on the team deserves more gratitude
than I can express here or with citrus fruit.
I am especially grateful to Phil Mixter of Washington State
University, Mary Jane Niles of the University of San Francisco,
Bronwen Steele of Estrella Mountain Community College, Jan
Miller of American River College, and Jane Reece for their
expertise and advice.
I am further indebted to Sam Schwarzlose for his excellent work
on the Video Tutor assessments and to Terry Austin for lending
his technical expertise to the project.
On the home front: Thank you, Jennie and Nick Knapp,
Elizabeth Bauman, Jeremy Bauman, Larry Latham, Josh Wood,
and Mike Isley. You keep me even-keeled. My wife Michelle
deserves more recognition than I can possibly express: “Many
are noble, but you excel them all.” Thank you.
Robert W. Bauman
Amarillo, Texas
xvii
Table of Contents

1
Acids and Bases 36
Salts 38
Organic Macromolecules 38
A Brief Functional Groups 39
Lipids 40
History of Carbohydrates 42
Proteins 44
­Microbiology 1 Nucleotides and Nucleic Acids 48
The Early Years of Chapter Summary 51 • Questions for Review 52  
Microbiology 2 Critical Thinking 53 • Concept Mapping 54
What Does Life Really Look
Like? 2

3
How Can Microbes Be
Classified? 3
The Golden Age of Microbiology 7
Does Microbial Life Spontaneously Generate? 7
What Causes Fermentation? 10
Cell Structure and
What Causes Disease? 11
How Can We Prevent Infection and Disease? 15
Function 55
The Modern Age of Microbiology 18 Processes of Life 56
What Are the Basic Chemical Reactions of Life? 18 Prokaryotic and Eukaryotic
How Do Genes Work? 18 Cells: An Overview 57
What Roles Do Microorganisms Play in the Environment? 20 External Structures of Bacterial
How Do We Defend Against Disease? 20 Cells 59
What Will the Future Hold? 21 Glycocalyces 59
Chapter Summary 22 • Questions for Review 22  
Flagella 59
Critical Thinking 24 • Concept Mapping 25 Fimbriae and Pili 62
Bacterial Cell Walls 63

2
Gram-Positive Bacterial Cell Walls 64
Gram-Negative Bacterial Cell Walls 66
Bacteria Without Cell Walls 66
Bacterial Cytoplasmic Membranes 66
The Chemistry of Structure 66
­Microbiology 26 Function 67
Cytoplasm of Bacteria 72
Atoms 27 Cytosol 72
Atomic Structure 27 Inclusions 72
Isotopes 27 Endospores 73
Electron Configurations 28 Nonmembranous Organelles 74
Chemical Bonds 30 External Structures of Archaea 74
Nonpolar Covalent Bonds 30 Glycocalyces 75
Polar Covalent Bonds 31 Flagella 75
Ionic Bonds 32 Fimbriae and Hami 75
Hydrogen Bonds 33 Archaeal Cell Walls and Cytoplasmic Membranes 76
Chemical Reactions 34 Cytoplasm of Archaea 76
Synthesis Reactions 34
External Structure of Eukaryotic Cells 77
Decomposition Reactions 34
Exchange Reactions 35 Glycocalyces 77
Water, Acids, Bases, and Salts 35 Eukaryotic Cell Walls and Cytoplasmic Membranes 77
Water 35
xviii
 Table of Contents xix

Cytoplasm of Eukaryotes 79 Carbohydrate Catabolism 131

Flagella 79 Glycolysis 131
Cilia 79 Cellular Respiration 133
Other Nonmembranous Organelles 80 Pentose Phosphate Pathway 139
Membranous Organelles 81 Fermentation 139
Endosymbiotic Theory 85 Other Catabolic Pathways 141
Lipid Catabolism 141
Chapter Summary 87 • Questions for Review  89  
Protein Catabolism 142
Critical Thinking 92 • Concept Mapping 93
Photosynthesis 143
Chemicals and Structures 143

4
Light-Dependent Reactions 144
Light-Independent Reactions 145
Other Anabolic Pathways 148
Carbohydrate Biosynthesis 148
Microscopy, Lipid Biosynthesis 149
Staining, and Amino Acid Biosynthesis 149
Nucleotide Biosynthesis 150
­Classification 94 Integration and Regulation of Metabolic Functions 151

Units of Measurement 95 Chapter Summary 153 • Questions for Review  155  

Microscopy 96 Critical Thinking 157 • Concept Mapping 159
General Principles of
Microscopy 96
Light Microscopy 97
Electron Microscopy 102
Probe Microscopy 103
6
Staining 104
Preparing Specimens for Staining 104
Microbial Nutrition
Principles of Staining 106 and Growth 160
Simple Stains 106
Differential Stains 107 Growth
Special Stains 108 Requirements 161
Staining for Electron Microscopy 109 Nutrients: Chemical
Classification and Identification of Microorganisms 110 and Energy
Requirements 161
Linnaeus and Taxonomic Categories 111
Physical
Domains 113
Requirements 164
Taxonomic and Identifying Characteristics 114
Associations and Biofilms 167
Taxonomic Keys 117
Culturing Microorganisms 169
Chapter Summary 118 • Questions for Review  119   Clinical Sampling 170
Critical Thinking 121 • Concept Mapping 121 Obtaining Pure Cultures 171
Culture Media 172
Special Culture Techniques 176

5
Preserving Cultures 176
Growth of Microbial Populations 177
Generation Time 178
Microbial Mathematical Considerations in Population Growth 178
Phases of Microbial Population Growth 178
Metabolism 122 Continuous Culture in a Chemostat 180
Measuring Microbial Reproduction 180
Basic Chemical Reactions
Underlying Metabolism 123 Chapter Summary 185 • Questions for Review  187  
Catabolism and Anabolism 123 Critical Thinking 188 • Concept Mapping 189
Oxidation and Reduction
Reactions 124
ATP Production and Energy
Storage 124
The Roles of Enzymes
in Metabolism 125
xx Table of Contents

Separating DNA Molecules: Gel Electrophoresis and the

7 Southern Blot 243

DNA Microarrays 243
Inserting DNA into Cells 244
Microbial Applications of Recombinant DNA Technology 245
Genetics 190 Genetic Mapping 245
Microbial Community Studies 248
The Structure and Pharmaceutical and Therapeutic Applications 249
Replication of Agricultural Applications 251
Genomes 191 The Ethics and Safety of Recombinant DNA
The Structure of Nucleic Technology 252
Acids 191
The Structure of Prokaryotic Genomes 191
Chapter Summary 254 • Questions for Review  255  
The Structure of Eukaryotic Genomes 193
Critical Thinking 256 • Concept Mapping 257
DNA Replication 195
Gene Function 200
The Relationship Between Genotype and Phenotype 200
The Transfer of Genetic Information 200
The Events in Transcription 201
9
Translation 204 Controlling
Regulation of Genetic Expression 209
Mutations of Genes 213 Microbial
Types of Mutations 214
Effects of Point Mutations 215
Growth in the
Mutagens 215
Frequency of Mutation 217
Environment 258
DNA Repair 218 Basic Principles of Microbial
Identifying Mutants, Mutagens, and Carcinogens 218 Control 259
Genetic Recombination and Transfer 221 Terminology of Microbial
Horizontal Gene Transfer Among Prokaryotes 222 Control 259
Transposons and Transposition 227 Microbial Death Rates 260
Action of Antimicrobial Agents 261
Chapter Summary 229 • Questions for Review  230   The Selection of Microbial Control Methods 261
Critical Thinking 233 • Concept Mapping 234 Factors Affecting the Efficacy of Antimicrobial Methods 261
Biosafety Levels 263
Physical Methods of Microbial Control 264

8 Heat-Related Methods 264

Refrigeration and Freezing 267
Desiccation and Lyophilization 267
Recombinant Filtration 268
Osmotic Pressure 269
DNA Radiation 269
Chemical Methods of Microbial Control 271
­Technology 235 Phenol and Phenolics 272
The Role of Recombinant Alcohols 272
DNA Technology in Halogens 272
Biotechnology 236 Oxidizing Agents 274
Surfactants 274
The Tools of Recombinant Heavy Metals 275
DNA Technology 236 Aldehydes 275
Mutagens 236 Gaseous Agents 275
The Use of Reverse Transcriptase to Synthesize cDNA 237 Enzymes 276
Synthetic Nucleic Acids 237 Antimicrobial Drugs 276
Restriction Enzymes 238 Methods for Evaluating Disinfectants and Antiseptics 276
Vectors 240 Development of Resistant Microbes 278
Gene Libraries 241
Techniques of Recombinant DNA Technology 241 Chapter Summary 278 • Questions for Review  279  
Multiplying DNA In Vitro: The Polymerase Chain Critical Thinking 281 • Concept Mapping 282
Reaction 241
Selecting a Clone of Recombinant Cells 243
 Table of Contents xxi

Survey of Bacteria 325

10 Deeply Branching and Phototrophic Bacteria 325

Low G + C Gram-Positive Bacteria 327
High G + C Gram-Positive Bacteria 330
Controlling Gram-Negative Proteobacteria 332
Other Gram-Negative Bacteria 341
Microbial Growth Chapter Summary 342 • Questions for Review  343  
in the Body: Critical Thinking 345 • Concept Mapping 345
Antimicrobial
Drugs 283
The History of Antimicrobial
12
Agents 284
Mechanisms of Antimicrobial
Characterizing
Action 285 and Classifying
Inhibition of Cell Wall Synthesis 286
Inhibition of Protein Synthesis 288 Eukaryotes 346
Disruption of Cytoplasmic Membranes 289
General Characteristics of
Inhibition of Metabolic Pathways 290
Eukaryotic Organisms 347
Inhibition of Nucleic Acid Synthesis 291
Prevention of Virus Attachment, Entry, or Uncoating 293 Reproduction of
Eukaryotes 347
Clinical Considerations in Prescribing Antimicrobial Classification of Eukaryotic
Drugs 293 Organisms 350
Spectrum of Action 293
Protozoa 351
Effectiveness 294
Routes of Administration 296 Distribution of Protozoa 351
Safety and Side Effects 297 Morphology of Protozoa 352
Nutrition of Protozoa 352
Resistance to Antimicrobial Drugs 298 Reproduction of Protozoa 353
The Development of Resistance in Populations 298 Classification of Protozoa 353
Mechanisms of Resistance 298
Fungi 357
Multiple Resistance and Cross Resistance 301
Retarding Resistance 301 The Significance of Fungi 358
Morphology of Fungi 358
Chapter Summary 313 • Questions for Review  314   Nutrition of Fungi 359
Critical Thinking 315 • Concept Mapping 316 Reproduction of Fungi 360
Classification of Fungi 361
Lichens 364

11
Algae 366
Distribution of Algae 366
Morphology of Algae 366
Reproduction of Algae 366
Characterizing Classification of Algae 367
and Classifying Water Molds 369
Other Eukaryotes of Microbiological Interest: Parasitic
Prokaryotes 317 Helminths and Vectors 370
General Characteristics of Arachnids 370
Prokaryotic Organisms 318 Insects 370
Morphology of Prokaryotic Chapter Summary 372 • Questions for Review  373  
Cells 318 Critical Thinking 375 • Concept Mapping 376
Endospores 318
Reproduction of Prokaryotic Cells 319
Arrangements of Prokaryotic Cells 320
Modern Prokaryotic Classification 322
Survey of Archaea 322
Extremophiles 323
Methanogens 325
xxii Table of Contents

The Invasion and Establishment of Microbes in Hosts:

13 Infection 411
Exposure to Microbes: Contamination and Infection
Portals of Entry 411
411

Characterizing The Role of Adhesion in Infection 413

and ­Classifying The Nature of Infectious Disease 414

Manifestations of Disease: Symptoms, Signs, and
Viruses, Viroids, Syndromes 414
Causation of Disease: Etiology 415
and ­Prions 377 Virulence Factors of Infectious Agents 417
The Stages of Infectious Diseases 420
Characteristics of Viruses 378 The Movement of Pathogens Out of Hosts: Portals of
Genetic Material of Viruses 379 Exit 422
Hosts of Viruses 379
Sizes of Viruses 380 Modes of Infectious Disease Transmission 422
Capsid Morphology 380 Contact Transmission 422
Viral Shapes 380 Vehicle Transmission 422
The Viral Envelope 382 Vector Transmission 423
Classification of Viruses 383 Classification of Infectious Diseases 424
Viral Replication 385 Epidemiology of Infectious Diseases 426
Lytic Replication of Bacteriophages 385 Frequency of Disease 426
Lysogenic Replication of Bacteriophages 388 Epidemiological Studies 427
Replication of Animal Viruses 388 Hospital Epidemiology: Healthcare-Associated (Nosocomial)
Infections 429
The Role of Viruses in Cancer 393
Epidemiology and Public Health 431
Culturing Viruses in the Laboratory 394
Culturing Viruses in Mature Organisms 395 Chapter Summary 434 • Questions for Review  435  
Culturing Viruses in Embryonated Chicken Eggs 396 Critical Thinking 437 • Concept Mapping 438
Culturing Viruses in Cell (Tissue) Culture 396
Are Viruses Alive? 397
Other Parasitic Particles: Viroids and Prions
Characteristics of Viroids 397
Characteristics of Prions 398
397
15
Chapter Summary 401 • Questions for Review  402   Innate
Critical Thinking 403 • Concept Mapping 404 Immunity 439
An Overview of the Body’s

14
Defenses 440
The Body’s First Line of
Defense 440
Infection, The Role of Skin in Innate
Immunity 440
Infectious The Role of Mucous Membranes
in Innate Immunity 441
Diseases, and The Role of the Lacrimal Apparatus in Innate Immunity 442
The Role of Normal Microbiota in Innate Immunity 442
Epidemiology 405 Other First-Line Defenses 443
Symbiotic Relationships Between The Body’s Second Line of Defense 444
Microbes and Their Hosts 406 Defense Components of Blood 444
Types of Symbiosis 406 Phagocytosis 447
Normal Microbiota in Hosts 407 Nonphagocytic Killing 448
How Normal Microbiota Become Opportunistic Nonspecific Chemical Defenses Against Pathogens 449
Pathogens 408 Inflammation 454
Fever 457
Reservoirs of Infectious Diseases of Humans 410
Animal Reservoirs 410 Chapter Summary 459 • Questions for Review  460  
Human Carriers 411 Critical Thinking 462 • Concept Mapping 463
Nonliving Reservoirs 411
 Table of Contents xxiii

Labeled Antibody Tests 507

16 Point-of-Care Testing 511

Chapter Summary 512 • Questions for Review  513  
Adaptive Critical Thinking 515 • Concept Mapping 516

Immunity 464
Overview of Adaptive
Immunity 465
18
Elements of Adaptive
Immunity 466 Immune
The Tissues and Organs of the
Lymphatic System 466
Disorders 517
Antigens 468 Hypersensitivities 518
Preparation for an Adaptive Type I (Immediate)
Immune Response 469 Hypersensitivity 518
T Lymphocytes (T Cells) 471 Type II (Cytotoxic)
B Lymphocytes (B Cells) and Antibodies 474 Hypersensitivity 522
Immune Response Cytokines 480 Type III (Immune Complex–
Cell-Mediated Immune Responses 481 Mediated)
Activation of Cytotoxic T Cell Clones and Their Functions 481 Hypersensitivity 525
The Perforin-Granzyme Cytotoxic Pathway 483 Type IV (Delayed or Cell-Mediated) Hypersensitivity 527
The CD95 Cytotoxic Pathway 483 Autoimmune Diseases 531
Memory T Cells 483 Causes of Autoimmune Diseases 531
T Cell Regulation 484 Examples of Autoimmune Diseases 531
Antibody Immune Responses 484 Immunodeficiency Diseases 532
Inducement of T-Dependent Antibody Immunity with Clonal Primary Immunodeficiency Diseases 533
Selection 484 Acquired Immunodeficiency Diseases 533
Memory Cells and the Establishment of Immunological
Memory 486 Chapter Summary 534 • Questions for Review  535  
Types of Acquired Immunity 487 Critical Thinking 537 • Concept Mapping 537
Naturally Acquired Active Immunity 487
Naturally Acquired Passive Immunity 487
Artificially Acquired Active Immunity 488
Artificially Acquired Passive Immunotherapy 488 19
• Questions for Review  491  
Chapter Summary 490
Critical Thinking 493 • Concept Mapping 494 Pathogenic
Gram-Positive
­Bacteria 538
17 Staphylococcus 539
Structure and
Immunization Physiology 539
Pathogenicity 539
and Immune Epidemiology 540
­Testing 495 Staphylococcal
Diseases 541
Immunization 496 Diagnosis, Treatment,
Brief History of Immunization 496 and Prevention 542
Active Immunization 497 Streptococcus 543
Passive Immunotherapy 502 Group A Streptococcus: Streptococcus pyogenes 544
Serological Tests That Use Group B Streptococcus: Streptococcus agalactiae 548
Antigens and Corresponding Antibodies 503 Other Beta-Hemolytic Streptococci 549
Precipitation Tests 504 Alpha-Hemolytic Streptococci: The Viridans
Turbidimetric and Nephelometric Tests 505 Group 549
Agglutination Tests 505 Streptococcus pneumoniae 549
Neutralization Tests 506 Enterococcus 551
The Complement Fixation Test 507 Structure and Physiology 551
xxiv Table of Contents

Pathogenesis, Epidemiology, and Diseases 551 Bordetella 598

Diagnosis, Treatment, and Prevention 552 Burkholderia 600
Bacillus 553 Pseudomonads 600
Structure, Physiology, and Pathogenicity 553 Francisella 602
Epidemiology 553 Legionella 603
Disease 553 Coxiella 604
Diagnosis, Treatment, and Prevention 554 Pathogenic, Gram-Negative, Anaerobic Bacilli 605
Clostridium 554 Bacteroides 605
Clostridium perfringens 555 Prevotella 605
Clostridium difficile 555
Chapter Summary 606 • Questions for Review  607  
Clostridium botulinum 556
Critical Thinking 609 • Concept Mapping 610
Clostridium tetani 558
Listeria 560

21
Mycoplasmas 560
Mycoplasma pneumoniae 561
Other Mycoplasmas 564
Corynebacterium 564
Pathogenesis, Epidemiology, and Disease 565
Rickettsias,
Diagnosis, Treatment, and Prevention 565 Chlamydias,
Mycobacterium 565
Tuberculosis 566 ­Spirochetes,
Leprosy 566
Other Mycobacterial Infections 567 and Vibrios 611
Propionibacterium 570 Rickettsias 612
Nocardia and Actinomyces 572 Rickettsia 612
Nocardia asteroides 572 Orientia tsutsugamushi 613
Actinomyces 572 Ehrlichia and Anaplasma 616
Chlamydias 617
Chapter Summary 573 • Questions for Review  575   Chlamydia trachomatis 617
Critical Thinking 576 • Concept Mapping 577 Chlamydophila
pneumoniae 620
Chlamydophila psittaci 620

20 Spirochetes 620
Treponema 621
Borrelia 624
Pathogenic Leptospira 627
Pathogenic Gram-Negative Vibrios 628
Gram-Negative Vibrio 628
Cocci and Campylobacter jejuni 630
Helicobacter pylori 630
Bacilli 578 Chapter Summary 633 • Questions for Review  634  
Pathogenic Gram-Negative Critical Thinking 636 • Concept Mapping 637
Cocci: Neisseria 579
Structure and Physiology of
Neisseria 579
The Gonococcus: Neisseria gonorrhoeae 580
The Meningococcus: Neisseria meningitidis 582
22
Pathogenic, Gram-Negative, Facultatively Anaerobic
Bacilli 583
Pathogenic
The Enterobacteriaceae: An Overview 583 Fungi 638
Coliform Opportunistic Enterobacteriaceae 586
Noncoliform Opportunistic Enterobacteriaceae 590 An Overview of Medical
Truly Pathogenic Enterobacteriaceae 591 Mycology 639
The Pasteurellaceae 595 The Epidemiology of
Mycoses 639
Pathogenic, Gram-Negative, Aerobic Bacilli 596
Categories of Fungal Agents:
Bartonella 597
True Fungal Pathogens and
Brucella 597
Opportunistic Fungi 639
 Table of Contents xxv

Clinical Manifestations of Fungal Diseases 640

The Diagnosis of Fungal Infections 640
Antifungal Therapies 641
Antifungal Vaccines 641
24
Systemic Mycoses Caused by Pathogenic Fungi 642 Pathogenic
Histoplasmosis 643
Blastomycosis 644 DNA
Coccidioidomycosis 645
Paracoccidioidomycosis 647
Viruses 700
Systemic Mycoses Caused by Opportunistic Fungi 647 Poxviridae 701
Pneumocystis Pneumonia 647 Smallpox 701
Candidiasis 648 Molluscum
Aspergillosis 648 Contagiosum 703
Cryptococcosis 652 Other Poxvirus
Zygomycoses 654 Infections 703
The Emergence of Fungal Opportunists in AIDS Patients 654 Herpesviridae 704
Superficial, Cutaneous, and Subcutaneous Mycoses 655 Infections of Human
Superficial Mycoses 655 Herpesvirus 1
Cutaneous and Subcutaneous Mycoses 656 and 2 705
Fungal Intoxications and Allergies 659 Human Herpesvirus 3
Mycotoxicoses 659 (Varicella-Zoster Virus)
Mushroom Poisoning (Mycetismus) 659 Infections 708
Allergies to Fungi 659 Human Herpesvirus 4 (Epstein-Barr Virus) Infections 710
Human Herpesvirus 5 (Cytomegalovirus) Infections 711
Chapter Summary 661 • Questions for Review 662   Other Herpesvirus Infections 712
Critical Thinking 664 • Concept Mapping 665 Papillomaviridae and Polyomaviridae 713
Papillomavirus Infections 713
Polyomavirus Infections 713

23 Adenoviridae 715
Hepadnaviridae 716
Hepatitis B Infections 717
Parasitic The Role of Hepatitis B Virus in Hepatic Cancer 719

Protozoa, Parvoviridae 719

• Questions for Review

Helminths, Chapter Summary 720
Critical Thinking 723 • Concept Mapping 724
721  

and Arthropod
Vectors 666
Parasitology 667 25
Arthropod Vectors 668
Protozoan Parasites of
Pathogenic
Humans 668 RNA
Ciliates 668
Amoebas 669 Viruses 725
Flagellates 670
Apicomplexans 675 Naked, Positive ssRNA
Viruses: Picornaviridae,
Helminthic Parasites of Humans 683
Caliciviridae, Astroviridae,
Cestodes 683
and Hepeviridae 726
Trematodes 687
Common Colds Caused by
Nematodes 689
Rhinoviruses 726
Chapter Summary 694 • Questions for Review 696   Diseases of
Critical Thinking 698 • Concept Mapping 699 Enteroviruses 727
Hepatitis A 730
Acute Gastroenteritis 730
Hepatitis E 731
xxvi Table of Contents

Enveloped, Positive ssRNA Viruses: Togaviridae, Industrial Products of Microorganisms 777

Flaviviridae, and Coronaviridae 732 Water Treatment 779
Diseases of +RNA Arboviruses 732 Environmental Microbiology 786
Other Diseases of Enveloped +ssRNA Viruses 736 Microbial Ecology 786
Enveloped, Positive ssRNA Viruses with Reverse Bioremediation 787
Transcriptase: Retroviridae 739 The Problem of Acid Mine Drainage 787
Oncogenic Retroviruses (Deltaretrovirus) 740 The Roles of Microorganisms in Biogeochemical Cycles 789
Immunosuppressive Retroviruses (Lentivirus) and Acquired Soil Microbiology 791
Immunodeficiency Syndrome 740 Aquatic Microbiology 793
Enveloped, Unsegmented, Negative ssRNA Viruses: Biological Warfare and Bioterrorism 795
Paramyxoviridae, Rhabdoviridae, and Filoviridae 747 Assessing Microorganisms as Potential Agents of Warfare
Measles 747 or Terror 795
Diseases of Parainfluenza Virus 749 Known Microbial Threats 796
Mumps 750 Defense Against Bioterrorism 797
Disease of Respiratory Syncytial Virus 750 The Roles of Recombinant Genetic Technology in
Rabies 751 Bioterrorism 797
Hemorrhagic Fevers 753
Chapter Summary 798 • Questions for Review 800  
Enveloped, Segmented, Negative ssRNA Viruses: Critical Thinking 803 • Concept Mapping 804
Orthomyxoviridae, Bunyaviridae, and Arenaviridae 753
Influenza 753 Answers to Questions for Review A-1
Diseases of Bunyaviruses 757
Diseases of Arenaviruses 757 Glossary G-1
Naked, Segmented dsRNA Viruses: Reoviridae 760 Credits C-1
Rotavirus Infections 760 Index I-1
Coltivirus Infections 761
Chapter Summary 762 • Questions for Review 764  
Critical Thinking 766 • Concept Mapping 767

26
Applied and
Environmental
Microbiology 768
Food Microbiology 769
The Roles of Microorganisms in
Food Production 769
The Causes and Prevention of
Food Spoilage 772
Foodborne Illnesses 776
Industrial Microbiology 776
The Roles of Microbes in Industrial
Fermentations 776
Feature Boxes
BENEFICIALMICROBES
Bread, Wine, and Beer 7
Architecture-Preserving Bacteria 37
Plastics Made Perfect? 72
Glowing Viruses 111
A Nuclear Waste–Eating Microbe? 167
Life in a Hot Tub 198
Hard to Swallow? 273
Probiotics: Using Live Microorganisms to Treat or Prevent
Disease 298
HIGHLIGHT
Emerging and Reemerging Diseases: “The New Normal” 8
Biofilms: Slime Matters 63
Studying Biofilms in Plastic “Rocks” 102
What’s That Fishy Smell? 142
Hydrogen-Loving Microbes in Yellowstone’s Hot Springs 164
How Do You “Fix” a Mosquito? 239
The Human Microbiome Project 250
Microbes in Sushi? 269
Microbe Altruism: Why Do They Do It? 285
EMERGING DISEASE CASE STUDY
Variant Creutzfeldt-Jakob
Disease 20
Necrotizing Fasciitis 117
Vibrio vulnificus Infection 211
Acanthamoeba Keratitis 263
Community-Associated MRSA  299
Pertussis 337
Aspergillosis 363
A Microtube of Superglue 335
Fungi for $10,000 a Pound 364
Good Viruses? Who Knew? 381
Prescription Bacteriophages? 387
A Bioterrorist Worm 407
Cowpox: To Vaccinate or Not to Vaccinate? 502
Microbes to the Rescue? 554
Eliminating Dengue 735
Oil-Eating Microbes to the Rescue in the Gulf 788
Your Teeth Might Make You Fat 330
Lymphocyte Receptor Diversity: The Star of the Show 476
Can Pets Help Decrease Children’s Allergy Risks? 518
When Kissing Triggers Allergic Reactions 522
Does “Killer Mold” Exist? 660
Catch a Cold and Catch Obesity? 716
Nipah Virus: From Pigs to Humans 748
Could Bioterrorists Manufacture Viruses from Scratch? 798
Chikungunya 394 Babesiosis 682
Hantavirus Pulmonary Snail Fever Reemerges
Syndrome 433 in China 690
Microsporidiosis 488 Monkeypox 704
Buruli Ulcer 570 Norovirus in the Dorm 731
Melioidosis 602 Tick-Borne Encephalitis 734
A New Cause of Spots 613 H1N1 Influenza 756
Pulmonary Blastomycosis 645 Attack in the Lake 780
xxvii
xxviii Feature Boxes
CLINICAL CASE STUDY
Remedy for Fever or Prescription for
Death? 16
Can Spicy Food Cause Ulcers? 21
Raw Oysters and Antacids: A Deadly
Mix? 38
The Big Game 68
Cavities Gone Wild 169
Boils in the Locker Room 179
Deadly Horizontal Gene Transfer 228
Antibiotic Overkill 294
Battling the Enemy 296
Tough Decision 302
Invasion from Within or Without? 400
MICROBE AT A GLANCE
Streptococcus pneumoniae 550
Clostridium botulinum 558
Neisseria gonorrhoeae 581
Treponema pallidum 622
Helicobacter pylori 631
Disease in Depth
Necrotizing Fasciitis 546
Listeriosis 562
Tuberculosis 568
Bacterial Urinary Tract Infections 588
A Deadly Carrier 411
TB in the Nursery 424
Legionella in the Produce Aisle 432
Evaluating an Abnormal CBC 447
The Stealth Invader 454
The First Time’s Not the Problem 527
A Fatal Case of Methicillin-Resistant
Staphylococcus aureus (MRSA) 543
This Cough Can Kill 566
A Painful Problem 586
A Heart-Rending Experience 587
A Sick Camper 595
When “Health Food” Isn’t 600
Histoplasma capsulatum 644
Aspergillus 652
Orthopoxvirus variola
(Smallpox Virus) 703
Rocky Mountain Spotted Fever 614
Candidiasis 650
Giardiasis 676
Malaria 678
Nightmare on the Island 603
The Case of the Lactovegetarians 632
What’s Ailing the Bird Enthusiast? 653
Disease from a Cave 654
A Protozoan Mystery 673
A Sick Soldier 675
A Fluke Disease? 689
Grandfather’s Shingles 711
A Child with Warts 715
The Eyes Have It 719
A Threat from the Wild 752
The Sick Addict 756
Adenovirus 717
Lentivirus human immunodeficiency
virus (HIV) 743
Morbillivirus measles virus 749
Papillomas 714
Ebola 754
Influenza 758
1
A Brief History
of Microbiology

LIFE AS WE KNOW IT WOULD NOT EXIST

without microorganisms. Plants depend
on microorganisms to help them obtain
the nitrogen they need for survival.
Animals such as cows and sheep need
microbes in order to digest the carbohy-
drates in their plant-based diets. Ecosys-
tems rely on microorganisms to enrich
soil, degrade wastes, and support life. We
use microorganisms to make wine and
cheese and to develop vaccines and anti-
biotics. Through recombinant DNA tech-
nology (genetic engineering), we are now
able to harness the power of these small
microbes to do big jobs like mass pro-
ducing important pharmaceuticals such
as blood-clotting factor VIII and insulin for
patients who desperately need them.
The human body is home to trillions
of microorganisms, many of which help
keep us healthy. Microorganisms are an
essential part of our lives. Of course,
some microorganisms do cause harm to
us, from the common cold to more seri-
ous diseases such as tuberculosis, malaria,
and AIDS. The threats of bioterrorism and
new or re-emerging infectious diseases
are real.
This textbook explores the roles—
both beneficial and harmful—that micro-
organisms play in our lives, as well as their
sophisticated structures and processes.
Aquatic This chapter will explore not only the
microorganisms, history of microbiology, but how new
such as these, ­discoveries have led to a number of new
thrilled early disciplines within the field of microbiology.
microscopists with We begin with the invention of crude
their beauty and microscopes that revealed, for the first
antics. time, the existence of this miraculous,
miniature world.
2 CHAPTER 1 A Brief History of Microbiology

Science is the study of nature that proceeds by posing ques-

tions about observations. Why are there seasons? What is the
function of the nodules at the base of this plant? Why does this
bread taste sour? What does plaque from between teeth look
like when magnified? What causes the spread of diseases?
Many early written records show that people have always
asked questions like these. For example, the Greek physician
Hippocrates (ca. 460–ca. 377 b.c.) wondered whether there is
a link between environment and disease, and the Greek histo-
rian Thucydides (ca. 460–ca. 404 b.c.) questioned why he and
other survivors of the plague could have close contact with vic-
tims and not fall ill again. For many centuries, the answers to
these and other fundamental questions about the nature of life
remained largely unanswered. But about 350 years ago, the in-
vention of the microscope began to provide some clues.
In this chapter, we’ll see how one man’s determination to
answer a fundamental question about the nature of life—What
does life really look like?—led to the birth of a new science
▲ FIGURE 1.1 Antoni van Leeuwenhoek. Leeuwenhoek reported
called microbiology. We’ll then see how the search for answers the existence of protozoa in 1674 and of bacteria in 1676. Why did
to other questions, such as those concerning spontaneous ­Leeuwenhoek discover protozoa before bacteria?
generation, the reason fermentation occurs, and the cause of Figure 1.1 Protozoa are generally larger than bacteria.
disease, prompted advances in this new science. Finally, we’ll
look briefly at some of the key questions microbiologists are
asking today.
he spent hours examining, reexamining, and recording every
detail of each object he observed.
Making and looking through his simple microscopes, really
The Early Years of Microbiology no more than magnifying glasses, became the overwhelming
passion of his life. His enthusiasm and dedication are evident
The early years of microbiology brought the first observations
from the fact that he sometimes personally extracted the metal
of microbial life and the initial efforts to organize them into log-
ical classifications.
Lens Specimen holder
What Does Life Really Look Like?
LEARNI N G OUT COM E S

1.1 Describe the world-changing scientific contributions of

Leeuwenhoek.
1.2 Define microbes in the words of Leeuwenhoek and as we
know them today.

A few people have changed the world of science forever. We’ve

all heard of Galileo, Newton, and Einstein, but the list also in-
cludes Antoni van Leeuwenhoek (lā´vĕn-huk; 1632–1723), a
Dutch tailor, merchant, and lens grinder, and the man who first
discovered the bacterial world (FIGURE 1.1).
Leeuwenhoek was born in Delft, the Netherlands, and lived
most of his 90 years in the city of his birth. What set Leeuwen-
hoek apart from many other men of his generation was an in-
satiable curiosity coupled with an almost stubborn desire to
do everything for himself. His journey to fame began simply
enough, when as a cloth merchant he needed to examine the ▲ FIGURE 1.2 Reproduction of Leeuwenhoek’s microscope.
quality of cloth. Rather than merely buying a magnifying lens, This simple device is little more than a magnifying glass with screws for
he learned to make glass lenses of his own (FIGURE 1.2). Soon manipulating the specimen, yet with it, Leeuwenhoek changed the way
we see our world. The lens, which is convex on both sides, is about the
he began asking, “What does it really look like?” of everything size of a pinhead. The object to be viewed was mounted either directly
in his world: the stinger of a bee, the brain of a fly, the leg of a on the specimen holder or inside a small glass tube, which was then
louse, a drop of blood, flakes of his own skin. To find answers, mounted on the specimen holder.

for a microscope from ore. Further, he often made a new
microscope for each specimen, which remained mounted so
that he could view it again and again. Then one day, he turned
a lens onto a drop of water. We don’t know what he expected to
see, but certainly he saw more than he had anticipated. As he
reported to the Royal Society of London1 in 1674, he was sur-
prised and delighted by
some green streaks, spirally wound serpent-wise, and or-
derly arranged. . . . Among these there were, besides, very
many little animalcules, some were round, while others a
bit bigger consisted of an oval. On these last, I saw two little
legs near the head, and two little fins at the hind most end
of the body. . . . And the motion of most of these animal-
cules in the water was so swift, and so various, upwards,
downwards, and round about, that ’twas wonderful to see.2
Leeuwenhoek had discovered the previously unknown micro-
bial world, which today we know to be populated with tiny an-
imals, fungi, algae, and single-celled protozoa (FIGURE 1.3). In a
later report to the Royal Society, he noted that
the number of these animals in the plaque of a man’s teeth,
are so many that I believe they exceed the number of men
in a kingdom. . . . in a quantity of matter no bigger than the
1/100 part of a [grain of] sand.
From the figure accompanying his report and the precise
description of the size of these organisms from between
his teeth, we know that Leeuwenhoek was reporting the
existence of ­bacteria. By the end of the 19th century, Leeuwen-
hoek’s “beasties,” as he sometimes dubbed them, were called
microorganisms, and today we also know them as microbes.
LM
50 μm
▲ FIGURE 1.3 The microbial world. Leeuwenhoek reported seeing
a scene very much like this, full of numerous fantastic, cavorting creatures.
1The
Royal Society of London for the Promotion of Natural Knowledge, granted a royal
charter in 1662, is one of the older and more prestigious scientific groups in Europe.
2Antony von Leeuwenhoek, in a letter to the Royal Society of London for the Promotion of
Natural Knowledge.
CHAPTER 1 A Brief History of Microbiology 3
Both terms include all organisms that are too small to be seen
without a microscope.
Because of the quality of his microscopes, his profound ob-
servational skills, his detailed reports over a 50-year period, and
his report of the discovery of many types of microorganisms,
Antoni van Leeuwenhoek was elected to the Royal Society in
1680. He was one of the more famous scientists of his time.
How Can Microbes Be Classified?
L EA RN IN G OU TCOM ES
1.3 List six groups of microorganisms.
1.4 Explain why protozoa, algae, and nonmicrobial parasitic
worms are studied in microbiology.
1.5 Differentiate prokaryotic from eukaryotic organisms.
Shortly after Leeuwenhoek made his discoveries, the Swedish
botanist Carolus Linnaeus (1707–1778) developed a taxonomic
system—a system for naming plants and animals and group-
ing similar organisms together. For instance, Linnaeus and
other scientists of the period grouped all organisms into either
the animal kingdom or the plant kingdom. Today, biologists
still use this basic system, but they have modified Linnaeus’s
scheme by adding categories that more realistically reflect the
relationships among organisms. For example, scientists no
­longer classify yeasts, molds, and mushrooms as plants but
­instead as fungi. (We examine taxonomic schemes in more
­detail in Chapter 4.)
The microorganisms that Leeuwenhoek described can be
grouped into six basic categories: bacteria, archaea, fungi, pro-
tozoa, algae, and small multicellular animals. The only types of
microbes not described by Leeuwenhoek are viruses,3 which are
too small to be seen without an electron microscope. We briefly
consider organisms in the first five categories in the following
sections.
Bacteria and Archaea
Bacteria and archaea are prokaryotic,4 meaning that their cells
lack nuclei; that is, their genes are not surrounded by a mem-
brane. Bacterial cell walls are composed of a polysaccharide
called peptidoglycan, though some bacteria lack cell walls. The
cell walls of archaea lack peptidoglycan and instead are com-
posed of other chemicals. Members of both groups reproduce
asexually. (Chapters 3, 4, and 11 examine other differences
­b etween bacteria and archaea, and Chapters 19–21 discuss
pathogenic [disease-causing] bacteria.)
Most archaea and bacteria are much smaller than eukary-
otic cells (FIGURE 1.4). They live singly or in pairs, chains, or
clusters in almost every habitat containing sufficient moisture.
Archaea are often found in extreme environments, such as the
highly saline and arsenic-rich Mono Lake in California, acidic
3Technically, viruses are not “organisms,” because they neither replicate themselves nor
carry on the chemical reactions of living things.
4From Greek pro, meaning “before,” and karyon, meaning “kernel” (which, in this case,
refers to the nucleus of a cell).
4 CHAPTER 1 A Brief History of Microbiology
Prokaryotic Nucleus of
bacterial cells eukaryotic cheek cell
LM
20 μm
▲ FIGURE 1.4 Cells of the bacterium Streptococcus (dark blue)
and two human cheek cells. Notice the size difference.
hot springs in Yellowstone National Park, and oxygen-depleted
mud at the bottom of swamps. No archaea are known to cause
disease.
Though bacteria may have a poor reputation in our world,
the great majority do not cause disease in animals, humans,
or crops. Indeed, bacteria are beneficial to us in many ways.
For example, without beneficial bacteria, our bodies would be
much more susceptible to disease. Also, bacteria (and fungi) de-
grade dead plants and animals to release phosphorus, sulfur,
nitrogen, and carbon back into the air, soil, and water to be used
by new generations of organisms. Without microbial recyclers,
the world would be buried under the corpses of uncountable
dead organisms.
Fungi
Fungi (fŭn´jī)5 are eukaryotic;6 that is, each of their cells con-
tains a nucleus composed of genetic material surrounded by
a distinct membrane. Fungi are different from plants because
fungi obtain their food from other organisms (rather than
making it for themselves). They differ from animals by having
cell walls.
Microscopic fungi include some molds and yeasts. Molds
are typically multicellular organisms that grow as long filaments
that intertwine to make up the body of the mold. Molds repro-
duce by sexual and asexual spores, which are cells that produce
a new individual without fusing with another cell (FIGURE 1.5a).
The cottony growths on cheese, bread, and jams are molds. Pen-
icillium chrysogenum (pen-i-sil´ē-ŭm krī-so´jĕn-ŭm) is a mold
that produces penicillin.
Yeasts are unicellular and typically oval to round. They
­reproduce asexually by budding, a process in which a daughter
cell grows off the mother cell. Some yeasts also produce sex-
ual spores. An example of a useful yeast is Saccharomyces cere-
visiae (sak-ă-rō-mī´sēz se-ri-vis´ē-ī; FIGURE 1.5b), which causes
Hyphae Spores Budding cells
SEM SEM
(a) 5 μm (b) 5 μm
▲ FIGURE 1.5 Fungi. (a) The mold Penicillium chrysogenum, which
produces penicillin, has long filamentous hyphae that intertwine to form
its body. It reproduces by spores. (b) The yeast Saccharomyces cerevisiae.
Yeasts are round to oval and typically reproduce by budding.
bread to rise and produces alcohol from sugar (see Beneficial
­Microbes: Bread, Wine, and Beer on p. 7). Another example of
a yeast is Candida albicans (kan´did-ă al´bi-kanz), which causes
most cases of yeast infections in women. (Chapters 12, 22,
and 26 discuss fungi and their significance in the environment,
in food production, and as agents of human disease.)
Protozoa
Protozoa are single-celled eukaryotes that are similar to ­animals
in their nutritional needs and cellular structure. In fact, protozoa is
Greek for “first animals,” though scientists ­today ­classify them in
their own groups rather than as animals. Most protozoa are capa-
ble of locomotion, and one way scientists c­ ategorize protozoa is
according to their locomotive structures: pseudopods,7 cilia,8 or fla-
gella.9 Pseudopods are extensions of a cell that flow in the direc-
tion of travel (FIGURE 1.6a). Cilia are numerous short protrusions
of a cell that beat rhythmically to propel the protozoan through its
environment (FIGURE 1.6b). Flagella are also extensions of a cell
but are fewer, longer, and more whiplike than cilia (FIGURE 1.6c).
Some protozoa, such as the malaria-causing Plasmodium (plaz-
mō´dē-ŭm), are nonmotile in their mature forms.
Many protozoa live freely in water, but some live inside
animal hosts, where they can cause disease. Most protozoa
reproduce asexually, though some are sexual as well. ­(Chapters 12
and 23 further examine protozoa and some ­diseases they cause.)
5Plural of the Latin fungus, meaning “mushroom.”
6From Greek eu, meaning “true,” and karyon, meaning “kernel.”
7Plural Greek pseudes, meaning “false,” and podos, meaning “foot.”
8Plural of the Latin cilium, meaning “eyelid.”
9Plural of the Latin flagellum, meaning “whip.”
 CHAPTER 1 A Brief History of Microbiology 5

▶ FIGURE 1.6 Locomotive structures of protozoa. (a) Pseudo- Nucleus Pseudopods

pods are cellular extensions used for locomotion and feeding, as seen in
Amoeba proteus. (b) Blepharisma americana moves by means of cilia.
(c) Flagella are whiplike extensions that are less numerous and longer
than cilia, as seen in ­Peranema. How do cilia and flagella differ?
flagella are long and relatively few in number.
FIGURE 1.6 Cilia are short, numerous, and often cover the cell, whereas

Algae
Algae10 are unicellular or multicellular photosynthetic eukary-
otes; that is, like plants, they make their own food from carbon
dioxide and water using energy from sunlight. They differ from
plants in the relative simplicity of their reproductive structures.
Algae are categorized on the basis of their pigmentation and the
composition of their cell walls.
Large algae, commonly called seaweeds and kelps, are
LM
common in the world’s oceans. Manufacturers use gelatinous (a) 200 µm
chemicals from the cell walls of some large algae as thicken-
ers and emulsifiers in many foods and cosmetics. Scientists use
the algae-derived chemical called agar to solidify laboratory
media.
Unicellular algae (FIGURE 1.7) are common in freshwater
ponds, streams, and lakes and in the oceans as well. They are Cilia
the major food of small aquatic and marine animals and pro-
vide most of the world’s oxygen as a by-product of photosyn-
thesis. The glasslike cell walls of diatoms provide grit for many
polishing compounds. (Chapter 12 discusses other aspects of
the biology of algae.)

Other Organisms of Importance to Microbiologists

Microbiologists also study parasitic worms, which range in size
from microscopic forms (FIGURE 1.8) to adult tapeworms over
10 meters (approximately 33 feet) in length. Even though most LM
parasitic worms are not microscopic as adults, many of them (b) 10 µm

cause diseases that were studied by early microbiologists, so mi-

crobiology books and classes often discuss parasitic worms. Fur- Flagellum
ther, laboratory scientists diagnose infections of parasitic worms
by finding microscopic eggs and immature stages in blood, fe-
cal, urine, and lymph specimens. (Chapter 23 discusses parasitic
worms.)
The only type of microbe that remained hidden from Leeu-
wenhoek and other early microbiologists was the virus, which
is typically much smaller than the smallest prokaryote and is
not usually visible by light microscopy (FIGURE 1.9). Viruses
were not seen until the electron microscope was invented in
1932. All viruses are acellular (not composed of cells) obligatory
parasites composed of small amounts of genetic material (either
DNA or RNA) surrounded by a protein coat. (Chapter 13 exam-
ines the general characteristics of viruses, and Chapters 24 and
25 discuss specific viral pathogens.)
Leeuwenhoek first reported the existence of most types
LM
of microorganisms in the late 1600s, but microbiology did not (c) 20 µm

10Plural of the Latin alga, meaning “seaweed.”

6 CHAPTER 1 A Brief History of Microbiology

(a) LM (b) LM
10 μm 10 μm

▲ FIGURE 1.7 Algae. (a) Spirogyra. These microscopic algae grow as chains of cells containing
helical photosynthetic structures. (b) Diatoms. These beautiful algae have glasslike cell walls.

develop significantly as a field of study for almost two cen- one that demanded experimental evidence rather than mere
turies. There were a number of reasons for this delay. First, acceptance of traditional knowledge. This fresh philosophi-
Leeuwenhoek was a suspicious and secretive man. Though cal foundation, accompanied by improved microscopes, new
he built over 400 microscopes, he never trained an apprentice, laboratory techniques, and a drive to answer a series of piv-
and he never sold or gave away a microscope. In fact, he never otal questions, propelled microbiology to the forefront as a
let anyone—not his family or such distinguished visitors as the ­scientific discipline.
czar of Russia—so much as peek through his very best instru-
ments. When Leeuwenhoek died, the secret of creating superior
microscopes was lost. It took almost 100 years for scientists to
make microscopes of equivalent quality.
Another reason that microbiology was slow to develop as Virus
a science is that scientists in the 1700s considered microbes to
be curiosities of nature and insignificant to human affairs. But
in the late 1800s, scientists began to adopt a new philosophy,

Bacterium
Red blood cell

Viruses
assembling
inside cell

TEM
75 nm

▲ FIGURE 1.9 A colorized electron microscope image of viruses

LM
30 µm
infecting a bacterium. Viruses, which are acellular obligatory parasites,
are generally too small to be seen with a light microscope. Notice how
▲ FIGURE 1.8 An immature stage of a parasitic worm in blood. small the viruses are compared to the bacterium.
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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