GLOBAL
EDITION
Brock Biology of
Microorganisms
FIFTEENTH EDITION
Madigan • Bender • Buckley • Sattley • Stahl
Brief Contents
UNIT 1 CHAPTER
CHAPTER
1
2
The Microbial World 37
Microbial Cell Structure and Function 70
The Foundations of CHAPTER 3 Microbial Metabolism 109
Microbiology CHAPTER 4 Molecular Information Flow and Protein Processing 138
UNIT 2 CHAPTER
CHAPTER
5
6
Microbial Growth and Its Control 173
Microbial Regulatory Systems 209
Microbial Growth CHAPTER 7 Molecular Biology of Microbial Growth 238
and Regulation CHAPTER 8 Viruses and Their Replication 259
UNIT 3 CHAPTER
CHAPTER
9
10
Microbial Systems Biology 277
Viral Genomics, Diversity, and Ecology 310
Genomics and Genetics CHAPTER 11 Genetics of Bacteria and Archaea 342
CHAPTER 12 Biotechnology and Synthetic Biology 368
UNIT 4 CHAPTER
CHAPTER
13
14
Microbial Evolution and Systematics 399
Metabolic Diversity of Microorganisms 428
Microbial Evolution CHAPTER 15 Functional Diversity of Microorganisms 487
and Diversity CHAPTER 16 Diversity of Bacteria 530
CHAPTER 17 Diversity of Archaea 566
CHAPTER 18 Diversity of Microbial Eukarya 593
UNIT 5 CHAPTER
CHAPTER
19
20
Taking the Measure of Microbial Systems 619
Microbial Ecosystems 651
Microbial Ecology CHAPTER 21 Nutrient Cycles 687
and Environmental CHAPTER 22 Microbiology of the Built Environment 708
Microbiology CHAPTER 23 Microbial Symbioses with Microbes, Plants, and Animals 732
UNIT 6 CHAPTER
CHAPTER
24
25
Microbial Symbioses with Humans 765
Microbial Infection and Pathogenesis 793
Microbe–Human CHAPTER 26 811
Interactions and the CHAPTER 27 834
Immune System CHAPTER 28 Clinical Microbiology and Immunology 866
Unit 7 CHAPTER
CHAPTER
29
30
Epidemiology 902
Person-to-Person Bacterial and Viral Diseases 923
Infectious Diseases and CHAPTER 31 Vectorborne and Soilborne Bacterial and Viral Diseases 955
Their Transmission CHAPTER 32 Waterborne and Foodborne Bacterial and Viral Diseases 973
CHAPTER 33 Eukaryotic Pathogens: Fungi, Protozoa, and Helminths 994
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Where cutting edge science meets
state of the art learning.
The Fifteenth Edition of the world-renowned Brock Biology of Microorganisms
introduces today’s students to cutting edge microbiology research and ensures
core concept mastery, enhanced by .
NeW & Revised! Microbiology
5
Now chapter opening features
highlight cutting edge, engaging
Microbial Growth
and Its Control research that is important to how we
understand microbiology today. Paired
assessments in MasteringMicrobiology
microbiologynow
Picking Apart a Microbial Consortium
engage students in the course material
In nature, certain metabolic processes are carried out by microbes and foster deep concept mastery.
that team up to get the job done, a cozy arrangement called a con-
sortium. Such is the case with the oxidation of methane (CH4) linked
to the reduction of sulfate (SO42-) in anoxic marine sediments. The
overall reaction (CH4 + SO42- S HCO3- + HS- + H2O) is exergonic and
the small amount of energy released is shared between two distinct
microbes. The methane oxidizer in the consortium is a species of
Archaea nicknamed ANME (for anaerobic methanotroph, blue in
photo), and its sulfate-reducing partner is a species of Bacteria
(brown in photo). The consortium is thought to play a key role in the
carbon cycle as a major methane sink, and thus a detailed picture of
how it works is important to our understanding of the global carbon
economy, climate change, and marine biogeochemistry.
Researchers have tried for years to separate the consortium into
its components but always found that methane oxidation required
both organisms. However, some researchers hypothesized that it
might be possible to replace the sulfate reducer with an artificial
electron acceptor and that this might unlock the consortium and
allow the methanotroph to grow in pure culture. Using an electron
acceptor called AQDS, the scientists discovered that they could turn
off sulfate reduction in the consortium while maintaining CH4 oxi-
dation. During this process, the methanotroph used electrons from
CH4 to reduce AQDS rather than passing them on to its sulfate-
reducing partner. Several other electron acceptors known to support
anaerobic respiration also sustained methane oxidation, giving hope
that ANME may eventually be obtained in pure culture. I Cell Division and Population Growth 174
The ability to grow a microbe in pure culture is the “gold standard” for the II Culturing Microbes and Measuring
study of its physiology, biochemistry, regulation, and several other aspects of its Their Growth 180
25
biology. In the case of the ANME–sulfate reducer consortium, several physiologies
III Environmental Effects on Growth:
were active at once, and resolving these many reactions proved to be a major
Temperature 188
scientific challenge. However, if further work shows that ANME can be removed
from the consortium and grown in pure culture, detailed aspects of its biology IV Environmental Effects on Growth:
pH, Osmolarity, and Oxygen 194
Microbial Infection
can be studied that were not possible when the organism was tightly coupled to
its partner in the consortium (photo). V Controlling Microbial Growth 200
Source: Scheller, S., H. Yu, G.L. Chadwick, S.E. McGlynn, and V.J. Orphan. 2016.
Artificial electron acceptors decouple archaeal methane oxidation from sulfate reduction.
and Pathogenesis
Science 351: 703–706.
173
microbiologynow
The Microbial Community That Thrives on Your Teeth
Few people have such superb oral hygiene that they lack dental
plaque, the microbial biofilm that forms on and between teeth and
along or below the gumline. If not removed regularly, dental plaque
invariably leads to dental caries (cavities), the condition in which por-
tions of tooth enamel and dentin break down from the onslaught of
bacterial activities. Dental plaque and dental caries develop from the
natural tendency of oral bacteria such as Streptococcus mutans and its
close relative S. sobrinus to attach firmly to the teeth and gums and
ferment sucrose (table sugar) to lactic acid, which attacks the teeth
and slowly rots them away.
Until recently, dental plaque was thought to consist largely of the
aforementioned streptococci. Both species could easily be isolated
from dental plaque and both light and electron microscopy typically
showed large numbers of cocci in chains, a hallmark of the genus
Streptococcus. But a recent molecular ecology study of the microbial
diversity of dental plaque revealed that this material is composed of
more than just streptococci and develops in a precisely structured way.
The photo here is a light micrograph of a section through human
dental plaque stained by fluorescence in situ hybridization (FISH). I Human–Microbial Interactions 794
Different oligonucleotides, each specific for a different major phylum II Enzymes and Toxins of Pathogenesis 800
of Bacteria and containing a distinct fluorescent dye, were allowed to
hybridize to the ribosomal RNA in cells in the plaque and then
observed by fluorescence microscopy. Surprisingly, instead of seeing
primarily streptococci, the researchers saw a diverse and highly organized
microbial community. The micrograph shows streptococci (stained green)
located primarily at the periphery of the plaque beyond several other bacteria
that combine to form a scaffold emerging from the tooth surface. These
include Corynebacterium (purple), Capnocytophaga (red), Fusobacterium
(yellow), Leptotrichia (blue-green), and Haemophilus (orange), among others.
A major conclusion that emerged from this study was that the scaffolding
microbes likely function to position the streptococci out into the oral cavity
where sucrose should be more available.
New views of old problems often reveal surprising results. In the case of
dental plaque, FISH technology has revealed a whole new microbial world in a
habitat previously thought to be dominated by only two species of well-
characterized bacteria.
Source: Mark Welch, J.L., et al. 2016. Biogeography of a human oral microbiome at the
micron scale. Proc. Natl. Acad. Sci. (USA) 113: doi: 10.1073/pnas.1522149113.
793
Microbiology today and tomorrow.
Genomics, and the various “omics” it has spawned, support content throughout the
Fifteenth Edition ensuring that today’s students understand the transformation that
fast paced nature of the science.
CONTENTS 3
Authoritative. Accurate. Accessible.
The Fifteenth Edition continues its legacy of authoritative, accessible writing; beautiful
and clear art; and student-focused pedagogy, engaging learners in the science.
Student focused pedagogy informs the
organization and design of each chapter feature
Chapter Review
I fundamentals of Host Defense Q What is the origin of the phagocytes and
lymphocytes active in the immune response? Track the
26.1 Innate immunity is an inborn protective response to maturation of B cells and T cells.
infection characterized in part by recognition and
elimination of common pathogens, primarily through the
III Phagocyte Response Mechanisms
activity of phagocytes. Adaptive immunity is the acquired
ability of the immune system to eliminate specific 26.5 Pathogens may colonize host tissues when appropriate
pathogens from the body via lymphocyte-mediated nutrients and growth conditions are present, such as on
responses, including the production of antibodies that mucosal surfaces, especially where the composition of the
bind foreign antigens on pathogens or their products. normal microbiota has been altered. Innate responses to
Q List two different types of phagocytes. How do T microbial invasion and tissue damage are initiated by the
cells and B cells differ in their functions? from where in release of chemokines, which recruit phagocytes and
the human body do all of these cells originate and which other immune cells to sites of infection.
require maturation before they are functional? Q Describe a scenario in which microorganisms invade
26.2 The human body possesses numerous protective defenses body tissues. What factors allow for the migration of
against infectious agents. Natural host resistance to phagocytes to sites of infection?
infection includes physical barriers to infection posed by the 26.6 Innate recognition of common pathogens occurs through
skin and mucosa, as well as chemical barriers to infection pathogen-associated molecular patterns (PAMPs).
including acidic secretions, defensins, and lysozyme. The Phagocytes recognize PAMPs through preformed pattern
specificity of pathogens for particular tissues limits which recognition receptors (PRRs). The recognition and
hosts and tissues might be susceptible to infection. interaction process stimulates phagocytes to destroy the
Q How does the human normal microbiota play a role pathogens through a signal transduction mechanism that
in preventing disease? induces phagocytosis of the infectious agent.
Q Identify some PAMPs that are recognized by PRRs.
II Cells and organs of the Immune System Which cells express PRRs? How do PRRs associate with
PAMPs to promote innate immunity?
26.3 Cells involved in innate and adaptive immunity originate
from hematopoietic stem cells in bone marrow. The blood 26.7 Phagocytosis is the engulfing of infectious particles by
and lymph systems circulate cells and proteins that are phagocytes. Engulfed pathogens are bathed in toxic
important components of the immune response. Diverse oxygen compounds inside the phagolysosome, killing
leukocytes participate in immune responses in all parts of and degrading them. However, some pathogens have
the body. developed various defense mechanisms to avoid or inhibit
phagocytes, including secretion of leukocidins, the
Q Describe the significance of bone marrow, blood, and
presence of a capsule, and biosynthesis of carotenoid
lymph to cells and proteins associated with the immune
pigments, which combat oxidative stress.
system.
Q explain how phagocytes kill microorganisms, with
26.4 Leukocytes are differentiated white blood cells derived from
Continuous Learning
Before, during, and After Class
MasteringMicrobiology improves results by engaging students before, during,
and after class.
BefoRe CLAss
Reading Questions, art-based activities and
MCAT Prep, along with Quantitative Questions,
prepare students for in-depth class discussion.
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Microbial Symbioses
with Humans
microbiologynow
Frozen in Time: The Iceman Microbiome
Humans and their microbial associates—collectively called the human
microbiome—have coevolved for millennia. As we will see in this chapter,
24
the human microbiome influences a person’s health, disease, and predisposi-
tion to disease. Among our intimate microbial associates, the pathogenic
bacterium Helicobacter pylori is known to have developed a close relation-
ship with humans in the distant past and to have coevolved with humans.
H. pylori colonizes the stomachs of about half the human race. Although
this bacterium generally does not cause overt disease, it is a major risk factor
for the development of ulcers and stomach cancer. Moreover, because H.
pylori is transmitted primarily by contact within families, the distribution of
genetic variants of this bacterium may yield clues to past human migrations.
Unraveling the details of the H. pylori ancestry is complicated by the
ability of different strains of this bacterium to recombine their genetic
information. Because the DNA of various strains has mixed over long peri-
ods, the reconstruction of population movement inferred from genome
sequences of modern H. pylori strains is incomplete. One of the biggest
unanswered questions was the origin of strains now common among mod-
ern Europeans, which appear to be hybrids of strains originating in Asia
and Africa. Unfortunately, the sequence data did not point to a reliable
time interval in which that mingling of human populations occurred—an
important period of human migration that was estimated to have occurred
10,000–50,000 years ago.
This estimate has now been greatly refined following the remarkable
discovery of a well-preserved 5300-year-old European Copper Age mummy
frozen in the Italian Alps. Using the newest methods for DNA sequencing, it
was possible to reconstruct the genome of H. pylori preserved in the stom-
ach of the “Iceman” (see photo), the corpse discovered when melting ice
revealed the human remains on the side of a mountain. The Iceman H. pylori
genome sequence turned out to be an almost pure representative of the
Asian population, which means this H. pylori strain was present in Europe I Structure and Function of the Healthy Adult
before hybridization of African and Asian strains produced the modern Human Microbiome 766
European variant. Thus, by employing historical biogeography, we now
II From Birth to Death: Development of the
know this important period of human migration was much more recent
Human Microbiome 780
than previously thought.
III Disorders Attributed to the Human
Source: Maixner, F., et al. 2016. The 5300-year-old Helicobacter pylori genome of the Microbiome 783
Iceman. Science 351: 162–165.
IV Modulation of the Human Microbiome 788
765
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Microbiology is a dynamic
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