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About the Authors

Michael T. Madigan received his B.S. in Biology and Education from Wisconsin State Univer-
sity–Stevens Point (1971) and his M.S. (1974) and Ph.D. (1976) in Bacteriology from the University
of Wisconsin–Madison in the laboratory of Thomas Brock. Following a postdoc at Indiana University
with Howard Gest, Mike moved to Southern Illinois University Carbondale, where he taught courses
in introductory microbiology and bacterial diversity as a professor of microbiology for 33 years. In
1988 Mike was selected as the Outstanding Teacher in the College of Science and in 1993, the Out-
standing Researcher. In 2001 he received the SIUC Outstanding Scholar Award and Distinguished
Professor title. In 2003 Mike received the Carski Award for Distinguished Undergraduate Teaching from
the American Society for Microbiology (ASM), and he is an elected Fellow of the American Academy
of Microbiology (ASM) and the American Association for the Advancement of Science (AAAS). He has
also been recognized by the American Red Cross as a major volunteer blood donor for the 24 gallons
of blood he has donated since 1967. Mike’s research is focused on phototrophic bacteria that inhabit
extreme environments, and for the past 20 years his emphasis has been Antarctic microbiology. Mike
has co-edited a major treatise on phototrophic bacteria and served for 10 years as chief editor of the
journal Archives of Microbiology. He currently serves on the editorial board of the journals Environmen-
tal Microbiology and Antonie van Leeuwenhoek. Mike’s other interests include forestry, swimming, reading,
and caring for his dogs and horses. He lives on a small farm near a quiet lake with his wife, Nancy,
three dogs (Kato, Nut, and Merlyn), and three horses (Eddie, Georgie, and Roscoe).

Kelly S. Bender received her B.S. in Biology from Southeast Missouri State University (1999)
and her Ph.D. (2003) in Molecular Biology, Microbiology, and Biochemistry from Southern Illinois
University Carbondale. Her dissertation research focused on the genetics of perchlorate-reducing bac-
teria. During her postdoctoral fellowship, Kelly worked on the genetic regulation of sulfate-reducing
bacteria in the laboratory of Judy Wall at the University of Missouri–Columbia. She also completed a
transatlantic biotechnology fellowship at Uppsala University in Sweden researching regulatory small
RNAs in bacteria. In 2006, Kelly returned to her alma mater, Southern Illinois University Carbondale,
as an Assistant Professor in the Department of Microbiology and in 2012 was tenured and promoted
to Associate Professor. She has served as Chair of the SIUC Department of Microbiology since 2018.
Her lab studies a range of topics including regulation in sulfate-reducing bacteria, the microbial com-
munity dynamics of sites impacted by acid mine drainage, and diversity of phototrophic heliobacteria.
Kelly teaches courses in introductory microbiology and microbial diversity, has served on numerous
federal grant review panels, and is an active member of the American Society for Microbiology (ASM).
Her other interests include spending time with her daughter, Violet, and husband, Dick.

Daniel H. Buckley is a Professor at Cornell University in the School of Integrative Plant Science
and the Department of Microbiology. He earned his B.S. in Microbiology (1994) at the University of
Rochester and his Ph.D. in Microbiology (2000) at Michigan State University. His graduate research in
the laboratory of Thomas M. Schmidt explored environmental factors that influence microbial diversity
in soils. Dan then received a National Science Foundation Postdoctoral Fellowship to work with Pieter
T. Visscher, University of Connecticut, investigating linkages between microbial diversity and biogeo-
chemistry within microbial mats and stromatolites. Dan moved to Cornell in 2003 where he investi-
gates the ecology and evolution of the diverse microorganisms that live in soils. He has taught both
introductory and advanced courses in microbiology, microbial diversity, and microbial genomics. He
received a National Science Foundation Faculty Early Career Development (CAREER) award in 2005
for excellence in integrating research and education, and served as Co-Director of the MBL Microbial
Diversity summer course in Woods Hole, Massachusetts (2009–2013). He currently serves on the edi-
torial boards of Applied and Environmental Microbiology and Environmental Microbiology. Dan lives in
Ithaca, New York, with his wife, Merry, and sons, Finn and Colin.

iii
x P R E FA C E
Chapter 18
• Along with major updates on eukaryotic phylogeny, a new section
is devoted to the haptophytes, including the globally and ecologi-
cally important coccolithophore Emiliania huxleyi. Coccolitho-
phores play a major role in regulating global climate, illustrating
the power that microbes exert over our biosphere.
UNIT 5 Microbial Ecology and
Environmental Microbiology
Chapter 19
• The chapter begins a unit on ecology and environmental microbi-
ology. The modern tools of the microbial ecologist are described
with examples of how each has helped sculpt the science.
• Some highlights: A new method to visualize protein synthesis in
single cells allows study of microbial activity in the environment.
Metabolomics exploits new methods in mass spectrometry to
unravel the complex metabolic interactions sustaining microbial
communities. Nanosensor technologies are revealing how
microbes alter the chemical landscape of three-dimensional sur-
faces. A new section explores multi-omics, which combines mul-
tiple state-of-the-art analytical tools to more fully characterize
microbial communities.
Chapter 20
• The properties and microbial diversity of major microbial ecosys-
tems including soils and aquatic systems are compared and con-
trasted in exciting ways.
• Some highlights: Expansive coverage of surface-attached micro-
bial communities and how those communities are responding
to plastic pollution of the environment. New understanding of
the ecology of iron-oxidizing bacteria revealed by the isolation
of new members of this biogeochemically significant group. The
discovery in deep ocean sediments of novel Archaea that link this
domain with Eukarya. Extensive coverage of marine viruses, their
abundance and diversity, and how they alter the physiology of
organisms they infect. Humans traveling to 10,000-meter depths
in the oceans discover the most pressure-tolerant bacterium
known.
Chapter 21
• Extensive coverage of the major nutrient cycles in nature and the
microbes that catalyze them are presented in a fashion that allows
the cycles to be taught as individual entities or as interrelated met-
abolic loops.
• Some highlights: Expanded coverage of the biogeochemistry of
sulfur compounds highlights the importance of volatile microbial
products such as dimethyl sulfide for cloud formation. Advances
in the biochemistry of extracellular electron transfer add new
understanding to how the ecology and diversity of microorganisms
drive the biogeochemical cycling of iron and manganese. The mys-
tery of how methane is generated (typically a strictly anoxic pro-
cess) in highly oxygenated ocean surface waters is solved by
discoveries in the phosphorus cycle described in a new Explore the
Microbial World.
Chapter 22
• This chapter on the built environment shows how humans create
new microbial habitats through construction of buildings, sup-
porting infrastructure, and habitat modification, and which
microbes take advantage of these habitats and why.
• Some highlights: The microbial metabolism of biologically pro-
duced and manufactured chlorinated organics has been expanded,
as has the basis for the bioremediation of major chemical pollut-
ants. How microbes are responding to the mountains of plastics
contaminating the environment and the discovery of novel bacte-
ria capable of degrading plastic bottles are described. New technol-
ogy that improves the efficiency of wastewater treatment using
granular sludge technology is presented, and the microbial
response to the excessive use of common household cleansers is
considered.
Chapter 23
• A chapter devoted to nonhuman microbial symbioses describes the
major microbial partners that live in symbiotic associations with
other microbes, with plants, and with animals other than humans.
• Some highlights: Newly revised section on symbioses between
microorganisms addresses the ecological significance of phototroph
switching in lichens and how certain bacterial species use electri-
cally conductive structures to form intimate symbiotic associations.
Several updates include how insect symbionts are used to combat
transmission of major viral diseases of humans and how defensive
chemicals produced by symbionts protect insects from predation.
Detailed coverage is given to the elaborate “cross-talk” between
microbe and animal needed to establish the squid light organ.
UNIT 6 Microbe–Human Interactions
and the Immune System
Chapter 24
• A chapter on the human microbiome launches the unit on
microbe–human interactions and the immune system by introduc-
ing and updating advances in our understanding of the microbes
that inhabit the human body and their relationship to health and
disease.
• Some highlights: The discovery of ultrasmall bacteria in the mouth
parasitizing other bacteria brings a new twist to the microbial ecol-
ogy of the oral cavity. A new section on the human virome describes
how metagenomics is driving the discovery and isolation of inter-
esting new viruses. Extensive coverage is devoted to the impact of
early-life events on the development of the newborn gut microbi-
ome and of recent successes in probiotic therapy for preventing
newborn intestinal diseases.
Chapter 25
• Beginning with this chapter, the book shifts its focus to pathogenic
microorganisms, the immune system, and disease. Part I of this
chapter addresses microbial adherence, colonization and invasion,
and pathogenicity, including important sections on virulence and
virulence attenuation. Part II highlights key enzymes and toxins
produced by microbes that contribute to pathogenesis.

• Some highlights: The updated text includes expanded coverage of
bacterial adhesins supported by a new, two-part figure that high-
lights new discoveries in staphylococcal adherence. Revised cover-
age of virulence attenuation includes new artwork to show how
this principle can be exploited for development of effective vac-
cines. An updated discussion of botulinum toxins reflects new find-
ings and clearly presents both the neurotoxic mechanism and the
surprising clinical utility of these extremely potent substances.
Chapter 26
• Chapter 26 opens with an overview of the immune system and
the body’s first-line barriers to infection. This is followed by a
brief discussion of hematopoiesis before focusing on innate
immune responses to pathogen invasion. The chapter provides a
natural progression into adaptive immune responses covered in
Chapter 27.
• Some highlights: In addition to a new chapter opener highlighting
breakthroughs that link Alzheimer’s disease to microbial infection,
this chapter contains heavily edited text that includes a more com-
prehensive discussion of leukocyte diversity and an all-new
description of the role of amyloid-β protein as an innate defense
in the brain. Other highlights include expanded coverage of inter-
ferons and the role of natural killer cells as the primary effectors of
antibody-dependent cell-mediated cytotoxicity. Finally, a fascinat-
ing new Explore the Microbial World highlights the role of pattern
recognition receptors in establishing host–microbe mutualisms
using hydrothermal vent tube worms as an example.
Chapter 27
• Chapter 27 begins with an essential discussion of the principles
that define adaptive immunity: specificity, immune memory, lym-
phocyte selection, and immune tolerance. This is followed by sec-
tions that discuss the functional mechanisms of the key cells and
proteins (immunoglobulins, major histocompatibility complexes,
and T cell receptors) that drive adaptive immunity.
• Some highlights: The text has been heavily edited throughout, and
this has produced a clearer and more informative presentation of
B and T lymphocyte selection and tolerance, including a new dis-
cussion of T-dependent versus T-independent antigens. In addi-
tion, a new section dedicated to T cell activation and anergy clearly
presents the important concept of the second signal required for
T cell activation.
Chapter 28
• The newly reorganized Chapters 28 and 29 have emerged from
materials presented in Chapter 28 of the 15th edition. Treating
immune disorders and antimicrobial therapy (Chapter 28) sepa-
rately from clinical diagnostic methods (Chapter 29) has produced
a more teachable format, making these topics more accessible for
students and easier for the instructor to plan course assignments.
• Some highlights: The text progresses smoothly from immune dis-
orders and deficiencies to methods used to train and hone the
immune response for disease prevention and treatment. New cover-
age of mRNA and plant-based vaccines shares the latest innovations
in vaccinology. An all-new section on immunotherapy, supported
by vibrant new artwork, highlights exciting advancements in the use
of genetic engineering and molecular immunology to treat cancer.
P R E FA C E xi
UNIT 7 Infectious Diseases
Chapter 29
• To bring better focus to the material, this chapter is now solely
dedicated to the clinical microbiology laboratory and includes
information on lab safety, healthcare-associated infections, and a
wide array of both culture-dependent and culture-independent
techniques used to diagnose infectious diseases.
• Some highlights: The chapter launches with the description of an
exciting new method of diagnosing tuberculosis—humanity’s most
notorious scourge. The text has been edited throughout for better
organization and clarity, and art modifications help clarify com-
plex diagnostic techniques. Updated terminology includes an
introduction to point-of-care diagnostics.
Chapter 30
• This chapter introduces the topics and terminology of the science
of epidemiology and public health. Historical and modern exam-
ples throughout emphasize key concepts such as emerging (and
reemerging) diseases, epidemics and pandemics, and the public
health threat associated with the development and use of weapon-
ized microorganisms.
• Some highlights: incorporation of the most up-to-date statistics
available on disease incidence and outbreaks throughout the text
and in figures and tables, as well as an all-new section supported
by photos on the emergence of the important healthcare-associated
pathogen Clostridioides (Clostridium) difficile.
Chapter 31
• This is the first of four highly visual chapters that take an ecological
approach to pathogenic microorganisms by considering infectious
diseases based on their modes of transmission. Bacterial and viral
diseases transmitted person to person by way of airborne particles,
direct contact, or sexual contact are the focus here.
• Some highlights: Statistical data regarding key emerging and
reemerging diseases, including measles, pertussis, influenza, hepa-
titis, HIV/AIDS, gonorrhea, and syphilis have been updated to
reflect the most recent data available; an all-new discussion with
supporting photo of the neglected tropical disease yaws helps
impart knowledge and awareness of this lingering scourge.
Chapter 32
• In this chapter we examine pathogens transmitted to humans
through either an animal vector or soil-contaminated wounds or
objects. Many of these diseases have high morbidity and mortality
rates, and in most cases, effective vaccines are not yet available.
• Some highlights: The text and figures include the most up-to-
date statistics for diseases throughout the chapter, including
rabies, hantavirus, spotted fever rickettsiosis, ehrlichiosis and
anaplasmosis, Lyme disease, and the major tropical hemorrhagic
fevers. In addition, the text now includes updated discussions of
the emergence of key tickborne diseases in the United States and
coverage of new strategies against dengue fever, including
description of a new vaccine and the use of the bacterial endo-
symbiont Wolbachia to control the dengue virus–infected mos-
quito population.
xii P R E FA C E
Chapter 33
• Pathogens in contaminated water or food are easily transmitted to
humans, with waterborne diseases being especially common in
developing countries lacking adequate water treatment facilities.
This chapter highlights the most prevalent water- and foodborne
diseases and emphasizes the importance of clean water and proper
food preparation and preservation in preventing these physically
uncomfortable and occasionally fatal illnesses.
• Some highlights: Updated statistics have been incorporated for all
major water- and foodborne diseases, including Campylobacter
infections, which have now overtaken salmonellosis as the leading
cause of bacterial food infection in the United States. New discus-
sions cover recently elucidated norovirus pathology and new food
safety developments, including the use of eBeam technology and
bacteriophage sprays. A new overview figure of cholera infection
integrates photos with artwork to emphasize key aspects of this
devastating and all too common disease.
Chapter 34
• Eukaryotic pathogens present a special challenge to medicine
because, on a cellular level, they are not that different from our
own cells. Thus, it can be difficult to find selective targets for che-
motherapeutic drugs. Yet the microbes highlighted in this highly
visual chapter cause some of the most devastating and prevalent
diseases today.
• Some highlights: New color photos adorn the chapter, including
two stunning fluorescent micrographs of Entamoeba histolytica, the
causative agent of amebic dysentery. Broader coverage of distinc-
tive features of several diseases, including cyclosporiasis, toxoplas-
mosis, and malaria, has been seamlessly incorporated. All statistics
have been updated with the most recent surveillance data to yield
a global picture of fungal and parasitic diseases.
Acknowledgments
A n excellent textbook is an educational resource that can only
emerge from the combined contributions of a dedicated book
team. In addition to the authors, the Brock Biology of Microorganisms
(BBOM) team was composed of folks both inside and outside of Pear-
son. Content Manager Josh Frost paved the way for the 16th Edition of
BBOM and provided the resources necessary for the authors to produce
a spectacular revision in a timely fashion. Importantly, Josh also brought
to BBOM his extensive experience as Content Manager of Campbell Biol-
ogy, the leading textbook of biology worldwide. This greatly benefited
BBOM, a book whose educational philosophy has traditionally paral-
leled that of strong majors-level biology books. The BBOM coauthor
team greatly appreciated the guidance and input that Josh brought to
our book.
BBOM 16e editorial and production were headed up by Michele Man-
gelli (Mangelli Productions), and both cover and interior designs were
created by Gary Hespenheide (Hespenheide Design). Michele assembled
and managed the production team and kept editorial and production
on mission, on budget, and on time, and did so tirelessly in a helpful,
author-friendly, and accommodating manner. The artistic magic of Gary
is clearly visible in the outstanding internal design that smoothly ushers
the reader through the book with highly effective organizational and
navigational cues. Gary also designed the book’s cover—a spectacular
display of microbial diversity (photo courtesy of Professor Dr. Jörg
Overmann, Braunschweig, Germany). The art team at Imagineering Art
(Toronto, Canada) did an outstanding job in helping the authors link
art with text and provided many helpful suggestions and options for art
presentation, consistency, and style. Many thanks are extended to
Michele, Gary, Jörg, and Imagineering for their outstanding efforts on
BBOM 16e.
Many other people were part of the book production, editorial, or
marketing team, including Karen Gulliver, Jean Lake, Kristin Piljay, Betsy
Dietrich, Maureen Johnson, Susan Wang, Greg Anderson, and Elizabeth
McPherson. Karen was our excellent and highly efficient production
editor; she kept manuscript and pages moving smoothly through the
wheels of production, graciously tolerated the authors’ many requests,
and accommodated our time constraints. Jean was our art coordinator,
efficiently tracking and routing art and handling interactions between
the art studio and the authors to ensure quality control and a timely
schedule. Betsy worked with Jean and Karen to ensure an art program
and text free of both bloopers and subtle errors. Kristin was our photo
researcher who dug out some of the hard-to-find specialty photos that
grace BBOM 16e. Susan, Greg, and Elizabeth composed an excellent
accuracy review team and made numerous very helpful comments. The
authors thank Karen, Jean, Kristin, Betsy, Susan, Greg, and Elizabeth for
their combined contributions to the book you see in front of you today.
Special thanks go to Anita Wagner Hueftle, our spectacular copyeditor
and a key part of the book team. Anita is not only a master wordsmith;
her amazing gift of being able to keep track of where everything was said
in this book and how everything was said in this book has helped us
make the most readable, accurate, and consistent textbook of microbiol-
ogy available today. Thank you kindly, Anita; you made outstanding
contributions to this edition.
We thank Alysun Estes and Krista Clark for their much-appreciated
marketing support.
We are also grateful to the top-notch educators who constructed the
Mastering Microbiology program that accompanies this text; these
include: Candice Damiani, Jennifer Hatchel, Bryan Jennings, Ann Pater-
son, Emily Booms, Ines Rauschenbach, and Monica Togna.
And last but not least, no textbook in microbiology could be pub-
lished without reviewing of the manuscript and the gift of new photos
from experts in the field. We are therefore extremely grateful for the
assistance of the many individuals who kindly provided manuscript
reviews, unpublished results, and new photos. Complete photo credits
in this book are found either alongside a photo or in the photo credits
listed in the back of the book. Reviewers and photo suppliers included:
Jônatas Abrahão, Universidade Federal de Minas Gerais (Brazil)
Sonja-Verena Albers, University of Freiburg (Germany)
Rebecca Albright, California Academy of Sciences
Douglas Bartlett, Scripps Institution of Oceanography
Bonnie Bassler, Princeton University
David Battigelli, University of North Carolina Greensboro
Tom Ian Battin, Ecole Polytechnique Federale de Lausanne
(Switzerland)
J. Thomas Beatty, University of British Columbia (Canada)
Veysel Berk, Wallit! Inc.
Edward Bernard, University of Maine
Tanmay Bharat, University of Oxford (England)
Benjie Blair, Jacksonville State University
Ilka Bischofs, Max Planck Institute for Terrestrial Biology (Germany)
Robert Blankenship, Washington University in St. Louis
Antje Boetius, Max Planck Institute for Marine Microbiology
(Germany)
Gary Borisy, The Forsyth Institute
Ariane Briegel, Leiden University (Netherlands)
Volker Brinkmann, Max Planck Institute for Infection Biology, Berlin
Pamela Brown, University of Missouri
Jennifer Brum, Louisiana State University
Marie Bulínová, Charles University (Czech Republic)
Gustavo Caetano-Anollés, University of Illinois
Christian Cambillau, Centre National de la Recherche Scientifique
Aix-Marseille University Marseille (France)
Hans Carlson, Lawrence Berkeley National Laboratory
Dale Casamatta, University of North Florida
Clara S. Chan, University of Delaware
Matthew Wook Chang, National University of Singapore
Beat Christen, ETH Zürich (Switzerland)
Pascale Cossart, Pasteur Institute (France)
Matt Cruzen, Biola University
Michele Culumber, Weber State University
xiii
xiv ACKNOWLEDGMENTS
Ankur Dalia, Indiana University
Bertram Daum, University of Exeter (England)
Svetlana N. Dedysh, Winogradsky Institute of Microbiology (Russia)
David J. Des Marais, NASA Ames
Adam Deutschbauer, Lawrence Berkeley National Laboratory
Omar Din, University of California, San Diego
Alice Dohnalkova, Pacific Northwest National Laboratory
(Richland, WA)
Steven Dominy, Cortexyme, Inc.
Paul Dunlap, University of Michigan
David Emerson, Bigelow Laboratory for Ocean Sciences (Maine)
Susanne Erdmann, University of New South Wales (Australia)
Belinda Ferrari, University of New South Wales (Australia)
Derek J. Fisher, Southern Illinois University
Jason Flowers, Murraysmith (Seattle, WA)
Bruce Fouke, University of Illinois Urbana–Champaign
Andreas Giesen, Royal HaskoningDHV (The Netherlands)
Mariana Gomes de Pinho, Universidade Nova de Lisboa (Portugal)
Hans-Peter Grossart, Leibniz-Institute of Freshwater Ecology and Inland
Fisheries (Germany)
Ricardo Guerrero, University of Barcelona (Spain)
Jennifer Hatchel, College of Coastal Georgia
Roland Hatzenpichler, Montana State University
Jennifer Hess, Aquinas College
Donald Hilvert, ETH Zürich (Switzerland)
Jay Hodgson, Georgia Southern University
William Inskeep, Montana State University
Anthony James, University of California, Irvine
Joshua Jenkins, University of Bristol Dental School (England)
Grant Jensen, California Institute of Technology
Deborah O. Jung, Southeast Missouri State University
Kathryn Kauffman, Massachusetts Institute of Technology
Vjollca Konjufca, Southern Illinois University
Klaus Koren, University of Copenhagen (Denmark)
Michael Kovach, Baldwin Wallace University
Michael Kühl, University of Copenhagen (Denmark)
Philippe Laissue, University of Essex (England)
Christian Lesterlin, Institut de Biologie et Chimie des
Protéines (France)
Ruth E. Ley, Max-Planck Institute for Developmental Biology (Germany)
Shee-Mei Lok, Duke-NUS Medical School (Singapore)
E. Erin Mack, Dupont Corporate Remediation Group (Newark, DE)
Jessica Mark Welch, Marine Biological Laboratory, Woods Hole
Francis Martin, Lab of Excellence ARBRE, INRA-Nancy (France)
Sean McAllister, University of Delaware
Margaret McFall-Ngai, University of Hawaii at Manoa
Jeffrey McLean, University of Washington, Seattle
Nancy Moran, University of Texas
Phillip Nadeau, Massachusetts Institute of Technology
National Aeronautics and Space Administration (USA)
Jeniel Nett, University of Wisconsin
Daniela Nicastro, University of Texas Southwestern
Trent Northen, Lawrence Berkeley National Laboratory
Gal Ofir, Weizmann Institute of Science (Israel)
Catherine Oikonomou, California Institute of Technology
Jörg Overmann, Leibniz Institute DSMZ Braunschweig (Germany)
Niki Parenteau, NASA Ames
Donghyun Park, Yale University School of Medicine
Nicolás Pinel, Universidad de Antioquia (Colombia)
Marie Pizzorno, Bucknell University
Joe Pogliano, University of California, San Diego
Martin Polz, Massachusetts Institute of Technology
Tessa Quax, University of Freiburg (Germany)
Katherine Ralston, University of California
Ines Rauschenbach, Rutgers University
Tara Renbarger, Indiana Wesleyan University
Niels Peter Revsbech, Aarhus University (Denmark)
Ned Ruby, University of Hawaii at Manoa
Bernhard Schink, University of Konstanz (Germany)
Susan Schlimpert, John Innes Centre (England)
Andrey Shkoporov, University College Cork (Ireland)
Simon Silver, University of Illinois Chicago
Janice Speshock, Tarleton State University
Emily Stowe, Bucknell University
Shunichi Takahashi, National Institute for Basic Biology (Japan)
Francisco Tenllado, Centro de Investigaciones Biológicas (Spain)
Andreas Teske, University of North Carolina
David Valentine, University of California, Santa Barbara
Mikel Valle, Asociación Centro de Investigación Cooperativa en
Biociencias (Spain)
Nicholas Verola, Verola Studio (Vero Beach, FL)
Marilyn B. Vogel, Auburn University
Judy Wall, University of Missouri
Dave Ward, Montana State University
Bishop Wash, The Richards Group (Dallas TX)
Jillian L. Waters, Max Planck Institute for Developmental Biology
(Germany)
Nicki Watson, Massachusetts Institute of Technology
Gunter Wegener, Max Planck Institute of Marine Microbiology
(Germany)
Mari Winkler, University of Washington
Gerry Wright, McMaster University
Vamsi Yadavalli, Virginia Commonwealth University
Jeremy R. Young, University College London (England)
Carl Zeiss AG (Jena, Germany)
Jizhong (Joe) Zhou, University of Oklahoma
Steve Zinder, Cornell University
As hard as a publishing team may try, no textbook can ever be com-
pletely error-free. Although we are confident the reader will be hard
pressed to find errors in BBOM 16e, any errors that do exist, either of
commission or omission, are the responsibility of the authors. In past
editions, users have been kind enough to contact us when they spot an
error so we can fix it in a subsequent printing. Users should feel free to
continue to contact the authors directly about any errors, concerns, ques-
tions, or suggestions they have about the book. We are always happy to
hear from our readers; through the years, your comments have helped
make the book stronger.
Michael T. Madigan (madigan@siu.edu)
Kelly S. Bender (bender@siu.edu)
Daniel H. Buckley (dbuckley@cornell.edu)
W. Matthew Sattley (matthew.sattley@indwes.edu)
David A. Stahl (dastahl@uw.edu)
Contents

2
About the Authors iii
Preface vii Microbial Cell Structure
Acknowledgments xiii and Function 38
UNIT 1 The Foundations MICROBIOLOGYNOW Exploring the Microbial Cell 38
of Microbiology I  
• The Cell Envelope 39
2.1 The Cytoplasmic Membrane 39

1 The Microbial World 1 2.2

2.3
2.4
2.5
Transporting Nutrients into the Cell 42
The Cell Wall 44
LPS: The Outer Membrane 47
Diversity of Cell Envelope Structure 49
MICROBIOLOGYNOW Microbiology in Motion 1
II  • Cell Surface Structures and
I  
• Exploring the Microbial World 2 Inclusions 51
1.1 Microorganisms, Tiny Titans 2.6 Cell Surface Structures 51
of the Earth 2 2.7 Cell Inclusions 53
1.2 Structure and Activities of 2.8 Endospores 55
Microbial Cells 3
1.3 Cell Size and Morphology 5 III   • Cell Locomotion 58
1.4 An Introduction to Microbial Life 10 2.9 Flagella, Archaella, and Swimming Motility 58
1.5 Microorganisms and the Biosphere 12 2.10 Surface Motility 61
1.6 The Impact of Microorganisms on 2.11 Chemotaxis 63
Human Society 13 2.12 Other Forms of Taxis 65

II  • Microscopy and the Origins IV    • Eukaryotic Microbial Cells 66

of Microbiology 18 2.13 The Nucleus and Cell Division 66
1.7 Light Microscopy and the Discovery 2.14 Mitochondria and Chloroplasts 68
of Microorganisms 18 2.15 Other Eukaryotic Cell Structures 70
1.8 Improving Contrast in Light
Microscopy 20

3
1.9 Imaging Cells in Three Dimensions 22
1.10 Probing Cell Structure: Electron Microscopy 23 Microbial Metabolism 75
III   • Microbial Cultivation Expands the Horizon
of Microbiology 25
MICROBIOLOGYNOW Life Begins with Metabolism 75
1.11 Pasteur and Spontaneous
Generation 25 I  
• Fundamentals of Metabolism 76
1.12 Koch, Infectious Diseases,
3.1 Defining the Requirements for Life 76
and Pure Cultures 27
3.2 Electron Transfer Reactions 78
1.13 Discovery of Microbial Diversity 29
3.3 Calculating Changes in Free Energy 80
IV    • Molecular Biology and the Unity 3.4 Cellular Energy Conservation 82
and Diversity of Life 31 3.5 Catalysis and Enzymes 84
1.14 Molecular Basis of Life 31 II  • Catabolism: Chemoorganotrophs 85
1.15 Woese and the Tree of Life 32
3.6 Glycolysis, the Citric Acid Cycle,
Explore the Microbial World and the Glyoxylate Cycle 86
Tiny Cells 9 3.7 Principles of Fermentation 88

xv
xvi CONTENTS

5
3.8 Principles of Respiration: Electron Carriers 89
3.9 Principles of Respiration: Generating a Proton Viruses and Their
Motive Force 91 Multiplication 148
III   • Catabolism: Electron Transport
and Metabolic Diversity 94 MICROBIOLOGYNOW  hen Antibiotics Fail, Bacteriophage
W
3.10 Anaerobic Respiration and Metabolic Modularity 94 Therapy to the Rescue 148
3.11 Chemolithotrophy and Phototrophy 96
I  
• The Nature of Viruses 149
IV    • Biosynthesis 98 5.1 What Is a Virus? 149
3.12 Autotrophy and Nitrogen Fixation 98 5.2 Structure of the Virion 151
3.13 Sugars and Polysaccharides 101 5.3 Culturing, Detecting, and Counting Viruses 153
3.14 Amino Acids and Nucleotides 102
3.15 Fatty Acids and Lipids 103 II  • Overview of the Viral Replication Cycle 155
5.4 Steps in the Replication Cycle 155
5.5 Bacteriophage T4: A Model Lytic Virus 156

4
5.6 Temperate Bacteriophages and Lysogeny 159
Microbial Growth 5.7 An Overview of Viruses of Eukaryotes 159
and Its Control 108

MICROBIOLOGYNOW Growing Their Own Way 108 UNIT 2 Molecular Biology

I  
• Culturing Microbes and Measuring Their and Genetics
Growth 109
4.1
4.2
4.3
4.4
Feeding the Microbe: Cell Nutrition 109
Growth Media and Laboratory Culture 111
Microscopic Counts of Microbial Cell Numbers 114
Viable Counting of Microbial Cell Numbers 115
6 Molecular Information Flow
and Protein Processing 165

4.5 Turbidimetric Measures of Microbial Cell MICROBIOLOGYNOW Injectisomes: Salmonella’s Mode of Attack 165
Numbers 117
I  
• Molecular Biology and Genetic Elements 166
II  • Dynamics of Microbial Growth 118 6.1 DNA and Genetic Information Flow 166
4.6 Binary Fission and the Microbial Growth Cycle 118 6.2 Genetic Elements: Chromosomes
4.7 Quantitative Aspects of Microbial Growth 120 and Plasmids 169
4.8 Continuous Culture 122
4.9 Biofilm Growth 123 II  • Copying the Genetic Blueprint: DNA
4.10 Alternatives to Binary Fission 124 Replication 172
6.3 Templates, Enzymes, and the Replication Fork 172
III   • Environmental Effects on Growth: 6.4 Bidirectional Replication, the Replisome, and
Temperature 126 Proofreading 175
4.11 Temperature Classes of Microorganisms 126
4.12 Microbial Life in the Cold 127 III   • RNA Synthesis: Transcription 177
4.13 Microbial Life at High Temperatures 129 6.5 Transcription in Bacteria 177
6.6 Transcription in Archaea and Eukarya 181
IV    • Environmental Effects on Growth: pH, Osmolarity,
and Oxygen 131 IV    • Protein Synthesis: Translation 183
4.14 Effects of pH on Microbial Growth 132 6.7 Amino Acids, Polypeptides, and Proteins 183
4.15 Osmolarity and Microbial Growth 133 6.8 Transfer RNA 186
4.16 Oxygen and Microbial Growth 135 6.9 Translation and the Genetic Code 187
6.10 The Mechanism of Protein Synthesis 189
V  • Controlling Microbial Growth 137
4.17 General Principles and Microbial Growth Control by V  • Protein Processing, Secretion,
Heat 138 and Targeting 192
4.18 Other Physical Control Methods: Radiation and 6.11 Assisted Protein Folding and Chaperones 192
Filtration 139 6.12 Protein Secretion: The Sec and Tat Systems 193
4.19 Chemical Control of Microbial Growth 141 6.13 Protein Secretion: Gram-Negative Systems 194
 CONTENTS xvii

III   • Antibiotics and Microbial Growth 255

7 Microbial Regulatory
Systems 200
8.11
8.12
Antibiotic Targets and Antibiotic Resistance 255
Persistence and Dormancy 257

9
MICROBIOLOGYNOW  s Bacterial Cells Chatter,
A
Viruses Eavesdrop 200 Genetics of Bacteria
I  
• DNA-Binding Proteins and Transcriptional
and Archaea 261
Regulation 201
7.1 DNA-Binding Proteins 201 MICROBIOLOGYNOW L ive Cell Imaging Captures Bacterial
7.2 Transcription Factors and Effectors 202 Promiscuity 261
7.3 Repression and Activation 204 I  
• Mutation 263
7.4 Transcription Controls in Archaea 207
9.1 Mutations and Mutants 263
II  • Sensing and Signal Transduction 209 9.2 Molecular Basis of Mutation 265
7.5 Two-Component Regulatory Systems 209 9.3 Reversions and Mutation Rates 267
7.6 Regulation of Chemotaxis 210 9.4 Mutagenesis 268
7.7 Cell-to-Cell Signaling 213 II  • Gene Transfer in Bacteria 270
III   • Global Control 215 9.5 Genetic Recombination 271
7.8 The lac Operon 216 9.6 Transformation 273
7.9 Stringent and General Stress Responses 218 9.7 Transduction 275
7.10 The Phosphate (Pho) Regulon 220 9.8 Conjugation 278
7.11 The Heat Shock Response 221 9.9 The Formation of Hfr Strains and Chromosome
Mobilization 279
IV    • RNA-Based Regulation 222
III   • Gene Transfer in Archaea and Other
7.12 Regulatory RNAs 223
7.13 Riboswitches 224
Genetic Events 282
7.14 Attenuation 226 9.10 Horizontal Gene Transfer in Archaea 282
9.11 Mobile DNA: Transposable Elements 284
V  • Regulation of Enzymes and Other Proteins 227 9.12 Preserving Genomic Integrity and CRISPR 286
7.15 Feedback Inhibition 228

UNIT 3 Genomics, Synthetic Biology,

7.16 Post-Translational Regulation 228

and Evolution
8 Molecular Aspects of Microbial
Growth 234

MICROBIOLOGYNOW  embrane Vesicles: Nano Vehicles

M
Transporting Important Cargo 234
10 Microbial Genomics and Other
Omics 292

MICROBIOLOGYNOW  mics Tools Unravel Mysteries

O
I  
• Bacterial Cell Division 235 of “Fettuccine” Rocks 292
8.1 Visualizing Molecular Growth 235
8.2 Chromosome Replication and Segregation 236 I  
• Genomics 293
8.3 Cell Division and Fts Proteins 239 10.1 Introduction to Genomics 293
8.4 Determinants of Cell Morphology 241 10.2 Sequencing and Annotating Genomes 295
8.5 Peptidoglycan Biosynthesis 243 10.3 Genome Size and Gene Content in Bacteria and
Archaea 298
II  • Regulation of Development 10.4 Organelle and Eukaryotic Microbial Genomes 302
in Model Bacteria 246
8.6 Regulation of Endospore Formation 246 II  • Functional Omics 305
8.7 Regulation of Endospore Germination 247 10.5 Functional Genomics 305
8.8 Caulobacter Differentiation 248 10.6 High-Throughput Functional Gene
8.9 Heterocyst Formation in Anabaena 250 Analysis: Tn-Seq 308
8.10 Biofilm Formation 251
xviii CONTENTS

10.7 Metagenomics 308 12.3 Expressing Foreign Genes

10.8 Gene Chips and Transcriptomics 311 in Bacteria 362
10.9 Proteomics and the Interactome 314 12.4 Molecular Methods for
10.10 Metabolomics 316 Mutagenesis 364
12.5 Reporter Genes and Gene Fusions 365
III   • Systems Biology 317
10.11 Single-Cell Genomics 318 II  • Making Products from Genetically Engineered
10.12 Integrating Mycobacterium tuberculosis Omics 319 Microbes: Biotechnology 367
10.13 Systems Biology and Human Health 321 12.6 Somatotropin and Other
Mammalian Proteins 367
Explore the Microbial World 12.7 Transgenic Organisms in Agriculture and
DNA Sequencing in the Palm of Your Hand 300 Aquaculture 369
12.8 Engineered Vaccines and
Therapeutic Agents 371

11 Viral Genomics
and Diversity 325
12.9 Mining Genomes and
Engineering Pathways 375
12.10 Engineering Biofuels 377

MICROBIOLOGYNOW  acteriophages Mimicking Eukaryotes—

B
III   • Synthetic Biology and Genome Editing 379
Discovery of a Phage-Encoded Nucleus 12.11 Synthetic Metabolic Pathways, Biosensors,
and Spindle 325 and Genetic Circuits 380
12.12 Synthetic Cells 383
I  
• Viral Genomes and Classification 326 12.13 Genome Editing and CRISPRs 384
11.1 Size and Structure of Viral Genomes 326 12.14 Biocontainment of Genetically Modified
11.2 Viral Taxonomy and Phylogeny 328 Organisms 388

II  • DNA Viruses 330

13 Microbial Evolution
11.3 Single-Stranded DNA Bacteriophages: fX174 and
M13 330
11.4 Double-Stranded DNA Bacteriophages: T4, T7, and and Genome Dynamics 392
Lambda 332
11.5 Viruses of Archaea 335
MICROBIOLOGYNOW Exploring Viral Genesis 392
11.6 Uniquely Replicating DNA Animal Viruses 338
11.7 DNA Tumor Viruses 339 I  
• Early Earth and the Origin
and Diversification of Life 393
III   • RNA Viruses 341
13.1 Formation and Early History of Earth 393
11.8 Positive-Strand RNA Viruses 341
13.2 Photosynthesis and the Oxidation
11.9 Negative-Strand RNA Animal Viruses 343
of Earth 396
11.10 Double-Stranded RNA Viruses 345
13.3 Living Fossils: DNA Records the History
11.11 Viruses That Use Reverse Transcriptase 346 of Life 398
IV    • Subviral Agents 349 13.4 Endosymbiotic Origin of Eukaryotes 399
13.5 Viral Evolution 402
11.12 Viroids 349
11.13 Prions 350 II  • Mechanisms of Microbial
Evolution 403
13.6 The Evolutionary Process 403

12 Biotechnology and Synthetic

Biology 354
13.7
13.8
Experimental Evolution 405
Gene Families, Duplications,
and Deletions 407
13.9 Horizontal Gene Transfer 409
MICROBIOLOGYNOW  n Ingestible Biosensor: Using Bacteria to
A 13.10 The Evolution of Microbial Genomes 410
Monitor Gastrointestinal Health 354
III   • Microbial Phylogeny and
I  
• Tools of the Genetic Engineer 355 Systematics 412
12.1 Manipulating DNA: PCR and Nucleic Acid 13.11 Molecular Phylogeny: Making Sense
Hybridization 355 of Molecular Sequences 412
12.2 Molecular Cloning 358 13.12 Microbial Systematics 416
 CONTENTS xix

UNIT 4 Microbial Diversity

15 Ecological Diversity of
Bacteria 478

14 Metabolic Diversity
of Microorganisms 424 MICROBIOLOGYNOW Cyanobacterial Diversity
and Environmental Change 478

MICROBIOLOGYNOW F erreting Out the Peculiar Life I  

• Ecological Diversity Among
of Iron Bacteria 424 Microorganisms 479
15.1 Making Sense of Microbial Diversity 479
I • Introduction to Metabolic Diversity 425
14.1 Foundational Principles of Metabolic II  • Ecological Diversity of
Diversity: Energy and Redox 425 Phototrophic Bacteria 480
14.2 Autotrophic Pathways 428 15.2 Overview of Phototrophic Bacteria 480
II   • Phototrophy 430 15.3 Cyanobacteria 481
15.4 Purple Sulfur Bacteria 485
14.3 Photosynthesis and Chlorophylls 430
15.5 Purple Nonsulfur Bacteria and Aerobic
14.4 Carotenoids and Phycobilins 434
Anoxygenic Phototrophs 487
14.5 Anoxygenic Photosynthesis 435
15.6 Green Sulfur Bacteria 488
14.6 Oxygenic Photosynthesis 438
15.7 Green Nonsulfur Bacteria 490
III   • Respiratory Processes Defined by 15.8 Other Phototrophic Bacteria 491
Electron Donor 440 III   • Diversity of Bacteria Defined by
14.7 Oxidation of Sulfur Compounds 440 Metabolic Traits 492
14.8 Iron (Fe2+) Oxidation 442
15.9 Diversity of Nitrogen Fixers 492
14.9 Nitrification 443
15.10 Diversity of Nitrifiers and Denitrifiers 494
14.10 Anaerobic Ammonia Oxidation
15.11 Dissimilative Sulfur- and Sulfate-Reducers 496
(Anammox) 445
15.12 Dissimilative Sulfur-Oxidizers 498
IV    • Respiratory Processes Defined by 15.13 Dissimilative Iron-Reducers 502
Electron Acceptor 446 15.14 Dissimilative Iron-Oxidizers 503
14.11 Nitrate Reduction and Denitrification 446 15.15 Methanotrophs and Methylotrophs 504
14.12 Sulfate and Sulfur Reduction 448 IV    • Morphologically and Ecologically
14.13 Other Electron Acceptors 450 Distinctive Bacteria 506
V  • One-Carbon (C1) Metabolism 452 15.16 Microbial Predators 506
14.14 Acetogenesis 452 15.17 Spirochetes 508
14.15 Methanogenesis 454 15.18 Budding and Prosthecate/Stalked Bacteria 511
14.16 Methanotrophy 458 15.19 Sheathed Bacteria 514
15.20 Magnetic Microbes 515
VI   • Fermentation 460
14.17 Energetic and Redox

16 Phylogenetic Diversity of
Considerations 460
14.18 Lactic and Mixed-Acid
Fermentations 462 Bacteria 519
14.19 Fermentations of Obligate
Anaerobes 464 MICROBIOLOGYNOW Bacterial Diversity and Human Health 519
14.20 Secondary Fermentations 466
14.21 Fermentations That Lack Substrate-Level I  
• Proteobacteria 520
Phosphorylation 467 16.1 Alphaproteobacteria 521
14.22 Syntrophy 469 16.2 Betaproteobacteria 524
VII  • Hydrocarbon Metabolism 471 16.3 Gammaproteobacteria: Enterobacteriales 526
16.4 Gammaproteobacteria: Pseudomonadales
14.23 Aerobic Hydrocarbon
and Vibrionales 528
Metabolism 471
16.5 Deltaproteobacteria and
14.24 Anaerobic Hydrocarbon
Epsilonproteobacteria 529
Metabolism 472
xx CONTENTS

II  • Firmicutes, Tenericutes, and Actinobacteria 531 IV    • Evolution and Life at High Temperature 578
16.6 Firmicutes: Lactobacillales 531 17.12 An Upper Temperature Limit for Microbial Life 578
16.7 Firmicutes: Nonsporulating Bacillales and 17.13 Molecular Adaptations to Life at High
Clostridiales 533 Temperature 580
16.8 Firmicutes: Sporulating Bacillales and 17.14 Hyperthermophilic Archaea, H2, and Microbial
Clostridiales 534 Evolution 581
16.9 Tenericutes: The Mycoplasmas 535
16.10 Actinobacteria: Coryneform and Propionic Acid

18 Diversity of Microbial
Bacteria 536
16.11 Actinobacteria: Mycobacterium 538
16.12 Filamentous Actinobacteria: Streptomyces and Eukarya 585
Relatives 539

III   • Bacteroidetes 542 MICROBIOLOGYNOW Coccolithophores, Engineers

of Global Climate 585
16.13 Bacteroidales 542
16.14 Cytophagales, Flavobacteriales, and I  
• Organelles and Phylogeny
Sphingobacteriales 543 of Microbial Eukarya 586
IV    • Chlamydiae, Planctomycetes, and 18.1 Endosymbioses and the Eukaryotic Cell 586
Verrucomicrobia 544 18.2 Phylogenetic Lineages of Eukarya 588
16.15 Chlamydiae 544 II  • Protists 589
16.16 Planctomycetes 546 18.3 Excavates 589
16.17 Verrucomicrobia 547 18.4 Alveolata 591
V  • Hyperthermophilic Bacteria 548 18.5 Stramenopiles 593
18.6 Rhizaria 595
16.18 Thermotogae and Thermodesulfobacteria 548
18.7 Haptophytes 596
16.19 Aquificae 549
18.8 Amoebozoa 597
VI   • Other Bacteria 550
III   • Fungi 599
16.20 Deinococcus–Thermus 550
18.9 Fungal Physiology, Structure,
16.21 Acidobacteria and Nitrospirae 551
and Symbioses 599
16.22 Other Notable Phyla of Bacteria 552
18.10 Fungal Reproduction and Phylogeny 601
18.11 Microsporidia and Chytridiomycota 602

17
18.12 Mucoromycota and Glomeromycota 603
Diversity of Archaea 556 18.13 Ascomycota 604
18.14 Basidiomycota 605

IV    • Archaeplastida 606

MICROBIOLOGYNOW Methanogens and Global Climate Change 556 18.15 Red Algae 606
18.16 Green Algae 607
I  
• Euryarchaeota 558
17.1 Extremely Halophilic Archaea 558
17.2
17.3
Methanogenic Archaea 561
Thermoplasmatales 565
UNIT 5 Microbial Ecology and
17.4 Thermococcales and Archaeoglobales 566 Environmental Microbiology

19
II  • Thaumarchaeota and Cryptic Archaeal Phyla 567
17.5 Thaumarchaeota and Nitrification in Archaea 568 Taking the Measure
17.6 Nanoarchaeota and the “Hospitable Fireball” 569 of Microbial Systems 612
17.7 Korarchaeota, the “Secret Filament” 570
17.8 Other Cryptic Archaeal Phyla 571
MICROBIOLOGYNOW T ouring Microbial Biogeography Using
III   • Crenarchaeota 572 Combinatorial Imaging 612
17.9 Habitats and Energy Metabolism of I  
• Culture-Dependent Analyses of Microbial
Crenarchaeota 572
Communities 613
17.10 Crenarchaeota from Terrestrial Volcanic Habitats 574
19.1 Enrichment Culture Microbiology 613
17.11 Crenarchaeota from Submarine Volcanic Habitats 576
 CONTENTS xxi

19.2 Classical Procedures for Isolating Microbes 617 20.14 The Deep Sea 682
19.3 Selective Single-Cell Isolation: Laser Tweezers, Flow 20.15 Deep-Sea Sediments 685
Cytometry, Microfluidics, and High-Throughput 20.16 Hydrothermal Vents 687
Methods 618

21
II  • Culture-Independent Microscopic Analyses
of Microbial Communities 620 Nutrient Cycles 693
19.4 General Staining Methods 620
19.5 Microscopic Specificity: Fluorescence In Situ
Hybridization (FISH) 622
MICROBIOLOGYNOW  n Uncertain Future for Coral Reef
A
III   • Culture-Independent Molecular Analyses Ecosystems 693
of Microbial Communities 625 I  
• Carbon, Nitrogen, and Sulfur Cycles 694
19.6 PCR Methods of Microbial Community
21.1 The Carbon Cycle 694
Analysis 626
21.2 Syntrophy and Methanogenesis 697
19.7 Microarrays for Analysis of Microbial Phylogenetic
21.3 The Nitrogen Cycle 699
and Functional Diversity 630
21.4 The Sulfur Cycle 701
19.8 Environmental Multi-omics: Integration of Genomics,
Transcriptomics, Proteomics, and II  • Other Nutrient Cycles 702
Metabolomics 631
21.5 The Iron and Manganese Cycles: Reductive
IV    • Measuring Microbial Activities in Nature 637 Activities 702
19.9 Chemical Assays, Radioisotopic Methods, 21.6 The Iron and Manganese Cycles: Oxidative
Microsensors, and Nanosensors 638 Activities 706
19.10 Stable Isotopes and Stable Isotope Probing 641 21.7 The Phosphorus, Calcium, and Silicon Cycles 708
19.11 Linking Functions to Specific Organisms 643 III   • Humans and Nutrient Cycling 710
19.12 Linking Genes and Cellular Properties to
21.8 Mercury Transformations 711
Individual Cells 646
21.9 Human Impacts on the Carbon and Nitrogen
Cycles 713

20 Microbial Ecosystems 651 Explore the Microbial World

Solving the Marine Methane Paradox 710

MICROBIOLOGYNOW

I  
Living on Fumes 651

• Microbial Ecology 652

20.1 General Ecological Concepts 652
22 Microbiology of the Built
Environment 718

20.2 Ecosystem Service: Biogeochemistry MICROBIOLOGYNOW  ending Microbes to Clean Up

S
and Nutrient Cycles 653 after Polluters 718
II  • The Microbial Environment 654 I  
• Mineral Recovery and Acid Mine Drainage 719
20.3 Environments and Microenvironments 654 22.1 Mining with Microorganisms 719
20.4 Surfaces and Biofilms 656 22.2 Acid Mine Drainage 721
20.5 Microbial Mats 659
II  • Bioremediation 722
III   • Terrestrial Environments 661 22.3 Bioremediation of Uranium-Contaminated
20.6 Soils: General Properties 661 Environments 722
20.7 Prokaryotic Diversity in Soils 664 22.4 Bioremediation of Organic Pollutants:
20.8 The Terrestrial Subsurface 666 Hydrocarbons 723
22.5 Bioremediation and Microbial Degradation
IV    • Aquatic Environments 669 of Major Chemical Pollutants: Chlorinated
20.9 Freshwaters 669 Organics and Plastics 724
20.10 Oxygen Relationships in the Marine
Environment 671 III   • Wastewater and Drinking Water Treatment 727
20.11 Major Marine Phototrophs 674 22.6 Primary and Secondary Wastewater Treatment 728
20.12 Pelagic Bacteria and Archaea 677 22.7 Tertiary Wastewater Treatment: Further Removal
20.13 Pelagic Marine Viruses 680 of Phosphorus and Nitrogen 730
xxii CONTENTS

22.8 Sludge Processing and Contaminants

of Emerging Concern 732 UNIT 6 Microbe–Human Interactions
22.9 Drinking Water Purification and Stabilization 735 and the Immune System
22.10 Water Distribution Systems 736

24
IV    • Indoor Microbiology and Microbially
Influenced Corrosion 737
Microbial Symbioses with
22.11 The Microbiology of Homes and Humans 783
Public Spaces 737
22.12 Microbially Influenced Corrosion MICROBIOLOGYNOW  ne of the Most Abundant Viruses on
O
of Metals 739 Earth Discovered First in the Human
22.13 Biodeterioration of Stone and Concrete 740 Viral Microbiome 783

I  
• Structure and Function of the Healthy Adult

23 Microbial Symbioses with Gastrointestinal and Oral Microbiomes 784

24.1 Overview of the Human Microbiome 784
Microbes, Plants, and
24.2 Gastrointestinal Microbiota 785
Animals 744 24.3 Oral Cavity and Airways 791

II  • Urogenital Tract and Skin Microbiomes

MICROBIOLOGYNOW  oral Fluorescence Provides the Guiding
C and the Human Viral Microbiome 794
Light for Their Symbiotic Algae 744
24.4 Urogenital Tracts and Their Microbes 794
I  
• Symbioses Between Microorganisms 745 24.5 The Skin and Its Microbes 795
23.1 Lichens 745 24.6 The Human Virome 797
23.2 “Chlorochromatium aggregatum” 746 III   • From Birth to Death: Development of the
23.3 Methanotrophic Consortia: Direct Interspecies Human Microbiome 800
Electron Transfer 748
24.7 Human Study Groups and
II  • Plants as Microbial Habitats 749 Animal Models 800
23.4 The Legume–Root Nodule Symbiosis 749 24.8 Colonization, Succession, and Stability
of the Gut Microbiota 801
23.5 Mycorrhizae 755
23.6 Agrobacterium and Crown Gall Disease 757 IV    • Disorders Attributed to the Human
III   • Insects as Microbial Habitats 759 Microbiome 803
23.7 Heritable Symbionts of Insects 759 24.9 Syndromes Linked to the Gut Microbiota 804
23.8 Defensive Symbioses 762 24.10 Syndromes Linked to the Oral, Skin,
and Vaginal Microbiota 807
23.9 Termites 763

IV    • Other Invertebrates as Microbial Habitats 765
23.10 Bioluminescent Symbionts and the Squid
Symbiosis 765
23.11 Marine Invertebrates at Hydrothermal Vents
and Cold Seeps 769
23.12 Entomopathogenic Nematodes 770
23.13 Reef-Building Corals 771
V  • Modulation of the Human Microbiome 809
24.11 Antibiotics and the Human Microbiome 809
24.12 Probiotics, Prebiotics, and Synbiotics 810
Explore the Microbial World
The Gut–Brain Axis 790

V  • Mammalian Gut Systems as Microbial

Habitats 774
23.14 Alternative Mammalian Gut Systems 774
25 Microbial Infection and
Pathogenesis 814
23.15 The Rumen and Rumen Activities 776 MICROBIOLOGYNOW Killing Pathogens on Contact 814
23.16 Rumen Microbes and Their Dynamic
Relationships 777 I  
• Human–Pathogen Interactions 815
Explore the Microbial World 25.1 Microbial Adherence 815
Combating Mosquito-Borne Viral Diseases with 25.2 Colonization and Invasion 817
an Insect Symbiont 761
 CONTENTS xxiii

25.3 Pathogenicity, Virulence, and Virulence II  • Antibodies 862

Attenuation 819 27.3 Antibody Production and Structural
25.4 Genetics of Virulence and the Diversity 862
Compromised Host 820 27.4 Antigen Binding and the Genetics of
II  • Enzymes and Toxins of Pathogenesis 822 Antibody Diversity 866
25.5 Enzymes as Virulence Factors 822 III   • The Major Histocompatibility
25.6 AB-Type Exotoxins 824 Complex (MHC) 869
25.7 Cytolytic and Superantigen Exotoxins 827 27.5 MHC Proteins and Their Functions 869
25.8 Endotoxins 828 27.6 MHC Polymorphism, Polygeny, and Peptide
Binding 871

26
IV    • T Cells and Their Receptors 873
Innate Immunity: Broadly
27.7 T Cell Receptors: Proteins, Genes,
Specific Host Defenses 832 and Diversity 874
27.8 T Cell Subsets and Their Functions 877
MICROBIOLOGYNOW  eriodontal Disease and Alzheimer’s:
P
Evidence for Causation? 832

I  
• Fundamentals of Host Defense 833
26.1
26.2
Basic Properties of the Immune System 833
Barriers to Pathogen Invasion 834
28 Immune Disorders and
Antimicrobial Therapy 883

MICROBIOLOGYNOW  reventing Autoimmunity with . . . Parasitic

P
II  • Cells and Organs of the Worms? 883
Immune System 836
26.3 The Blood and Lymphatic Systems 836 I  
• Disorders and Deficiencies of the Immune
26.4 Leukocyte Production and Diversity 838 System 884
28.1 Allergy, Hypersensitivity, and Autoimmunity 884
III   • Phagocyte Response Mechanisms 840
28.2 Superantigens and Immunodeficiency 887
26.5 Pathogen Challenge and Phagocyte
Recruitment 840 II  • Vaccines and Immunotherapy 889
26.6 Pathogen Recognition and Phagocyte 28.3 Vaccination Against Infectious Diseases 889
Signal Transduction 841 28.4 Immunotherapy 892
26.7 Phagocytosis and Phagocyte
Inhibition 844 III   • Drug Treatments for Infectious Diseases 894
28.5 Antibacterial Drugs 894
IV    • Other Innate Host Defenses 846
28.6 Antimicrobial Drugs That Target Nonbacterial
26.8 Inflammation and Fever 846 Pathogens 900
26.9 The Complement System 848 28.7 Antimicrobial Drug Resistance and New Treatment
26.10 Innate Defenses Against Viruses 851 Strategies 902
Explore the Microbial World
Pattern Recognition Receptors of Hydrothermal
Vent Tube Worms Facilitate Endosymbiosis 843 UNIT 7 Infectious Diseases

27 Adaptive Immunity: Highly

Specific Host Defenses 856 29 Diagnosing Infectious
Diseases 907

MICROBIOLOGYNOW  hedding New Light on Diagnosing

S
MICROBIOLOGYNOW  ontrolling HIV through “Public” T Cell Receptors
C Tuberculosis 907
on CD4 T Cells 856
I  
• Microbiology and the Healthcare
I  
• Principles of Adaptive Immunity 857 Environment 908
27.1 Specificity, Memory, Selection Processes, 29.1 The Clinical Microbiology Laboratory 908
and Tolerance 857
29.2 Healthcare-Associated Infections 909
27.2 Immunogens and Classes of Immunity 860
xxiv CONTENTS

II  • Isolating and Characterizing Infectious II  • Airborne Viral Diseases 959
Microorganisms 910 31.6 MMR and Varicella-Zoster Infections 959
29.3 Workflow in the Clinical Laboratory 910 31.7 The Common Cold 961
29.4 Choosing the Right Treatment 916 31.8 Influenza 962

III   • Immunological and Molecular Tools III   • Direct-Contact Bacterial

for Disease Diagnosis 918 and Viral Diseases 964
29.5 Immunoassays and Disease 918 31.9 Staphylococcus aureus Infections 965
29.6 Precipitation, Agglutination, and 31.10 Helicobacter pylori and Gastric Diseases 966
Immunofluorescence 920 31.11 Hepatitis 967
29.7 Enzyme Immunoassays, Rapid Tests, and 31.12 Ebola: A Deadly Threat 969
Immunoblots 922
29.8 Nucleic Acid–Based Clinical Assays 925 IV    • Sexually Transmitted Infections 970
31.13 Gonorrhea, Syphilis, and Chlamydia 971
Explore the Microbial World
31.14 Herpes Simplex Viruses (HSV)
MRSA—A Formidable Clinical Challenge 912 and Human Papillomavirus (HPV) 975
31.15 Human Immunodeficiency

30
Virus (HIV) and AIDS 976
Epidemiology and Public
Health 929

MICROBIOLOGYNOW  New Urgent Threat is Emerging in Public

A
Health Microbiology 929
32 Vectorborne and
Soilborne Bacterial
and Viral Diseases 983
I  
• Principles of Epidemiology 930
30.1 The Language of Epidemiology 930 MICROBIOLOGYNOW T he Historical Emergence of an
30.2 The Host Community 932 Ancient and Deadly Pathogen 983
30.3 Infectious Disease Transmission
and Reservoirs 933 I  
• Animal-Transmitted Viral Diseases 984
30.4 Characteristics of Disease Epidemics 935 32.1 Rabies Virus and Rabies 984
32.2 Hantavirus and Hantavirus Syndromes 986
II  • Public and Global Health 937
30.5 Public Health and Infectious Disease 937 II  • Arthropod-Transmitted Bacterial and Viral
30.6 Global Health Comparisons 939 Diseases 987
32.3 Rickettsial Diseases 987
III   • Emerging Infectious Diseases, Pandemics, 32.4 Lyme Disease and Borrelia 989
and Other Threats 940 32.5 Yellow Fever, Dengue Fever, Chikungunya,
30.7 Emerging and Reemerging Infectious Diseases 940 and Zika 991
30.8 Examples of Pandemics: HIV/AIDS, Cholera, 32.6 West Nile Fever 993
and Influenza 943 32.7 Plague 994
30.9 Public Health Threats from Microbial Weapons 945
III   • Soilborne Bacterial Diseases 996
32.8 Anthrax 996

31 Person-to-Person Bacterial 32.9 Tetanus and Gas Gangrene 997

and Viral Diseases 950

MICROBIOLOGYNOW  eversing Antibiotic Resistance in a

R
Recalcitrant Pathogen 950 33 Waterborne and
Foodborne Bacterial
and Viral Diseases 1001
I  
• Airborne Bacterial Diseases 951
31.1 Airborne Pathogens 951
31.2 Streptococcal Syndromes 952 MICROBIOLOGYNOW Reverse Zoonosis in the Southern Ocean 1001
31.3 Diphtheria and Pertussis 955
I  
• Water as a Disease Vehicle 1002
31.4 Tuberculosis and Leprosy 956
33.1 Agents and Sources of Waterborne Diseases 1002
31.5 Meningitis and Meningococcemia 958
33.2 Public Health and Water Quality 1003
 CONTENTS xxv

II  • Waterborne Diseases 1004 II  • Visceral Parasitic Infections 1028

33.3 Vibrio cholerae and Cholera 1004 34.3 Amoebae and Ciliates: Entamoeba,
33.4 Legionellosis 1006 Naegleria, and Balantidium 1028
33.5 Typhoid Fever and Norovirus Illness 1007 34.4 Other Visceral Parasites: Giardia,
Trichomonas, Cryptosporidium,
III   • Food as a Disease Vehicle 1008 Toxoplasma, and Cyclospora 1029
33.6 Food Spoilage and Food Preservation 1008
33.7 Foodborne Diseases and Food Epidemiology 1010
III   • Blood and Tissue Parasitic
Infections 1031
IV    • Food Poisoning 1012 34.5 Plasmodium and Malaria 1031
33.8 Staphylococcal Food Poisoning 1012 34.6 Leishmaniasis, Trypanosomiasis,
33.9 Clostridial Food Poisoning 1013 and Chagas Disease 1033
34.7 Parasitic Helminths: Schistosomiasis
V  • Food Infection 1014 and Filariases 1034
33.10 Salmonellosis 1014
Photo Credits 1039
33.11 Pathogenic Escherichia coli 1015
33.12 Campylobacter 1016 Glossary Terms 1043
33.13 Listeriosis 1017
Index 1047
33.14 Other Foodborne Infectious Diseases 1018

34 Eukaryotic Pathogens:
Fungi, Protozoa, and
Helminths 1023

MICROBIOLOGYNOW  Silver Bullet to Kill Brain-Eating

A
Amoebae? 1023

I  
• Fungal Infections 1024
34.1 Pathogenic Fungi and Classes of Infection 1024
34.2 Fungal Diseases: Mycoses 1026
ASM Recommended Curriculum Guidelines
for Undergraduate Microbiology
T he American Society for Microbiology (ASM) endorses a concept-
based curriculum for undergraduate microbiology, emphasizing
skills and concepts that have lasting importance beyond the classroom
and laboratory. The ASM (in its Curriculum Guidelines for Understanding
Microbiology Education) recommends deep understanding of 27 key con-
cepts, 4 scientific thinking competencies, and 7 key skills. These guide-
lines follow scientific literacy reports and recommendations from the
American Association for the Advancement of Science and the Howard
Hughes Medical Institute by encouraging an active learning, student-
based course. Consider these guiding statements as you progress through
this book and master principles, problem solving, and laboratory skills
in microbiology.
ASM Guideline Concepts
and Statements
Evolution: Chapters 1, 9, 10–14, 20, 30
• Cells, organelles (e.g., mitochondria and chloroplasts), and all major
metabolic pathways evolved from early prokaryotic cells.
• Mutations and horizontal gene transfer, with the immense variety
of microenvironments, have selected for a huge diversity of microor-
ganisms.
• Traditional concept of species is not readily applicable to microbes
due to asexual reproduction and the frequent occurrence of horizon-
tal gene transfer.
• Evolutionary relatedness of organisms is best reflected in phylogenetic
trees.
• Human impact on the environment influences the evolution of
microorganisms (e.g., emerging diseases and the selection of antibi-
otic resistance).
Cell Structure and Function: Chapters 1, 2, 5, 8,
11, 18
• Structure and function of microorganisms have been revealed by the
use of microscopy (including bright-field, phase contrast, fluores-
cence, super-resolution, and electron).
• Bacteria have unique cell structures that can be targets for antibiotics,
immunity, and phage infection.
• Bacteria and Archaea have specialized structures (e.g., flagella, endo-
spores, and pili) that often confer critical capabilities.
• While microscopic eukaryotes (for example, fungi, protozoa, and
algae) carry out some of the same processes as bacteria, many of the
cellular properties are fundamentally different.
• Replication cycles of viruses (lytic and lysogenic) differ among viruses
and are determined by their unique genomes.
xxvi
Metabolic Pathways: Chapters 1, 3, 4, 7, 8, 12, 14
• Bacteria and Archaea exhibit extensive, and often unique, metabolic
diversity (e.g., nitrogen fixation, methane production, anoxygenic
photosynthesis).
• Interactions of microorganisms among themselves and with their
environment are determined by their metabolic abilities (e.g., quorum
sensing, oxygen consumption, nitrogen transformations).
• Survival and growth of any microorganism in a given environment
depends on its metabolic characteristics.
• Growth of microorganisms can be controlled by physical, chemical,
mechanical, or biological means.
Information Flow and Genetics: Chapters 1, 5–13
• Genetic variations can impact microbial functions (e.g., in biofilm
formation, pathogenicity, and drug resistance).
• Although the central dogma is universal in all cells, the processes of
replication, transcription, and translation differ in Bacteria, Archaea,
and eukaryotes.
• Regulation of gene expression is influenced by external and internal
molecular cues and/or signals.
• Synthesis of viral genetic material and proteins is dependent on
host cells.
• Cell genomes can be manipulated to alter cell function.
Microbial Systems: Chapters 1, 15–34
• Microorganisms are ubiquitous and live in diverse and dynamic
ecosystems.
• Many bacteria in nature live in biofilm communities.
• Microorganisms and their environment interact with and modify each
other.
• Microorganisms, cellular and viral, can interact with both human and
nonhuman hosts in beneficial, neutral, or detrimental ways.
Impact of Microorganisms: Chapters 1, 6–8, 12,
19–34
• Microbes are essential for life as we know it and the processes that
support life (e.g., in biogeochemical cycles and plant and/or animal
microbiota).
• Microorganisms provide essential models that give us fundamental
knowledge about life processes.
• Humans utilize and harness microorganisms and their products.
• Because the true diversity of microbial life is largely unknown, its
effects and potential benefits have not been fully explored.
Making Connections Across

UPDATED! Each chapter is carefully cross-referenced to

connect students with related material found earlier (◀ )
or later ( ▶) in the book.

NEW! Key Concept statements at the start of

each major part of a chapter give students a big picture
view of the content to come before they dive in and
immerse themselves in the details.
Another random document with
no related content on Scribd:
 ‘My dear Friend,—I cannot let Mrs. Young return without
sending you my best thanks for the second edition of your
excellent pamphlet. Indeed, if I were singular in approbation of it,
you might think me a cleverer fellow than I shall seem among the
crowd of your admirers. What is a single name in a list fifty yards
long? And if I were to tell you what numbers of first-rate judges
have spoke well of your performance, I should want more room
than Mr. Sheridan’s friends at Glasgow. I shall only just specify
those who would be at the head of a complete list, if I had time to
make one: Mr. Burke, Lord Orford, who, on my asking him if he
had seen your pamphlet, pointed to it, “There it is; I read nothing
else;” Mrs. Montagu the same; a large party of bluestockings at
Lady Hesketh’s all agreed that your book and Hannah More’s
“Chip”[155] were the best on the subject. When I made Mrs. Crewe
read the first edition, she wrote me word that she had perused it
with great attention, and that she found it contained stubborn
facts, to each of which she should say with the grave-digger,
“Answer me that and unyoke.” Last week, in a note she sent me
from Hampstead, she says: "Mr. Arthur Young’s pamphlet makes
a great noise, and, I think, I never knew any book take more; it is
reprinted, you know, with additions." In the communication of the
latter information she got the start of me; the second edition
could not have been out three days but you are meditating a
third. I like your additions to the second much, particularly what
concerns the reform of Parliament.
‘I wish you could overhaul Grey’s speech as well as Charles
Fox’s on that subject, and in an appendix expose the weakness
and inconsistence of both. Only observe how both confess that
there was danger to our constitution “from opinions favourable to
the principles and measures of France,” after so stubbornly and
pertinaciously denying in Parliament the existence of any such
danger; challenging Government to prove it, and saying that “the
Proclamation, call of Parliament, Alien and Traitorous
Correspondence Bills, were mere Ministerial juggles” to increase
influence, diminish liberty, and encourage excess of loyalty.
“But,” says Mr. G. “that danger must now be much lessened, as
all approbation of those principles, or imitation of that example, is
now improbable, totally discredited, and removed from all
political speculation and practice.” What, then, is all the defence
of France and Frenchmen by the Opposition? And why is every
measure condemned in Parliament that tends to put an end to
their anarchy and ambition? Why is war against them so
censured? Why is it always called the war of kings and despots?
Why is the Minister so importuned to make peace with regicides
and assassins, determined to force, if possible, every nation
upon earth to adopt their measures? Mr. Grey repeats in his
speech, “All dread of the example is completely removed, and
that none could suppose him, or any other party in this country,
favourable to that example.” What is this but open falsehood?
Mr. Fox allows that “there was a party whose wild theories
certainly aimed at an impracticable perfection, that could only
have been pursued by means subversive of every part of our
constitution.” Yet there never was any danger! Mr. G. says, in
express terms, “that his motion extended to an alteration in the
present government of the country.” But he had no specific plan
ready of his own, or that he chose to father. But as all the
petitions he and Mr. Sheridan brought in for a reform demanded
nothing less than universal suffrage, and as these gentlemen
either drew up or approved the contents of these petitions, we
may easily judge what was the general plan of our Jacobins, if
they could have had the tinkering of the constitution.
‘Grey seems to me a silly fellow, with a greater wish than
abilities to do mischief. Charles Fox’s speech is more a
panegyric on the constitution than on his friend’s motion. When
every man is left to himself to reform an old constitution or make
a new one, no two will be found of a mind on the subject.
Sherry’s speech was nothing to the purpose. There was no
attempt of the phalanx which I so much dreaded, as the doubling
our tiers état. Thank God, their great gun has flashed in the pan!
The mountain has laboured in vain. You know I hope that the
gang in Parliament, like Egalité’s creatures in the “Convention,”
is called the Mountain. And it has been called by a punster of the
party Mount Sigh-on. I fear the war will be long and bloody; and
how it will end who can tell? Nothing but a vigorous prosecution
of the war can save the whole civilised globe from destruction.
After disdaining in the House the principles which they had
suggested and encouraged out of it, I should not wonder if the
Scotch and English petitioners for reform on the basis of
universal suffrage should mob and September their friends the
demagogues whenever they can catch them. I don’t love
mischief, but I do cordially wish something of that kind were to
happen.
‘Adieu.
‘Ever yours sincerely,
‘Charles Burney.’

From Dr. Symonds, high encomiums of my ‘Example,’ &c.

‘Prince of Wales Coffee House: April 8, 1793.

‘Traveller Coxe[156] desired me to tell you how charmed he was

with your pamphlet; nay, he had begun to write five or six lines to
you, but thought afterwards it was taking too great a liberty. One
thing, however, he wishes you to expunge in your next edition,
viz.: a reflection on Sunday schools as not being founded on
truth. You must not be surprised at this, for he is a zealous
patron of them, and has explained the Catechism in print for that
purpose. Wherever I go I hear your “Example of France” spoken
of in the highest terms as to the matter.
‘Everyone agrees that no political writer whatever has set the
representation of property in so clear and just a light. Bishop
Douglas, who has written many good pamphlets, and is therefore
the best judge, makes no scruple to declare frequently that you
deserve from Government a most ample reward; but we both
wish, as well as others, for your sake, that the second edition
may be printed more correctly.
‘You should come to town and be presented, or, at least, take
an opportunity to walk on the Terrace at Windsor, where you
would not fail of being marked out. Bishop Watson’s appendix
has rendered him rectus in curia. A few days ago he was at
Court, talking with Lord Dartmouth, who mentioned the word
philosophy, which the King overhearing, came to the bishop, and
said, “I have read the best sort of philosophy, my lord, in your
sermon and appendix, which has wonderfully pleased me.” The
bishop, of course, made his bow, and then the King went on,
“You write so concisely and so forcibly, that everyone must be
convinced by your arguments;” on which the bishop replied, “I
like, Sir, to step forward in a moment of danger.” The King
rejoined, “You have shown a good spirit, and it could not be done
in a better manner.” Should the last volume of "Clarendon’s
Letters" come in your way, I would advise you to read the famous
one from Sir John Colepeper to Secretary Nicholas; which is
always esteemed as a wonderful instance of political sagacity, as
it foretold that the Restoration would be accomplished by Monk!
But I think there is another part of this letter which shows equal
sagacity, viz.: his desiring that Charles would not send over any
foreign troops into England, as this measure would not fail of
uniting the English against him; whereas, if they were left to
themselves, he would always have a strong party, and must
sooner or later be restored. I am fully convinced of the truth of
this reasoning; and of what use is history unless it be considered
as a school for modern politicians?
‘Why did you not let me know whether your second edition had
gone to the press or not? Before I left Cambridge I saw a
gentleman who told me that Sir William Scott had mentioned in a
letter to one of his friends there that it was by far the most
convincing and best pamphlet that had been published.
‘All I could wish is that you had not stigmatised all reformers
with the name of enemies to the state; or, at least, you intimated
it. I was always myself an enemy to reform in Parliament, and
continue to be so; yet I know some warm advocates for it, who
mean as well to the benefit of this country as you can possibly
do.
‘Dr. Hardy and Sir Henry Moncrief (a Scotch clergyman) are
come to solicit a Bill for the enlarging of the stipends of the
Scotch clergy. They do not apprehend much difficulty in carrying
it through the Houses, though the addition must be supplied out
of the tithes in the hands of lay proprietors. Hardy is Professor of
Ecclesiastical History at Edinburgh—a most sensible man, with
great liberality of mind. Sir Henry is a polished man, and likewise
a man of business. I hope to see them both at St. Edmund’s Hill,
and you must meet them. You should get Hardy’s pamphlet, the
“Patriot,” published in Scotland on the present emergency; there
are in it many excellent things.
‘You seem in your letter to be still apprehensive of some plots
and insurrections.
‘Plot! Plots! was the catch-word in King Charles II.’s time. Sir
H. Moncrief and Dr. Hardy laughed at Dundas’s account of the
political riots in Scotland. They absolutely denied the existence
of them—considered them as political; and when you read
Hardy’s pamphlet, you will see that he would not have failed
setting them forth if they had deserved any consideration.
‘Adieu! I should not have come to London had it not been on
account of my ecclesiastical foundling.
‘John Symonds.’

From Dr. Symonds

‘September 1, 1793.

‘My dear Sir,—I do not wonder that you smiled at the affected
secrecy of Macpherson concerning the Censomento. The book
to which he alludes cannot be the “Bilancio dello stato,” &c.,
which was written to please Count Firmian. I knew well the
gentleman who wrote it and gave it to me, as I often met him at
dinner at the count’s.
‘Sometimes he was too decisive. One day he said at the
count’s table that the Bresciano contained 800,000 inhabitants
now. As Count F. knew that I had just come from Brescia, and
had not lost my time there, he asked me what number there was;
on which I told him that there were 376,000 according to a
census taken a few years before. The count smiled, and looked
very attentively on Carpani (for that was the author’s name), who
never liked me so well after that day, nor had Count Firmian so
high an opinion of him. You possibly may not know the history of
Sir John Macpherson. He offered the Duke of Grafton, when he
was at the head of the Treasury, a vast collection of jewels, by
order of the Nabob of Arcot, which the duke absolutely refused,
and Bradshaw, his secretary, also. Sir John, thinking that the
nabob would not believe that he had offered the present,
published for his own vindication the answers of the duke and
Bradshaw, for which he was turned out of the company’s service,
as he was pursuing an interest then opposite to its interests.
‘Scotch influence not long after restored him. You will find the
letters in Lind’s appendix to the defence of Lord Pigot. You are
now, of course, so much of a politician as not to be surprised
(shall I say disgusted?) at Macpherson’s conduct. The opinion of
the King of Prussia as to your book I value not a straw; but that
of the Marshal de Castries certainly carries with it great weight.
‘He is one of the few who adhered to Necker from gratitude,
when the latter was turned out of his post about ten years ago;
and I heard a very good character of the maréchal when I was
last in France. St. Paul, as you and the duke are pleased to call
him, is finished, and the preface is on the stocks.[157]
‘Why do you wish Clarke had commented on the Epistle to the
Romans? Locke and Taylor have done it admirably; and easy as
you may think the Gospels are, they have been rendered much
more so by Clarke.
‘What do you mean by saying that the Gospels want no
explanation? St. John is extremely difficult in some parts,
notwithstanding Clarke’s paraphrase; and I think, with Markland,
that he is as yet very far from being perfectly understood. Adieu!
‘I remain,
‘Ever your sincere friend,
‘John Symonds.’
 CHAPTER XI

THE SECRETARYSHIP, 1794-95-96

The Secretaryship and its drawbacks—Social compensations—Illness and

death of Elizabeth Hoole—Letters of Jeremy Bentham and others—A visit
to Burke—Home travels—Enclosures.

The Board of Agriculture, meeting in February, arranged the

President’s plan for the attendance of their officers. By these
laws all the officers of the Board were bound to attend, with no
other exception than the months of August, September and
October, with one month at Christmas and three weeks at
Easter. These laws, ready cut and dried when the Board met,
were adopted with no other alterations than such as the
President himself had made in them, previously to their being
presented at the meeting. Lord Hawke had examined the rules
and orders of many societies, and found that in all letters
communications were addressed to the Secretaries, and
answers given by them. Sir John Sinclair struck this out, and
directed all such communications to be to the President
(himself), and for him also to sign all letters. This at once
converted the Secretary into nothing more than a first clerk. I
saw not at first the tendency of the alterations; but I soon felt
their effect. All letters were dictated by the Secretary and written
in a book; this book was altered and corrected at the will of the
President, and such alterations made as in respect of agriculture
were absurd enough; the whole done in such a manner as not to
be very pleasing.
 In addition to this, Sir John Sinclair gave the Board the use of
his house, which ensured another circumstance hostile to my
feelings. There was only one room for transacting the business,
by the Secretary, under-Secretary, two clerks, to which Sir J.
after added the constant attendance of an attorney, for assisting
in the business of a general Enclosing Act, about which the
President busied himself some years in vain. As I was
determined to pass all the vacations at my farm in Suffolk, six
journeys of myself and servants became necessary, and caused
a considerable expense. I also was compelled to hire lodgings at
the expense of two or two guineas and a half per week, and
when I experienced the full career[158] of all these circumstances,
I deliberated repeatedly and carefully with myself, whether it
would not be cheaper to me to throw up the employment. Long
after, upon review of the whole, I was amazed that I had not
done it, more especially as my plan for settling on the moors in
Yorkshire was offered to my choice. I was infinitely disgusted
with the inconsiderate manner in which Sir John Sinclair
appointed the persons who drew up the original reports, men
being employed who scarcely knew the right end of a plough;
and the President one day desired I would accompany him with
one of these men, a half-pay officer out of employment, to call on
Lord Moira to request his assistance in the Leicestershire
Report, when this person told his Lordship that he was out of
employment and should like a summer’s excursion. To do him
justice, he did not know anything of the matter. Still, however, he
was appointed, and amused himself with his excursion to
Leicester. But the most curious circumstance of effrontery was,
that the greater number of the reporters were appointed, and
actually travelled upon the business before the first meeting of
the Board took place, under the most preposterous of all ideas—
that of surveying the whole Kingdom and printing the Reports in
a single year; by which manœuvre Sir John thought he should
establish a great reputation for himself. Consequently by his sole
authority, who could not possibly know whether the members of
the Board would approve or not such a plan. I was a capital idiot
not to absent myself sufficiently to bring the matter to a question,
and leave them to turn me out if they pleased. Mr. Pitt would
probably have interfered and effected the object I wanted, and, if
not, would have provided for me in a better way. However, I
made use of the opportunities that offered to frequent the
company of those that were agreeable to me; for a part of the
time was pretty regularly passed at the conversaziones of Mrs.
Matthew Montagu and the Countess of Bristol, where I met an
assemblage of persons remarkable for every characteristic of the
bas-bleu mixed with great numbers of the highest rank. [I was]
also at many similar parties upon a smaller scale at Mr. Charles
Coles’, the intimate friend of Soame Jenyns, and to whom he left
the property of his works. The petits soupers at Mrs. Matthew
Montagu’s, and to which she asked a selection of eight or nine
persons, were very pleasant, the conversations interesting, and
this select number more agreeable than I ever found full rooms.
On my first coming to town in the spring of 1794, I enquired of
several members of the Board whether there was not a farmers’
club in London, and was surprised that there never had been any
institution of the kind. I determined to endeavour at establishing
one, and spoke to the Duke of Bedford and the Earls of
Egremont and Winchilsea, who much approved the idea, and
applying also to a few more, I directed cards to be sent them
from the Thatched House Tavern,[159] in order to establish a club.
This meeting was fully attended, and a book being called for, the
club was instituted, and several rules entered, and the meetings
appointed once a fortnight during the sittings of Parliament. This
club became very fashionable, and applications to be elected
were very numerous, from the members of both Houses of
Parliament; and it subsists to this day, but has for some time
been very ill attended. This was occasioned by too free an
election of all who offered. While the club was limited to fifty
members it was well attended, but afterwards such numbers
were received, and with so much facility, as greatly to injure the
establishment. I have one remark to make upon clubs; the life
and soul of them is limitation to a selected few, and to blackball
the great mass of applicants, selecting merely such as will form a
very valuable addition to the society, which probably may not
amount to more than one in twenty. The annual subscription was
two guineas: one to the house, one to form a fund at the
disposition of the club. The latter gradually accumulated till it
amounted to 700l. or 800l. Both Sir John Sinclair and I were
strenuous that this might be applied to some useful purpose, and
with difficulty we got an appropriation of fifty guineas as a reward
for the best plough that could be produced; but the money
assigned to advertisements being much too small, the offer was
unknown, and no plough produced.
A member once proposed that the 800l. might be given to
charitable institutions; but this was negatived in an instant, and
the sum is still left (1812) unemployed in the funds.
While the club flourished the members who most generally
attended were the Dukes of Bedford, Buccleugh, Montrose, the
Earls of Egremont, Winchester and Darnley, the Lords of
Wentworth, Somerville, de Dunstanville, Sheffield, &c. &c.
I often dined at Charles Coles’, where I met repeatedly Jacob
Bryant,[160] Mrs. Montagu, Mrs. York, Mrs. Garrick, Hannah More,
Mrs. Orde and Soame Jenyns. The conversation at these parties
on the publications of the day, anecdotes of the time, with the
conduct of many of the great men of the age, was usually very
interesting. Alas! alas! how few of these persons are now left. I
was very eager in listening to every word that fell from Hannah
More, though not nearly so much so as I should have been many
years after.
I had an incessant round of dinners and many evening parties,
and generally with people of the highest rank and consequence,
but I was not pleased, being discontented with my employment,
and disgusted with the frivolous business of the Board, which
seemed to me engaged in nothing that could possibly produce
the least credit with the public. After five months’ residence at
London, I went to the Duke of Bedford’s at Woburn on my way to
Bradfield, spending some days very agreeably in company that
could not fail of being interesting.
This year my second daughter Elizabeth, who, as I have
mentioned before, was married to the Rev. John Hoole, died of
consumption. She was of a most amiable, gentle temper, and in
a resigned frame of mind, which gave me much satisfaction. The
last visit I paid her at Abinger, in Surrey, she was very weak, yet
not suspected to be so near her end. But at the last parting with
me, she did it in so feeling and affectionate a manner as seemed
to imply that she thought she should see me no more. It made
me, for a time, extremely melancholy, which was shaken off with
great difficulty. I took a tour into Hampshire, where I passed
several days with Mr. Poulett at Sombourne, taking an account of
the agriculture of that district, the result of which examination
was printed as an appendix to the original Hampshire Report.
On the meeting of the Board in 1793, Sir John Sinclair had
particularly requested me to draw up a Report for the County of
Suffolk, to effect which I took several journeys into different parts
of the county at some expense, and formed the Report which
was printed in 1794. I never executed any work more
commended in Suffolk than this. I had no remuneration.
Letters received this year:—
From Jeremy Bentham, Esq., enquiries into the landed
property of Great Britain and into the rental and value of houses:
—

‘Hendon, Middlesex: Sept. 1794.

‘Dear Sir,—Permit my ignorance to draw upon your science on

an occasion that happens just now to be a very material one to
me. I have a sort of floating recollection of a calculation, so
circumstanced, either in point of authority or argument, as to
carry weight with it, in which the total value of the landed
property in this country (Scotland, I believe, included) was
reckoned at a thousand millions, and that of the movable
property at either a thousand millions or twelve hundred millions.
Public debt did not come, I think, at least, it ought not to come,
into the account; it being only so much owned by one part of the
proprietors of the two thousand or the two thousand two hundred
millions to another.
‘Upon searching your book on France, which was the source
from whence I thought I had taken the idea, I can find no
calculation of the value of the movable property, nor even of the
immovable in an explicit form; on the contrary, in the instance of
the immovable, I find suppositions with which any such estimate
appears to be incompatible. The land tax at four shillings, I find,
you suppose, were it to be equal all over the country, would be
equivalent to as much as three shillings, on which supposition
the rental (the tax of four shillings producing no more than two
thousand millions) would amount to no more than 13,000,000l.
nor consequently the value, at so many years’ purchase, say
twenty-eight, to more than three hundred and sixty-four millions;
or at thirty, to three hundred and ninety millions; to which, in
order to complete the calculation of the landed property of Great
Britain, that of Scotland would have to be added.
‘The population of the three kingdoms you reckon in two
places at eleven millions; but in another place at fifteen. Is the
latter a slip of the pen? or, in the two former places, was only two
kingdoms (England and Scotland) in your view, though three are
mentioned? A circumstance that seems to favour the latter
supposition is, that the population of Ireland is well known (if I do
not much misrecollect) from recent and authentic sources to be a
little more than four millions; and as Scotland turns out to contain
a million and a half, this would leave nine and a half millions for
England, which, I should suppose, would quadrate in round
numbers with Mr. Howlett’s calculations, to which we refer; a
book which, from forgetfulness, I have never made myself
master of, and to which, being in the country, I have no speedy
means of recurring.
‘Now what I wish for is as follows: (1) a calculation (or, I should
rather say, the result) of the value of the landed property of Great
Britain reckoned at [so many] years’ purchase, two prices—a
peace price and a war price—could they be respectively of
sufficient permanence to be ascertained, would be of use.
‘(2) A calculation of the value of the personal, i.e. immovable
property of Great Britain.
‘(3) The amount of the population of Great Britain.
‘What I am a petitioner for is the benefit of your judgment and
authority upon the three several subjects; by reference, if there
be any other person’s calculation that you are satisfied with;
otherwise from your own notes; and, in either case, a word or
two just to indicate the sources from which they are taken would
be an additional help and satisfaction.
‘The occasion of the trouble I am attempting to give you I
expressly forbear mentioning; not only for want of space and
time, but more particularly that it may be impossible, and might,
upon occasion, be known to be impossible, that the response of
the Oracle should have received any bias from the consideration
of the purpose for which it was consulted.
‘I am, dear sir, with never failing esteem and regard,
‘Yours ever,
‘Jeremy Bentham.’
‘Q.I.P.: Sept. 30, 1794.

‘Dear Sir,—A thousand thanks for your kind letter—sorry you

should fancy you have been bathing[161] for health—hope it was
not true—only idleness—we can’t afford to have you otherwise
than well.
‘Must prefer[162] you once more, “Rental of England twenty-four
millions.” Good! but houses, such as those in town, and others
that have a separate rent, are included? I suppose not; since for
them you would have given a separate and different price in
number of years’ purchase.
‘In one of your tours you guess this article at five millions. Do
you abide by that guess? I think the number must have
increased since then considerably; that was, I believe, about
twenty years ago. London and the environs must since then
have increased, I should think, at least a quarter of a million.
How many years’ purchase would you reckon houses at, upon
an average, old and young together? Shall we say sixteen? I
should think, at the outside.
‘I am, dear Sir,
‘Your much obliged,
‘Jeremy Bentham.
‘A. Young, Esq.’
 The two following letters are from Mrs. Hoole, Dr. Burney’s
favourite, Miss Bessy, to her father:—
To Arthur Young, Esq.

‘Sidmouth: Feb. 22, 1794.

‘Dear Sir,—We came hither from Lynn near three weeks since,
as Mr. Hoole informed you. We are in very warm and comfortable
lodgings, and the woman of the house is very attentive and
obliging. The air of this place is very mild and very moist, but
they tell us the healthiest of any upon the coast. Mr. Hoole has
been on to Exmouth, which, upon the whole, he does not like so
well. We do not find that Devonshire is cheaper the further you
go, but the contrary, at least on the coast.
‘With regard to myself, I do not find I am any better for this
journey, indeed I have had more fever and cough since I came
here than ever I had. I am at present better, but I know that is
owing to a very strict regimen which I have lately taken to. The
weather has been very unfavourable, for though it has not been
cold we have had almost constantly either rain or wind. We have
been absent from home near nine weeks, and Mr. H. must very
soon return to his curacy; he will either take me with him or leave
me here, and we wish very much to know what you advise,
considering all circumstances.
‘This place is certainly warmer than Surrey, but we have heard
here, as at Lynn, that it sometimes proves unfavourable in
consumptive cases. I do not think Abinger at all in fault; I have
been well or better there than anywhere. But I am not unwilling to
be left here, if it should still be thought advisable. Will you have
the goodness to write as soon as you can, as we shall not
determine till we hear? Mr. Hoole has had but one letter from you
about a month ago. This I mention lest you should have sent any
which may have miscarried.
‘Believe me, dear Sir,
‘Your affectionate Daughter,
‘Elizabeth Hoole.
 ‘Perhaps you may like to know something of the price of
provisions: Meat 4½d. per lb.; poultry is reasonable; chickens
from 2s. to 2s. 6d. a couple; milk 2d. a quart; butter 10d. per lb.’

Postscript from Mr. Hoole

‘I fear this journey will be of no avail. I do not think our dear

Bessy is in any immediate danger, but I much fear this cruel
disease is gradually preying on her strength.
‘S. H.’

‘Sidmouth: March 18, 1794.

‘My dear Sir,—I was very sorry to find from your letter that
what I had written had made you uneasy; I am certain you think
me worse than I am; indeed it is very foolish to write my
symptoms to my friends, as they give way perhaps, or some of
them, in a short time, as is my case. I am now quite free from
pain, and can sleep on one side as well as the other; I think the
last blister was of use. I have been twice in the warm bath since
Mr. Hoole went. My cough must have its course.
‘I had a very kind letter from Agnes yesterday; she offers, if
she can get permission, to come and stay with me until Mr.
Hoole returns, and adds, if she cannot, Mrs. Forbes says she is
at liberty, and would willingly come; but I would not bring them
down upon any account, as I am more comfortably settled than
anybody would suppose, and I am sure Mr. Hoole will be back in
a short time.
‘Sidmouth is certainly very mild; we have had no cold winds,
but this clear weather suits me better than that warm moist
weather we had in February. But I cannot walk by the seaside;
there is always wind, and it seems colder than anywhere else.
‘I would not blame Mrs. F. in the least; I might have been the
same or worse anywhere; if anything in the air disagreed with
me, it was the moisture. We have no post from hence, either
Monday or Tuesday. I wrote to Mr. Hoole last Sunday, or would
have answered yours sooner. The quickness, or rather rapidity,
with which our letters arrive from town, seems surprising—a
letter put in one night we have the next. It is not the custom
indeed to deliver them at night, as the post comes in so late as
nine, but if you send they will give you them. At Lynn, which is
about the same distance from town, they deliver them at six in
the evening, but we have here a cross-post to send for them nine
miles.
‘I beg you will not think me worse than I am, and believe me,
‘Your affectionate Daughter,
‘Elizabeth Hoole.’

From J. Symonds, Esq.

‘Cambridge: March 27, 1795.

‘My dear Sir,—I am to thank you for two letters, which should
not have lain unanswered if a retirement like mine would have
furnished me with any materials. However, I must take notice of
your way of arguing. You say “the people in France are starved,
and assignats are destroyed,” with significant dashes. You told
me just the same in 1793 and 1794, and venture it once more.
Assuredly you seem to reason like the old wizard Tiresias in
Horace, “Quicquid dicam aut erit aut non.” Whether your
predictions be verified or not, you assume, like Tiresias, to speak
the truth.
‘I always thought with you, that Mr. Pitt would receive no real
benefit from his new friends; but I have heard the Duke of
Grafton say that he would not have entered on the war if he had
not been able to detach some from the Opposition. If this be so,
there is great reason to lament that he could detach them.
‘We have received here the Bishop of Llandaff’s speech on the
Duke of Bedford’s motion, published by Debrett. It amazed me to
find that the Bishop of Durham ventured to speak after him. A
gentleman who heard them both says that Watson’s was rich,
clouted cream, and Barrington’s thin, meagre, blue skim milk,
frothed up with an egg, but with so weak a froth that it rose only
to fall instantly. We are told that after Æschines was banished, in
consequence of Demosthenes’ speech de coronâ, one of
Æschines’ friends carried to him in his banishment a copy of
Demosthenes’ speech; on which the former said, “But what if you
had heard it?”
‘Two fellows of this college, who heard Watson, bear the same
ample testimony to the excellent manner in which he delivered it.
‘You tell me “that our situation is prosperous beyond all
example;” I should think so too if it were unnecessary to multiply
loans. The complaints of the dearness of the necessaries of life
seem to pervade the whole island, and I fear they must still be
dearer. If we be forced to persist in this war (and how are we to
get out of it, it is difficult to see) the middle class of the people, of
which you and I form a part, must be driven down to the lower.
They hold it is a principle not to tax the lower, but to tax luxuries,
so that the middle class will be forced to abandon everything but
necessaries, and then the upper class must pay all. This, to use
your words, “must render us prosperous beyond all example.” I
rather accede to Charles Coles’ declaration in his last letter to
me: “Alas! our glory is gone to decay.” A day or two ago I was
looking into the famous pamphlet of my old friend, Israel
Mauduit,[163] on the German war, in which I stumbled on the
following sentence, very applicable to our entering into this just
war to save the Dutch: “Is Britain to make itself the general
knight errant of Europe, to rescue oppressed States, and
exhaust itself in order to save men in spite of themselves, who
will not do anything towards their own deliverance?” Adieu!
‘Yours sincerely,
‘J. Symonds.’

1796.—In the spring of this year I waited on Mr. Pitt, by his

appointment, in order to answer some enquiries of his relative to
the propriety of any regulations by Parliament of the price of
labour.
I answered all his enquiries, and could not but admire the
wonderful quickness of his apprehension of all those collateral
difficulties which I started, and of which he seemed in a moment
to comprehend the full extent. I found him hostile to the idea.
March.—Among various dinners [was] at Mr. Burke’s and at
Mrs. Barrington’s parties. In May dinners at Duke of Bedford’s,
Duke of Buccleugh’s, and Mr. Jenkinson and Lady Louisa’s; her
manner is not the most agreeable, [but] she has ease and
elegance. I have long known her at Ickworth.
May 1.—For some time past the following advertisement has
appeared in many of the London papers: ‘Speedily will be
published a letter from the Bight Hon. Edmund Burke to Arthur
Young, Esq., Secretary to the Board of Agriculture, on some
projects talked of in Parliament, for regulating the price of labour.’
The appearance of this advertisement induced Sir John Sinclair
to write to Mr. Burke to propose to him that he should undertake
to draw up for the Board the chapter of a general Report which
was intended to treat on the subject of labour and provisions.
The question in the House of Commons was decided before
the publication could appear, and it was supposed that Mr. Burke
had, in consequence, abandoned the intention of publishing his
ideas. But Sir John, not having received any answer, or, at least,
any that was satisfactory to him, requested me to take his chariot
and go to Gregory’s, in order that I might discover whether that
celebrated character continued his intention of throwing his
thoughts upon paper.
I reached Mr. Burke’s before breakfast, and had every reason
to be pleased with my reception.
‘Why, Mr. Young, it is many years since I saw you, and, to the
best of my recollection, you have not suffered the smallest
change; you look as young as you did sixteen years ago. You
must be very strong; you have no belly; your form shows
lightness; you have an elastic mind.’
I wished to myself that I could have returned anything like the
compliment, but I was shocked to see him so broken, so low, and
with such expressions of melancholy. I almost thought that I was
come to see the greatest genius of the age in ruin.
And I had every reason to think, from all that passed on this
visit, that the powers of his mind had suffered considerably.
He introduced me to his brother, Mr. W. Burke, to Mrs. B., and
to the Count de la Tour du Pin, an emigrant philosopher and
naturalist.
 After breakfast he took me a sauntering walk for five hours
over his farm, and to a cottage where a scrap of land had been
stolen from the waste. I was glad to find his farm in good order,
and doubly so to hear him remark that it was his only
amusement, except the attention which he paid to a school in the
vicinity for sixty children of noble emigrants. His conversation
was remarkably desultory, a broken mixture of agricultural
observations, French madness, price of provisions, the death of
his son, the absurdity of regulating labour, the mischief of our
Poor-laws, and the difficulty of cottagers keeping cows. An
argumentative discussion of any opinion seemed to distress him,
and I, therefore, avoided it. And his discourse was so scattered
and interrupted by varying ideas, that I could bring away but few
of his remarks that were clearly defined.
Speaking on public affairs he said that he never looked at a
newspaper; ‘but if anything happens to occur which they think
will please me, I am told of it.’ I observed there was strength of
mind in this resolution. ‘Oh, no!’ he replied, ‘it is mere weakness
of mind.’ It appeared evident that he would not publish upon the
subject which brought me to Gregory’s; but he declared himself
to be absolutely inimical to any regulation whatever by law; that
all such interference was not only unnecessary but would be
mischievous. He observed that the supposed scarcity was
extremely ill understood, and that the consumption of the people
was a clear proof of it; this, in his neighbourhood, was not
lessened in the material articles of bread, meat, and beer, which
he learnt by a very careful examination of many bakers,
butchers, and excisemen; nor had the poor been distressed
further than what resulted immediately from that improvidence
which was occasioned by the Poor-laws.
Mr. Burke had not read Lord Sheffield’s Memoirs of Gibbon.
On my observing that Mr. Gibbon declares himself of the same
opinion with him on the French Revolution, he said that Gibbon
was an old friend of his, and he knew well that before he (Mr. G.)
died, that he heartily repented of the anti-religious part of his
work for contributing to free mankind from all restraint on their
vices and profligacy, and thereby aiding so much the spirit which
produced the horrors that blackened the most detestable of all
revolutions.
Upon my mentioning Monsieur de Mounier and Lally Tollendal,
he exclaimed, ‘I wish they were both hanged!’
He seemed to bear hard upon the Duke de Liancourt, and to
allude indistinctly to some report of my having opened an
hospitable door to that nobleman, and having received a bad
return. I defended the duke, and had not the conversation been
interrupted I should have discovered what he meant by the
remark. The same observation has met my ear on other
occasions, but was never explained.
Mrs. Crewe arrived just before dinner, and though she exerted
herself with that brilliancy of imagination which renders her
conversation so interesting, it was not sufficient to raise the
drooping spirits of Mr. Burke; it hurt me to see the languid
manner in which he lounged rather than sat at table, his dress
entirely neglected, and his manner quite dejected; yet he tried
once or twice to rally, and once even to pun. Mrs. Crewe,
observing that Thelwel was to stand for Norwich, said it would be
horrid for Mr. Wyndham to be turned out by such a man. ‘Aye,’
he replied, ‘that would not tell well.’
She laughed at him in the style of condemning a bad pun.
Somebody said it was a fair one, he said it was neither very bad
nor good.
He gave more attention to her account of Charles Fox than to
any other part of her conversation. She spoke slightingly of him,
and gave us some account of his life at Mrs. Armstead’s. She
says he lives very little in the world, or in any general society, for
years past; that his pleasure is to be at the head of a little society
of ten or twelve toad eaters, and seems to contract his mind to
such a situation.
The conversation would have become more interesting had
not Mrs. Crewe been so full of a plan for Ladies’ Subscriptions
for the Emigrants, and consulting him so much on the means of
securing the money from the fangs of the Bishop of St. Pol de
Léon, whose part, however, Mr. Burke took steadily. This
business was so discussed as to preclude much other