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Denise G. Anderson
U NIV ERSITY OF WAS H I N GTON
Sarah N. Salm
BOROU GH OF MAN H AT TA N
COMMU NITY COLLE GE
Mira Beins
U NIV ERSITY OF WAS H I N GTON
Eugene W. Nester
U NIV ERSITY OF WAS H I N GTON
Tenth Edition
Published by McGraw Hill LLC, 1325 Avenue of the Americas, New York, NY 10121. Copyright © 2022
by McGraw Hill LLC. All rights reserved. Printed in the United States of America. Previous editions
© 2019, 2016, and 2012. No part of this publication may be reproduced or distributed in any form or by any
means, or stored in a database or retrieval system, without the prior written consent of McGraw Hill LLC,
including, but not limited to, in any network or other electronic storage or transmission, or broadcast for
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Some ancillaries, including electronic and print components, may not be available to customers outside the
United States.
1 2 3 4 5 6 7 8 9 LWI 23 22 21
All credits appearing on page or at the end of the book are considered to be an extension of the copyright
page.
The Internet addresses listed in the text were accurate at the time of publication. The inclusion of a website
does not indicate an endorsement by the authors or McGraw Hill LLC, and McGraw Hill LLC does not
guarantee the accuracy of the information presented at these sites.
mheducation.com/highered
PA RT I I I
Microorganisms and Humans
14. The Innate Immune Response 360
15. The Adaptive Immune Response 385
16. Host-Microbe Interactions 415
17. Applications of Immune Responses 439
18. Immunological Disorders 464
19. Epidemiology 481
20. Antimicrobial Medications 504
iii
Richard Moore
niques, and medical mycology/parasitology Deborah Allen
laboratory for over 30 years. Equipped with
Deborah Allen is a Professor at Jefferson
a diverse educational background, includ-
College in Missouri, where she teaches
ing undergraduate work in nutrition and graduate work in food sci-
microbiology as well as several other
ence and in microbiology, she first discovered a passion for teaching
courses for students entering allied health
when she taught microbiology laboratory courses as part of her
careers. Her graduate work was in zoology
graduate training. Her enthusiastic teaching style, fueled by regu-
at the University of Oklahoma and in neuro-
lar doses of Seattle’s famous coffee, received high reviews from her
biology and behavior at Cornell U niversity.
students. Mike Bohrer
She participated in cancer research at the
Denise now relaxes in the Yorkshire Dales of England, where
University of Arkansas Medical Center before embarking on a career
she lives with her husband, Richard Moore. When not editing text-
in publishing, working in acquisitions and development for books in
book chapters, she can usually be found walking scenic footpaths,
the life sciences. She is now thrilled to be working on the other end of
chatting with friends, fighting weeds in her garden, or enjoying a
the desk with the Nester team. Away from campus, Deborah reads or
fermented beverage at the local pub.
listens to her favorite Eve Dallas novels, floats the rivers and listens
to folk music in the Ozarks, and fully appreciates the local microbes
while visiting Missouri wineries.
Sarah Salm
Sarah Salm is a Professor at the Borough of
Manhattan Community College (BMCC)
Eugene Nester
of the City University of New York, Gene (Eugene) Nester was instrumental in
where she teaches microbiology, anatomy establishing the text’s reputation for excel-
and physiology, and general biology. She lence over the decades. Although no lon-
earned her undergraduate and doctoral ger an active member of the author team,
degrees at the University of the Witwa- he wrote the original version of the present
Sandy Coetzee
tersrand in Johannesburg, South Africa. text with Evans Roberts and Nancy Pearsall
She later moved to New York, where she did postdoctoral work at more than 30 years ago. That text, Microbi-
Courtesy Eugene Nester
the NYU School of Medicine. Her research background is diverse ology: Molecules, Microbes and Man, pio-
and includes plant virology, prostate cancer, and bacteria in con- neered the organ system approach to the study of infectious disease
taminated water sources. and was developed specifically for allied health sciences.
iv
Top: Jenner Images/Getty Images, Left: Hero Images/Getty Images, Right: Hero Images/Getty Images
Development
Dev
velopmen
nt
Hematopoietic stem cell
Immature B cells: As
these develop, a
functionally limitless
assortment of B-cell
receptors is randomly Antigen X
generated.
Effector action
Antibodies:
These neutralize the Lymphoid organ
Plasma cells invader and tag it for
destruction.
(effector B cells):
These descendants of
activated B cells
secrete large quantities
of antibody molecules MHC class I molecule
that bind to antigen X. Dendritic cells presenting Dendritic cells presenting “self”
Co-stimulatory molecule microbial peptides produce peptides or other harmless
Memory B cells: co-stimulatory molecules. material do not produce
These long-lived MHC class II molecule co-stimulatory molecules.
descendants of
activated B cells
recognize antigen X
when it is encountered
again. T-cell receptor T-cell receptor T-cell receptor T-cell receptor
Effector action
CD4 CD8 CD4 CD8
Antibodies:
These neutralize the
invader and tag it for
destruction.
FIGURE 15.10
by dendritic cells expressing co-stimulatory molecules not expressing co-stimulatory molecules become anergic or, in
can become activated. the case of CD4+ cells, may become regulatory T cells.
(a)
(a
a))
a
Targeted delivery
of a “death package”
Virally infected “self” cells TC cell recognizes viral peptide Target cell undergoes apoptosis.
present viral peptides on presented by an infected “self” cell
MHC class I molecules. and initiates apoptosis in that target.
It also releases cytokines that alert
neighboring cells.
(b)
1 2 3 CD4
Cytokine delivery T-cell receptor
FIGURE 15.14
Secretion
of cytokines
B
B-cell receptor binds B cell internalizes antigen. B cell degrades antigen into
to antigen.
to peptide fragments.
5a T-cell receptor
FIGURE 15.16 4
Antigen
fragment
MHC class II
molecule TH cell recognizes
antigen fragment
Microbial and activates B cell.
antigen
Cytokine delivery
Harmless 5b
antigen
viii
Walk through the processes a characteristic set of signs and symptoms—such as fever,
fatigue, and headache—that result from the systemic immune
response to the infecting agent. 1 The microorganism is present in every case of the disease.
Step-by-step figures direct the student using numbered icons, The suffix -emia means “in the blood.” Thus, bacteremia
indicates that bacteria are circulating in the bloodstream. Note
that this term does not necessarily imply a disease state. A
often with corresponding icons in the text. person can become bacteremic for a short period of time after
forceful tooth brushing. On the other hand, infection-induced
bacteremia can lead to a life-threatening systemic inflamma-
tory response, a condition called sepsis. Toxemia indicates that
2 The microorganism must be grown in pure culture from diseased hosts.
toxins are circulating in the bloodstream. The organism that
causes tetanus, for instance, produces a localized infection, yet
its toxins circulate in the bloodstream. The term viremia indi-
178 Chapter 7 The Blueprint of Life, from DNA to Protein cates that viral particles are circulating in the bloodstream.
FIGURE 7.3 Three Functional Types Protein-encoding gene rRNA gene tRNA gene MicroAssessment 16.3
of RNA Molecules The different
functional types of RNA required for gene
A primary pathogen can cause disease in an otherwise healthy
individual; an opportunist causes disease in an immunocompromised
“The text and illustrations are ‘tight’ and give each other
expression—messenger RNA (mRNA), DNA Transcription
host. The course of infectious disease includes an incubation period,
ribosomal RNA (rRNA), and transfer RNA
illness, and a period of convalescence. Infections can be acute,
(tRNA)—are transcribed from different
good support.”
chronic, or latent; they can be localized or systemic. 3 The same disease can be produced when a pure culture of the
genes. The mRNA is translated, and the microorganism is introduced into susceptible hosts.
Messenger RNA (mRNA) Ribosomal RNA (rRNA) Transfer RNA (tRNA)
tRNA and rRNA fold into characteristic 5. Why are diseases caused by opportunists becoming more
frequent?
—Richard Shipee, Vincennes University
three-dimensional structures that each play
Translation
a role in protein synthesis. 6. Give an example of a microbe that causes a latent infection.
? Ribosomal RNA is a component of ribosomes. 7. What factors might contribute to a long incubation period?
What are ribosomes? Protein
(see figure 2.13). Another distinction is that RNA contains proteins; instead the RNA molecules themselves are the final 16.4 ■ Determining the Cause
the nucleobase uracil in place of the thymine found in DNA products, and each type plays a different but critical role in of an Infectious Disease
(see figure 2.28). Also, RNA is usually a single-stranded linear protein synthesis. 4 The same microorganism must be recovered from the experimentally
molecule much shorter than DNA. Learning Outcome infected hosts.
RNA is synthesized using a region of one of the two Regulating Gene Expression 6. List Koch’s postulates, and compare them to the molecular
strands of DNA as a template. In making the RNA molecule, Although a cell’s DNA can encode thousands of different pro- Koch’s postulates. FIGURE 16.3 Koch’s Postulates These criteria provide a foundation
or transcript, the base-pairing rules apply except that uracil, teins, not all of them are needed at the same time or in equal for establishing that a given microbe causes a specific disease.
rather than thymine, pairs with adenine. The interaction of quantities. Because of this, cells require mechanisms to regu- Criteria are needed to guide scientists as they try to determine
? To fufill Koch’s postulates, why must an organism suspected of causing the
DNA and RNA is only temporary, however, and the transcript the cause of an infectious disease. They can also be helpful
late the expression of certain genes. A fundamental aspect of disease be able to grow in laboratory medium?
when studying the disease process.
quickly separates from the template. the regulation is the cell’s ability to quickly destroy mRNA.
Three different functional types of RNA are required Within minutes of being produced, transcripts are degraded
for gene expression, and these are transcribed from different by cellular enzymes. Although this might seem wasteful, it Koch’s Postulates 1 The microorganism is present in every case of the disease.
sets of genes (figure 7.3). Most genes encode proteins and actually provides cells with a means to control gene expres- The steps that Robert Koch used to show that Bacillus 2 The microorganism must be grown in pure culture from
are transcribed into messenger RNA (mRNA). The infor- sion. If transcription of a gene is turned “on,” transcripts will anthracis causes anthrax (see A Glimpse of History) are diseased hosts.
mation encrypted in mRNA is decoded according to the be available for translation; if it is then turned “off,” the num- now known as Koch’s postulates. Although they were never 3 The same disease can be produced when a pure culture
genetic code, which correlates each set of three nucleotides ber of transcripts will rapidly decline. By simply regulating meant to be applied rigidly, they still provide scientists with of the microorganism is introduced into susceptible hosts.
to a particular amino acid. The genes for ribosomal RNA the synthesis of mRNA molecules, a cell can quickly change a logical framework for establishing that a given microbe 4 The microorganism must be recovered from the experi-
(rRNA) and transfer RNA (tRNA) are never translated into the levels of protein production (figure 7.4). causes a certain infectious disease (figure 16.3): mentally infected hosts.
Translation of each of
Encourage deeper understanding
24.1 ■ Anatomy, Physiology, by first physically breaking down food into small particles,
then chemically degrading those particles even further, and
the gene A transcripts and Ecology of the finally absorbing the available nutrients. The waste material
generates some protein A.
Figures Digestive
have accompanying
System questions that encourage that remains is eliminated as feces.
The digestive system includes two general components:
students to think more carefully about the concept illus-
Learning Outcomes
Translation of each of the gene C transcripts the digestive tract and the accessory organs (figure 24.1). The
generates many molecules of protein C. 1. Describe the functions of the main components of the upper digestive tract is a hollow tube that starts at the mouth and
and lower digestive tract.
trated in a figure.
2. Identify the functions of the liver and other accessory organs.
ends at the anus. When referring only to the stomach and the
intestines, the term gastrointestinal tract is often used. The
3. Describe the significance of the normal intestinal microbiota. accessory organs—which include the salivary glands, liver,
and pancreas—support the digestive process by producing
The main purpose of the digestive system is to convert the vital enzymes and other substances that help break down food.
food we eat into a form that the body’s cells can use as a Like the respiratory system and the skin, the digestive tract
FIGURE 7.4 The Level of Gene Expression Can Be Controlled source of energy and raw materials for growth. It does this is one of the body’s major boundaries with the environment.
? How does the fact that mRNA is quickly degraded help a cell control gene expression?
Organ Function
Each disease chapter includes a stunning figure that intro- Gallbladder Pancreas
Pancreatitis
in fat digestion
Food
molecules Large Absorbs some water
intestine and minerals;
Epithelial cells prepares waste
with microvilli Villus
FIGURE 24.1 The Digestive System Some of the disease conditions that can affect the system are shown in red.
? How do the accessory organs of the gastrointestinal tract support digestion?
ix
Key Terms for each chapter are defined on the KEY TERMS
Blood-Brain Barrier Cells Meninges Membranes covering the
opening page. that function together to create a
protective semipermeable border
brain and spinal cord.
Meningitis Inflammation of the
that separates the CNS from the meninges.
bloodstream.
Peripheral Nervous System
Central Nervous System (PNS) Division of the nervous
(CNS) Brain and spinal cord. system that carries information to
and from the CNS.
Cerebrospinal Fluid (CSF) Fluid
produced in the brain that flows Transmissible Spongiform
within and around the CNS. Encephalopathy (TSE) Chronic
degenerative brain disease caused
Encephalitis Inflammation of the by prions; characterized by spongy
A Glimpse of History opens each chapter, featur- Because the word leprosy carries centuries of grim overtones,
many people prefer to use the term Hansen’s disease, a name that
ing engaging stories about the men and women who Structure of West Nile virus particles. ©Science Picture Co/Getty Images
honors the discoverer of the causative bacterium. Today, the disease
can be treated.
sions designed to highlight the relevance of the information. was productive, meaning that it contained
sputum (pronounced spew-tum). She was
particularly concerned that the sputum was
2. What is the significance of the mucoid
phenotype of the colonies?
3. Siderophores help the bacterium obtain
iron from the host. Recall that the
body produces iron-binding proteins,
a blue-green color. His doctor immediately 3. How would the siderophore (the iron-
including lactoferrin and transferrin;
suspected a lung infection by Pseudomonas binding compound) benefit the bacterium?
this prevents microbes from using the
aeruginosa—a common complication of 4. Why would the boy’s lung infection host’s iron and thereby limits their
cystic fibrosis. A sputum sample was col- make his pre-existing respiratory growth. Microorganisms that make sid-
lected and sent to the clinical laboratory. problems even worse? erophores essentially engage in a “tug-
In the clinical laboratory, the sample was of-war” with the body over iron. This
Discussion
plated onto MacConkey agar and blood agar tug-of-war is especially important for
and incubated. Mucoid colonies surrounded 1. Cystic fibrosis patients often have P. aeruginosa because iron levels influ-
by a bluish-green color grew on both types of an accumulation of thick mucus in ence biofilm formation. When iron is
agar media. The colonies on MacConkey had their lungs, which interferes with the limiting, P. aeruginosa cells are motile
no pink coloration, so the medical technolo- mucociliary escalator and other first- and do not initiate biofilm formation.
244 gist concluded that the cells did not ferment
Chapter 9 Biotechnology line defenses. With a compromised
4. In response to a bacterial infection in the
lactose. She noted the blue-green color on the (weakened) mucociliary escalator,
Provide perspective
lungs, an inflammatory response devel-
agar plates and in the sputum, knowing that microbes that are inhaled are not eas-
F O C U S Y O U R P.Paeruginosa E R S Pmakes
E C Tseveral
I V Epigmented
9 .1com- ily removed. In addition, the accumu-
ops. The capillaries in the area become
leaky, allowing fluids from the blood to
pounds that give rise to lated mucus serves as a nutrient source
The COVID-19 Response—The Power ofcolors ranging from
Biotechnology enter the tissues. Those fluids cover the
yellow to blue. One of the pigments functions for bacteria.
Part
respiratory IV Infectious
surfaces, therebyDiseases
interfering 551
Focus Your Perspective boxes show how micro- The COVID-19 response as serves
binds iron.
lent illustration of the power
as an excel-
a siderophore, which istechnologies;
Another is important
of biotechnol-
a molecule that 2. The
not only do CRISPR-Cas-
give resultsbacterium
for biofilm
based tests in about anproduces
hour
facilitated
mucoid colonies suggest that research
an extracellu-
targeted
the aimed
with gas
antiviral therapies,
at exchange.
developing
as described
mation recruits
In addition, inflam-
neutrophils to the area.
ogy. Because ofMicroAssessment
formation. 21.3 or less, they do not compete with PCR- in Focus on the FutureSome
Further
several of the technologies testing showed that the lar material that forms a capsule or a 20.1.ofBy
the analyzing
neutrophils will die, releas-
organisms and their products influence our lives in described in this chapter,
global outcome—although
bacterium
The common
ative rod,
particularly
was an oxidase-positive,
the pandemic’s
cold may bewith
consistent
devastating—
caused
the
Gram-neg-
based tests with respect
by manyinitial
physician’s
reagents.
slime
to layer.
different viruses,
The first CRISPR-Cas-based
rhinoviruses. Transmission requires close personal
This material,
the required thealso
viralreferred
genome, scientists
7. Why aresub-
to as extracellular polymeric people most
amino acid sequence of
a for
cold?
determined
ing infectious
the
key proteins
granules
the
destructive
duringenzymes
essen-
as a result.
the first in
fewtheir
days of
suspicions. stancesfor(EPS), allowstial
Pseudomonas
many different ways.
resulted in fewer deaths than many feared diagnostic test was approved use only replication of the virus. Relatively
contact during which respiratory droplets from an infected person How is an adenovirus infectionstructure
treated?
or predicted. in certified laboratories, but researchers 8. soon thereafter, the 3-dimensional
or respiratory secretions on contaminated
SARS-CoV-2, the virus that causes also worked hands toward are transferred
developing similar 9. of Contrary
two of those proteins was determined—
to your grandmother’s warnings, going outside
to the
an nasalRNA epithelium.
genome. IfSymptoms typically last forversions about a week oneinthat
COVID-19, has a but instrument-free that can be coldtheweather
virus uses to attach
without a coat to and
is not then
likely to cause a cold.
researcher needs before a DNA the disease copy isofstopped that bycompleted
immune responses. on-site (comparable Adenovirus to home enter host cells and one it uses to replicate
Why not?
infections
genome, the enzyme reverse transcriptase may
infection, resemble
the regulatingcolds, but
center fever
“sets”
pregnancy tests). is present.
the body’s thermostat at Fever-inducing
its genome. Knowing cytokines
those and
proteinother substances are pyro-
struc-
is used to make cDNA. a higher
When level.the virus Anwas oral temperature Data obtained above 37.8°C via is regarded
high-throughput gens (pyro
tures allowsmeans “fire”toorfocus
scientists “heat” andefforts
their gen means “to gener-
first discovered in China, as fever. a cDNA copy of sequencing were used to track the global ate”). Pyrogenic cytokines
on developing compounds arethat
endogenous
specifically pyrogens, meaning
its genome was cloned and A then
higher temperaturespread
sequenced. settingofresults SARS-CoV-2.
when macrophages The tracking thetarget bodythe parts them,
makes essential for the microbial
whereas structure’s products such as
be used to prevent certain diseases. In fact, immunization has discuss the applications of immunotherapies, methods designed
probably had the greatest impact on human health of any medi- to either enhance or suppress specific immune responses as a
cal procedure, and it is just one example of how knowledge is means to treat certain diseases. Finally, we will explore some use-
power with respect to fighting disease (figure 17.1). We will also ful applications of immunological reactions in diagnostic tests.
xi
IMMUNIZATION AND IMMUNOTHERAPY
17.1 ■ Principles of Immunization antibodies produced by other people, animals, or even modi-
fied B cells growing in culture. It is used to prevent disease
Learning Outcome immediately before or after likely exposure to a pathogen or its
and35508_fm_i-xxxii.indd 11 1. Compare and contrast naturally acquired active immunity, toxins. Note that passive immunity provides no 10/14/20
memory; once 3:57 AM
the transferred antibodies are degraded, the protection is lost.
FOCUS ON UNDERSTANDING . . .
Student-Friendly Descriptions
Include analogies
WHY? Analogies provide students a comfortable framework for making
sense of difficult topics. Here’s an example from chapter 14.
Innate Immunity The innate immune system has three143general Part I Life and Death of Microorganisms Steve Cole/E+/Getty Images
components:
TABLE 6.1
first-line defenses, sensor systems, and innate effector
Electron Carriers
Reduced electron carriers represent reducing power lipids, carbohydrates, and nucleic acids. When E. coli cells Catabolism
Certain intermediates of catabolic pathways can be used in degrade glucose molecules, the pathways they use release
because they can easily transfer their electrons to another Catabolism of glucose, the preferred energy source of many
anabolic pathways; therefore they link these two types of energy and also form a dozen or so precursor metabolites.
cells, includes two key sets of processes:
WHY? Descriptions that emphasize the logic of processes make it easier for
to the electron transport chain, which then uses the energy
to generate a proton motive force. In turn, this drives the
metabolite pyruvate can be converted to any one of three
amino acids: alanine, leucine, or valine.
A cell’s metabolic pathways make it easy for that cell
to use glucose for multiple purposes. Think of the cells as
■ Transferring the electrons carried by NADH and FADH2
to the terminal electron acceptor, which occurs as part of
synthesis of ATP in the process of oxidative phosphoryla- Moodboard/Brand X either
extensive biological recycling centers that routinely process Pictures/Getty
cellular respiration orImages
fermentation (recall that the
which all cell components are made—including proteins, the following general reaction (figure 6.10):
once a precursor metabolite is made in catabolism, it can GLUCOSE
be further oxidized to release energy, or it can be used in C6H12O6 + 62 O2Pentose →phosphate 6 CO2 + 6 H 2O
Yields
(glucose) (oxygen) (carbon dioxide) Oxidizes
(water)
1 Glycolysis
pathway ~ ~ + Reducing
biosynthesis.
glucose to pyruvate power
Starts the oxidation of glucose
ATP
by substrate-level
The pathways are catabolic, but the precursor metabolites and phosphorylation
Glucose-6-phosphate Lipopolysaccharide Glycolysis it to form two molecules of pyruvate. It provides the cell
Biosynthesis 5 Fermentation
Fructose-6-phosphate Peptidoglycan Glycolysis with a small amount of energy in the form of ATP, some Reduces pyruvate
or a derivative
Transition step
3a
3-Phosphoglycerate Proteins (the amino acids cysteine, glycine, and serine) Glycolysis To: To: (called the Entner-Doudoroff Yields
pathway) that generates CO aCO
2 2
Lipid Amino acid slightly different set of intermediates and end products.
Reducing
power
synthesis synthesis
Phosphoenolpyruvate Proteins (the amino acids phenylalanine, tryptophan, and tyrosine) Glycolysis
Acetyl- Acetyl-
■ Pentose phosphate pathway. 2 This also breaks down CoA CoA
Pyruvate Proteins (the amino acids alanine, leucine, and valine) Glycolysis glucose, but its primary role is to produce compounds
used in biosynthesis, including two precursor metabo-
Ribose-5-phosphate Nucleic acids and proteins (the amino acid histidine) Pentose phosphate cycle lites as well as reducing power in the form of NADPH. A
x2 CO 2
α-Ketoglutarate Proteins (the amino acids arginine, glutamate, glutamine, and proline) TCA cycle cycle, a single reaction called the transition step converts
the pyruvate from glycolysis into acetyl-CoA. One mol-
Yields
~ ~ + Reducing
power
Oxaloacetate Proteins (the amino acids aspartate, asparagine, isoleucine, lysine, TCA cycle ecule of CO2 is released as a result. 3b Theby substrate-level TCAATP
cycle 4 Cellular respiration
Uses the electron transport
phosphorylation
methionine, and threonine) then accepts the 2-carbon acetyl group, ultimately oxidiz- chain to convert reducing
power to proton motive force
FIGURE 6.10 Overview of Catabolism (1) Glycolysis, (2) the pentose phosphate pathway,
ing it to release two
(3a) themolecules of theCO
transition step, and (3b) . The
tricarboxylic
2 acidtransition
cycle (TCA cycle) are step
used to gradually
oxidize glucose completely to CO . Together, these pathways produce ATP, reducing power,
and the TCA cycle together generate
as precursor the most reducing
2
and intermediates that function metabolites (depicted as colored ovals, as shown Yields
CO2 molecules + energy ~ ~
Note: The colored icons in the table are used in figures throughout the chapter to represent the respective precursor metabolites. in table 6.2). (4) Cellular respiration uses the reducing power to generate a proton motive force
power of all thethatcentral
is then used tometabolic
make ATP by oxidative pathways; theypassing
phosphorylation, ultimately alsothe electrons ATP
to O or another terminal electron acceptor. (5) Fermentation stops short of oxidizing glucose by oxidative
FIGURE 6.9 Cells Use Glucose for Multiple Purposes The During the oxidation of glucose, a relatively small
millions of glucose molecules that continually enter a cell can have amount of ATP is made by substrate-level phosphorylation.
different fates. Some may be oxidized completely to release the The reducing power accumulated during the oxidation steps,
maximum amount of energy, and others will be used in biosynthesis. however, can be used in cellular respiration (discussed next)
xii
Signs and
Dengue and Severe Dengue
Often asymptomatic; fever, headache, rash,
Chikungunya
Similar to dengue fever, but
Zika Virus Disease
Amebiasis 666
Usually asymptomatic; mild disease with
Disease discussions are separated into consistent subsec-
CENTRAL NERVOUS SYSTEM INFECTIONS
Science Picture Co/Getty Images
Incubation Period
Causative Agents
Usually 4 to 7 days
Treatment and Treatment: analgesics for pain; oral rehydration Treatment: analgesics for pain Treatment: no specific treatment.
Prevention therapy and blood or platelet transfusions if and oral rehydration. Prevention: Prevention: vector control. Disease Causative Agent Comment Summary Table
bleeding occurs. Prevention: vector control; vector control. BACTERIAL INFECTIONS OF THE UPPER RESPIRATORY TRACT
vaccine in limited areas. Conjunctivitis (pink eye), Usually Haemophilus Often occur together; factors involved in the transmission are
otitis media (earache), sinus influenzae or unknown.
infection Streptococcus pneumoniae
Streptococcal pharyngitis Streptococcus pyogenes Treated with antibiotics, partly to avoid sequelae; must be Table 21.3
(“strep throat”) (group A streptococcus) distinguished from viral pharyngitis, which cannot be treated with
antibiotics.
Diphtheria Corynebacterium Toxin-mediated disease characterized by pseudomembrane in the Table 21.4
diphtheriae upper respiratory tract. Preventable by vaccination.
VIRAL INFECTIONS OF THE UPPER RESPIRATORY TRACT
Common cold Rhinoviruses and other Runny nose, sore throat, and cough are due to the inflammatory Table 21.5
viruses response and cell destruction.
Adenovirus pharyngitis Adenoviruses Similar to the common cold but with fever; spread to the lower Table 21.6
respiratory tract can result in severe disease.
Summarize each disease’s characteristics BACTERIAL INFECTIONS OF THE LOWER RESPIRATORY TRACT
Pneumococcal pneumonia Streptococcus pneumoniae Organism common in the throat of healthy people; causes disease
when mucociliary escalator is impaired or with underlying conditions.
Vaccine that protects against multiple strains is available.
Table 21.7
that includes an outline of the disease process keyed to a human figure, Tuberculosis (“TB”) Mycobacterium
tuberculosis
Most infections result in latent tuberculosis infection (LTBI), but these
can reactivate to cause tuberculosis disease (TB disease). Treated
using combination drug therapy, but drug resistance is an increasing
Table 21.9
Each disease chapter ends with a table that summarizes the key features of COVID-19, SARS and MERS
Hantavirus pulmonary
Coronaviruses
Hantaviruses
Emerging infectious diseases characterized by severe lower
respiratory symptoms; zoonotic
Acquired via inhaled dust contaminated with rodent saliva, urine, or
Table 21.14
Table 21.15
xiii
Chapter 1 – Humans and the Microbial World Chapter 5 – Control of Microbial Growth
■ Added SARS-CoV-2 and Candida auris to the section on ■ Combined the physical methods of microbial control into
emerging pathogens one section
■ Added the African swine fever to the list of epidemics in ■ Expanded the discussion of biosafety levels
non-human populations
■ Added the recent FDA rulings that limit the use of many
■ Expanded the coverage of the human microbiome previously allowed ingredients in antiseptic lotions until
■ Defined the term strain they are shown to be safe and effective
■ Moved the information about bacterial cell shape from
chapter 3 to section 1.3 Chapter 6 – Microbial Metabolism: Fueling
■ Added a MicroByte about the Microbiome Conservancy Cell Growth
collecting/storing fecal samples from populations around ■ Rearranged the information about energy sources and
the world terminal electron acceptors so that the more conceptually
simple information comes first.
Chapter 2 – The Molecules of Life ■ Revised tables 6.2 (Precursor Metabolites) and 6.4 (Some
■ Consolidated and expanded the information on water’s Vitamins and Their Use in Coenzymes)
characteristics ■ Added new figure (6.11) to emphasize the difference in
■ Added a subsection on short-chain fatty acids, to allow a energy yield between aerobic respiration and fermentation
description of butyrate
xiv
xv
Weekly Report
Response
■ Added the URL for the CDC’s National Notifiable Dis-
■ Extensive revision; reorganized the chapter to create a eases Surveillance System (NNDSS)
more linear flow (T cells and their activation are now
described before B cells)
■ Added COVID-19 and Candida auris infection to the sec-
tion on emerging diseases
■ Expanded and rearranged the overview to reflect the new
chapter organization Chapter 20 – Antimicrobial Medications
■ Expanded the discussion of immune tolerance to distin- ■ Added a Focus on the Future Box: The Race to Develop
guish between central tolerance and peripheral tolerance COVID-19 Treatments
Explained that oral administration of poorly absorbed
Chapter 16 – Host-Microbe Interactions
■
xvii
Denise Anderson
Sarah Salm
Mira Beins
xviii
PA RT I Covalent Bonds 23
Life and Death of Microorganisms Hydrogen Bonds 24
Molarity 24
1 Humans and the Microbial World 1 Chemical Reactions 25
2.3 Water, pH, and Buffers 26
A Glimpse of History 1
Water 26
Key Terms 1
pH of Aqueous Solutions 27
1.1 T
he Dispute over Buffers 27
Spontaneous Generation 2
2.4 Organic Molecules 28
Early Experiments 2
INTERFOTO/Alamy, Stock Photo Carbohydrates 29
Experiments of Pasteur 2
Lipids 30
Experiments of Tyndall 2
Proteins 33
The Golden Age of Microbiology 3
Nucleic Acids 38
The Scientific Method 3
1.2 M
icrobiology: A Human Perspective 5 FOCUS ON A CASE 2.1 32
The Human Microbiome 5 FOCUS YOUR PERSPECTIVE 2.1: Right-Handed and Left-Handed
Molecules 36
Microorganisms in the Environment 6
Commercial Benefits of Microorganisms 6 SUMMARY 41
Microbes as Research Tools 7 REVIEW QUESTIONS 42
Microbes and Disease 7
1.3 Members of the Microbial World 10
3 Cells and Methods to Observe Them 44
Scientific Names 11
A Glimpse of History 44
Bacteria 13
Key Terms 44
Archaea 14
Eukarya 14 PROKARYOTIC CELL STRUCTURES
Acellular Infectious Agents 15 AND THEIR FUNCTIONS
xix
SUMMARY 114
FOCUS ON A CASE 3.1 55
REVIEW QUESTIONS 115
FOCUS YOUR PERSPECTIVE 3.1: Pathogens Hijacking Actin 67
SUMMARY 86
REVIEW QUESTIONS 88
SUMMARY 172
6 Microbial Metabolism: Fueling Cell Growth 137 REVIEW QUESTIONS 173
10.2 Identification Methods Based on Phenotype 259 11.5 Aerobic Chemoorganotrophs 282
Microscopic Morphology 259 Obligate Aerobes 282
Culture Characteristics 260 Facultative Anaerobes 284
Metabolic Capabilities 260
ECOPHYSIOLOGICAL DIVERSITY
Serological Characteristics 262
Protein Profile 262 11.6 T
hriving in Terrestrial Environments 286
Bacteria That Form a Resting Stage 286
10.3 Identification Methods Based on Genotype 264
Bacteria That Associate with Plants 287
Detecting Specific Nucleotide Sequences 264
Sequencing Ribosomal RNA Genes 264 11.7 Thriving in Aquatic Environments 289
Whole Genome Sequencing 265 Sheathed Bacteria 289
Prosthecate Bacteria 289
10.4 Characterizing Strain Differences 266
Bacteria That Derive Nutrients from Other Organisms 290
Biochemical Typing 266
Bacteria That Move by Unusual Mechanisms 291
Serological Typing 266
Bacteria That Form Storage Granules 292
Whole Genome Sequencing 266
Phage Typing 267 11.8 Animals as Habitats 293
Antibiograms 267 Bacteria that Inhabit the Skin 293
Bacteria That Inhabit Mucous Membranes 294
10.5 Classifying Microorganisms 269
Obligate Intracellular Parasites 296
Sequence Analysis of Ribosomal Components 269
DNA-DNA Hybridization (DDH) 270 11.9 Archaea That Thrive in Extreme Conditions 299
Sequence Analysis of Genomes 270 Extreme Halophiles 299
G + C Content 270 Extreme Thermophiles 300
Phenotypic Methods 271
FOCUS ON A CASE 11.1 294
FOCUS ON THE FUTURE 11.1: Astrobiology: Searching for Life Beyond Filamentous Phage Infections: M13 Phage as a
Earth 301 Model 338
SUMMARY 301 13.3 The
Roles of Bacteriophages in Horizontal Gene
REVIEW QUESTIONS 303 Transfer 339
Generalized Transduction 339
12 The Eukaryotic Members of the Microbial Specialized Transduction 339
World 305
13.4 Methods Used to Study Bacteriophages 340
A Glimpse of History 305 13.5 Animal Virus Replication 341
Key Terms 305 Attachment 341
12.1 Fungi 306 Entry and Uncoating 341
Characteristics of Fungi 307 Synthesis of Viral Proteins and Replication of the
Classification of Fungi 309 Genome 342
Groups of Medically
Steve Gschmeissner/ Science Photo Assembly (Maturation) 345
Library/Getty Images
Important Fungi 310 Release 346
Economic Importance of Fungi 311 13.6 Categories
of Animal Virus Infections 347
Symbiotic Relationships of Fungi 312 Acute Infections 347
12.2 Protozoa 313 Persistent Infections 347
Characteristics of Protozoa 313 13.7 Viruses
and Human Tumors 349
Groups of Medically Significant Protozoa 314 Cancer-Causing Viruses 349
Other Protozoan Groups 315 Cancer-Fighting Viruses 350
12.3 Algae 318 13.8 Cultivating
and Quantitating Animal Viruses 351
Characteristics of Algae 318 Cultivating Animal Viruses 351
Types of Algae 319 Quantitating Animal Viruses 352
Exceptions to the Rule 320
13.9 Plant Viruses 353
12.4 M
ulticellular Parasites: Helminths 321
Life Cycles and Transmission of Helminths 321 13.10 Other Infectious Agents: Viroids and Prions 354
Roundworms (Nematodes) 322 Viroids 354
Tapeworms (Cestodes) 322 Prions 354
Flukes (Trematodes) 324 FOCUS ON A CASE 13.1 346
12.5 Arthropods 325 FOCUS YOUR PERSPECTIVE 13.1: Microbe Mimicker 335
FOCUS ON THE FUTURE 13.1: The Potential of Phage Therapy 356
FOCUS ON A CASE 12.1 317
FOCUS YOUR PERSPECTIVE 12.1: What Causes River SUMMARY 357
Blindness? 322 REVIEW QUESTIONS 358
SUMMARY 326
REVIEW QUESTIONS 327
PA RT III
13 Viruses, Viroids, and Prions 329 Microorganisms and Humans
14.3 The Cells of the Immune System 364 15.3 The T-Cell Response: Cell-Mediated Immunity 396
Granulocytes 365 General Characteristics of T Cells 396
Mononuclear Phagocytes 366 Activation of T Cells 397
Dendritic Cells 367 Effector Functions of TC (CD8) Cells 398
Lymphocytes 367 Effector Functions of TH (CD4) Cells 399
14.4 Cell Communication 368 15.4 The B-Cell Response: Humoral Immunity 402
Surface Receptors 368 General Characteristics of B cells 402
Cytokines 368 B-Cell Activation 402
Adhesion Molecules 368 Characteristics of Antibodies 402
14.5 Pattern Recognition Receptors (PRRs) 369 Evolution of the Humoral Response
to T-Dependent Antigens 404
Pattern Recognition Receptors (PRRs) That Monitor a
Cell’s Surroundings 370 The Response to T-Independent Antigens 407
Pattern Recognition Receptors (PRRs) That Monitor 15.5 Lymphocyte Development 408
Material Ingested by a Cell 370 Generation of Diversity 408
Pattern Recognition Receptors (PRRs) That Monitor a Negative Selection of Self-Reactive B Cells 408
Cell’s Cytoplasm 371 Positive and Negative Selection of Self-Reactive
An Outcome of Cytoplasmic Pattern Recognition: The T Cells 408
Interferon Response 371
15.6 Natural Killer (NK) Cells 409
14.6 The Complement System 372
Complement System Activation 373 FOCUS ON A CASE 15.1 410
Effector Functions of the Complement System 374 FOCUS YOUR PERSPECTIVE 15.1: What Flavors Are Your Major
Regulation of the Complement System 374 Histocompatibility Complex (MHC) Molecules? 401
16.6 Invasion—Breaching the Anatomical Barriers 423 17.5 Common Types of Immunoassays 453
Penetrating the Skin 423 Immunoassays That Use Labeled Antibodies 454
Penetrating Mucous Membranes 423 Immunoassays That Involve Visible Antigen-Antibody
Aggregates 457
16.7 Avoiding the Host Defenses 424
Hiding Within a Host Cell 424 FOCUS ON A CASE 17.1 453
Avoiding Destruction by Phagocytes 425 FOCUS YOUR PERSPECTIVE 17.1: Obtaining Monoclonal Antibodies 449
Avoiding Killing by Complement System Proteins 426
SUMMARY 461
Avoiding Recognition by Antibodies 427
REVIEW QUESTIONS 462
16.8 Damage to the Host 427
Exotoxins 427 18 Immunological Disorders 464
Endotoxin and Other Bacterial Cell Wall Components 430
Damaging Effects of the Immune Response 432 A Glimpse of History 464
Key Terms 464
16.9 Mechanisms of Viral Pathogenesis 433
Binding to Host Cells and Invasion 433 18.1 Hypersensitivities 464
Avoiding Immune Responses 433 Type I Hypersensitivities:
Damage to the Host 434 Immediate IgE-Mediated 465
©bubutu/Shutterstock
Type II Hypersensitivities:
16.10 Mechanisms of Eukaryotic Pathogenesis 434 Cytotoxic 468
Fungi 434 Type III Hypersensitivities: Immune Complex–Mediated 470
Protozoa and Helminths 435 Type IV Hypersensitivities: Delayed-Type
FOCUS ON A CASE 16.1 429 Cell-Mediated 471
FOCUS ON THE FUTURE 16.1: The Potential of Probiotics 435 18.2 Autoimmune Disease 473
SUMMARY 436 Systemic Autoimmune Diseases 474
REVIEW QUESTIONS 437 Organ-Specific Autoimmune Diseases 474
18.3 Immunodeficiency Disorders 476
17 Applications of Immune Responses 439 Primary Immunodeficiencies 476
Secondary Immunodeficiencies 477
A Glimpse of History 439
Key Terms 439 FOCUS ON A CASE 18.1 476
FOCUS YOUR PERSPECTIVE 18.1: The Fetus as an Allograft 473
IMMUNIZATION AND
IMMUNOTHERAPY SUMMARY 478
REVIEW QUESTIONS 479
17.1 Principles of ©Kevin Horan/The Image Bank/Getty
Immunization 440
Active Immunity 440
Images
19 Epidemiology 481
Passive Immunity 440 A Glimpse of History 481
17.2 Vaccines and Immunization Procedures 441 Key Terms 481
Attenuated Vaccines 441 19.1 Basic Concepts of
Inactivated Vaccines 442 Epidemiology 482
The Importance of Vaccines 443
An Example of Vaccination Strategy— 19.2 Chain of Infection 483 Source: CDC/James Gathany
The Campaign to Eliminate Poliomyelitis 444 Reservoirs of Infection 483
Portals of Exit 484
17.3 Immunotherapies 446 Disease Transmission 484
Immunotherapies for Cancer 446 Portals of Entry 487
Immunotherapies for Immunological Disorders 449
Immunotherapies for Infectious Diseases 450 19.3 Factors That Influence the Epidemiology of
Disease 487
IMMUNOLOGICAL TESTING The Dose 488
17.4 Principles of Immunoassays 451 The Incubation Period 488
Quantifying Antigen-Antibody Reactions 452 The Host Population 488
Obtaining Known Antibodies 452 The Environment 488
FOCUS ON A CASE 21.1 546 23.3 Diseases Due to Anaerobic Bacterial Wound
Infections 617
FOCUS YOUR PERSPECTIVE 22.1: A Global Lesson in Microbiology:
The COVID-19 Pandemic 571 Tetanus (“Lockjaw”) 617
DISEASES IN REVIEW 21.1: Respiratory System Diseases 578 Clostridial Myonecrosis (“Gas Gangrene”) 619
KEY TERMS
Domain The highest level in Prokaryote Single-celled organism
biological classification. There are consisting of a prokaryotic cell;
three domains: Bacteria, Archaea, members of the domains Bacteria
and Eukarya. and Archaea are prokaryotes.
Eukaryote Organism composed Prokaryotic Cell Cell type
of one or more eukaryotic cells; characterized by the lack of a
members of the domain Eukarya are membrane-bound nucleus.
eukaryotes. Viroid An acellular infectious agent
Eukaryotic Cell Cell type consisting only of RNA.
characterized by a membrane-bound Virus An acellular infectious agent
nucleus. consisting of nucleic acid surrounded
Prion An acellular infectious agent by a protein coat.
consisting only of protein.
others yet were ashed grey. And the motion of most of these
animalcules in the water was so swift, and so various, upwards,
downwards, and round about, that ‘twas wonderful to see.
M
(INTERFOTO/Alamy, Stock Photo)
icrobiology is the study of an amazing world made
up of members too small to be seen without the aid
A Glimpse of History of a microscope. Antonie van Leeuwenhoek described
this world when he observed what he called “animalcules”
Microbiology as a science was born in 1674 when Antonie van
through his simple microscope (figure 1.1). What he saw were
Leeuwenhoek, an inquisitive Dutch fabric merchant, looked at a drop
microorganisms (organisms too small to see with the naked
of lake water through a glass lens he had carefully made. Although
many people before him had used curved glass to magnify objects,
eye), including bacteria, protozoa, and some fungi and algae.
Leeuwenhoek’s skilled hands made a lens that uncovered a startling The microbial world also includes viruses and other infectious
and amazing sight—the world of microbes. As van Leeuwenhoek agents that are not considered organisms because they are not
wrote in a letter to the Royal Society of London, he saw composed of cells; they are acellular. When referring to general
members of the microbial world, the term microbe is often used.
Very many little animalcules, whereof some were roundish,
Microorganisms are the foundation for all life on Earth.
while others a bit bigger consisted of an oval. On these last, I
saw two little legs near the head, and two little fins at the hind
They have existed on this planet for about 3.5 billion years,
most end of the body. Others were somewhat longer than an oval, and over this time, plants, animals, and modern microorgan-
and these were very slow a-moving, and few in number. These isms have evolved from them. Even today, they continue to be
animalcules had diverse colours, some being whitish and trans- a driving force in the evolution of all living things. Microor-
parent; others with green and very glittering little scales, others ganisms may be small, but as you are about to learn, our life
again were green in the middle, and before and behind white; depends on their activities.
cork. At that time, brief boiling was thought to kill all organ-
isms, so this suggested that microorganisms did indeed arise
Handle spontaneously.
Focus screw In 1776, the animal physiologist and priest Lazzaro
Spallanzani obtained results that contradicted Needham’s
experiments; no bacteria appeared in Spallanzani’s broths
Specimen holder after boiling. His experiments differed from Needham’s in
Lens two significant ways: Spallanzani boiled the broths for lon-
ger periods, and he sealed the flasks by melting their glass
necks closed. Using these techniques, he repeatedly demon-
FIGURE 1.1 Model of van Leeuwenhoek’s Microscope The strated that broths remained sterile (free of microorganisms).
original made in 1673 could magnify an object almost 300 times. The However, if the neck of the flask cracked, the broth rapidly
object is brought into focus with the adjusting screws. Tetra Images/Alamy became cloudy due to the growth of organisms. Spallanzani
Stock Photo concluded that microorganisms had entered the broth with the
? What kinds of organisms did van Leeuwenhoek observe through his microscope? air, and the corks used by Needham and other investigators
did not keep them out.
Spallanzani’s experiments did not stop the controversy.
Some people argued that the heating process destroyed a
“vital force” in the air that was necessary for spontaneous
1.1 ■ he Dispute over Spontaneous
T generation, and so the debate continued.
Generation
Learning Outcomes Experiments of Pasteur
1. Describe the key experiments of scientists who disproved One giant in science who helped disprove spontaneous gen-
spontaneous generation.
eration was Louis Pasteur, the French chemist considered by
2. Explain how the successful challenge to the idea of many to be the father of modern microbiology. In 1861, he
spontaneous generation led to the Golden Age of
did a series of clever experiments. First, he demonstrated
Microbiology.
that air contains microorganisms. He did this by filtering
3. Describe the scientific method, using Pasteur’s swan-necked
air through a cotton plug, trapping microorganisms. He then
flask experiment as an example.
examined the trapped microorganisms with a microscope and
The discovery of microorganisms in various specimens found that many looked identical to those described by oth-
raised an interesting question: “Where did these microscopic ers who had been studying broths. When Pasteur dropped the
forms originate?” Some people believed that worms and cotton plug into a sterilized broth, the broth became cloudy
other life-forms arise from non-living material in a process from the growth of these microorganisms.
known as spontaneous generation. This was challenged by Most important, Pasteur demonstrated that sterile broths
an Italian biologist and physician, Francesco Redi. In 1668, in specially constructed swan-necked flasks remained sterile
he used a simple experiment to show that worms found on even when left open to air (figure 1.2). Microorganisms from
rotting meat originated from fly eggs, not from the decay- the air settled in the bends of the flask necks, never reaching
ing meat as supporters of spontaneous generation believed. the broth. Only when the flasks were tipped would microor-
In his experiment, Redi covered the meat with fine gauze ganisms enter the broth and grow. Pasteur's simple and ele-
that prevented flies from depositing their eggs; when he did gant experiments ended the arguments that unheated air or
this, no worms appeared. Despite Redi’s work, it took more the broths themselves contained a “vital force” necessary for
than 200 years and many experiments to amass conclusive spontaneous generation. They led to the theory of biogenesis,
evidence that microorganisms did not arise by spontaneous the production of living things from other living things (bio
generation. means “life”; genesis means “to create”).
Years Hours/days
1 Broth sterilized— 2 Broth allowed 3 Broth stays sterile 4 Flask tilted so that 5 Microorganisms
air escapes. to cool slowly— indefinitely. the sterile broth comes multiply in broth.
air enters. in contact with micro-
organisms from air.
showed that Pasteur was correct. Tyndall found that various Microbiology, during which time the field of microbiology
types of broths required different boiling times to be steril- blossomed. Many important advances were made during this
ized. Some were sterilized by boiling for 5 minutes, whereas period, including discoveries that led to the acceptance of
others, most notably broths made from hay, still contained the suggestion that microorganisms cause certain diseases, a
living microorganisms even after boiling for 5 hours! Even principle now called the Germ Theory of Disease.
when hay was merely present in the laboratory, broths that Figure 1.3 lists some of the important advances in micro-
had previously been sterilized by boiling for 5 minutes biology made over the years in the context of other historical
could not be sterilized by boiling for several hours. What events. Rather than cover more history now, we will return to
was going on? Tyndall finally realized that the hay con- many of these milestones in brief stories called “A Glimpse of
tained heat-resistant forms of microorganisms. When hay History” that open each chapter.
was brought into the laboratory, dust particles must have
transferred these heat-resistant forms to the broths. Tyndall
concluded that some microorganisms exist in two forms: a
The Scientific Method
cell easily killed by boiling, and one that is heat resistant. In The dispute over spontaneous generation offers an excellent exam-
the same year (1876), a German botanist, Ferdinand Cohn, ple of the process of science. This process, called the scientific
discovered endospores, the heat-resistant forms of some method, separates science from intuition and beliefs. The scien-
bacteria. tific method involves a series of steps, including:
The extreme heat resistance of endospores explains the ■■ Making an observation and asking a question about
differences between Pasteur’s results and those of other inves- that situation. An example from this chapter was the
tigators. Organisms that produce endospores are commonly observation that microorganisms were present in various
found in the soil and were likely present in broths made examined specimens. This observation led to the ques-
from hay. Pasteur used only broths made with sugar or yeast tion, “Where did the microorganisms originate?”
extract, so his experiments probably did not have endospores.
■■ Developing an explanation and then devising an
Scientists at the time did not appreciate the importance of the
experiment that tests the explanation. A testable
source of the broth, but in hindsight, the source was critical.
explanation of an observation is called a hypothesis,
This points out an important lesson for all scientists: When
and experiments are done to test the hypothesis. The
repeating an experiment, all conditions must be reproduced
dispute over spontaneous generation led to two oppos-
as closely as possible. What may seem like a trivial difference
ing hypotheses: biogenesis and spontaneous generation.
might be extremely important.
Various people designed different experiments to test
the hypotheses.
The Golden Age of Microbiology ■■ Doing the experiment, collecting the data, and draw-
The work of Pasteur and others in disproving spontane- ing a conclusion. Experiments such as the one illus-
ous generation started an era called the Golden Age of trated in figure 1.2 provided data about the growth of
1650 1700
1674
van Leeuwenhoek observes
microorganisms (Glimpse of
History, Chapter 1)
American Revolution
1775–1783
1750
FIGURE 1.3 Historical Events in Microbiology Some major milestones in microbiology—and their timeline in relation to other historical events.
The gold band indicates the Golden Age of Microbiology.
? What is the Golden Age of Microbiology?
The Human Microbiome Project changed the way scientists Cellulose-degrading microorganisms in the specialized stomach
view the human body and also revealed how much more there of ruminants (a group of plant-eating animals that includes cat-
is to discover about our microbial partners. To understand their tle, sheep, and deer) help those animals digest plant material.
significance, think of Earth’s ecosystems (the environments and Without the assistance of microbes, the ruminants would starve.
their interacting inhabitants). Over time, an interacting assort- In recognition of the important role that microorgan-
ment of organisms has evolved to live in a given environment, isms play in all aspects of life, additional programs promise
resulting in a relatively stable community. Sudden changes can to expand the scope of existing DNA-based studies. In 2016,
alter individual populations, often with negative consequences to the National Microbiome Initiative (NMI) was started to sup-
the community as a whole. In turn, a disturbance in one ecosys- port research on the microbiomes of humans as well as the
tem can affect the overall health of the planet. The human body, surrounding environment. Perhaps the most ambitious DNA
like a planet, is composed of various ecosystems—for example, sequencing program so far is the Earth BioGenome Project,
the desert-like dry areas of the skin, and the nutrient-rich envi- an international effort launched in 2018 to sequence all the
ronment of the intestinal tract. An important part of these eco- known animal, plant, protozoan, and fungal species.
systems is a population of interacting microbes. Disturbances in
a microbial population can create an imbalance that may have Commercial Benefits of Microorganisms
negative consequences to that community, which, in turn, can In addition to the crucial roles microorganisms play in our
harm a person’s health. Observations such as these have led very existence, they also have made life more comfortable for
some scientists to suggest that the human body be considered humans over the centuries.
a superorganism, meaning that our own cells interact with the
body’s normal microbiota to form a single cooperative unit. Food Production
The human microbiome’s effect on health and disease is Microorganisms have been used in food production since
an exciting area of active research, but it is more difficult to ancient times. In fact, Egyptians used yeast to make bread and
understand than it might seem. For example, researchers have beer. Virtually every population that raised milk-producing ani-
found that the intestinal microbiome of people diagnosed with mals such as cows and goats also developed procedures to fer-
depression differs from those who report a good quality of ment milk. This allowed them to make foods such as yogurt,
life, but this correlation could be an effect of mood—perhaps cheeses, and buttermilk. Today, the bacteria added to some fer-
even dietary changes associated with certain moods—rather mented milk products are advertised as probiotics (live micro-
than a cause. Likewise, bacterial species associated with gum organisms that provide a health benefit), protecting against
disease have been found in the brains of people with Alzheim- digestive disruptions.
er’s disease, but again, this could be effect rather than cause.
Continuing studies aim to clarify the situation. Biodegradation
Microorganisms play essential roles in degrading various envi-
MicroByte ronmental pollutants. These include materials in wastewater, as
The Global Microbiome Conservancy is collecting fecal samples
well as toxic chemicals in contaminated soil and water. Bacteria
from people around the world in an effort to study and preserve the
diversity of intestinal bacteria. also lessen the damage from oil spills. In some cases, microor-
ganisms are added to pollutants to hasten their decay, a process
called bioremediation.
Microorganisms in the Environment Commercially Valuable Products from Microorganisms
Microorganisms are the masters of recycling, and without them Microorganisms synthesize a wide variety of commercially
we would run out of certain nutrients. For instance, humans and valuable products. Examples include: antibiotics used to treat
other animals all require nitrogen, an essential part of nucleic infectious diseases, ethanol used as a biofuel, hydrogen gas
acids and proteins. A plentiful source of nitrogen is N2—the and certain oils potentially used as biofuels, amino acids used
most common gas in the atmosphere—yet neither plants nor as dietary supplements, insect toxins used in insecticides, cel-
animals can use it. Instead, we depend on certain microbes that lulose used in headphones, and polyhydroxybutyrate used in
convert N2 into a form of nitrogen that other organisms can the manufacture of disposable diapers and plastics.
use, a process called nitrogen fixation. Without nitrogen-fixing
microbes, life as we know it would not exist. Biotechnology
Microorganisms are also important because they can Biotechnology—the use of microbiological and biochemical
degrade certain materials that other organisms cannot. techniques to solve practical problems—depends on mem-
Cellulose (an important component of plants) is an excel- bers of the microbial world. Information learned by studying
lent example. Although humans and other animals cannot microorganisms led to easier production of many medications,
digest cellulose, certain microorganisms can, which is why including the insulin used to treat diabetes. In the past, insu-
leaves and fallen trees do not pile up in the environment. lin was isolated from the pancreatic glands of cattle and pigs,
but now certain microorganisms have been genetically engi- 40 states have
neered to make human insulin. The microbe-produced insulin 1,000
health departments
is easier to obtain, and patients who use it have fewer aller- Influenza pandemic
gic reactions than occurred with the animal-derived product.
800
Biotechnology also allows scientists to genetically engineer
plants to give them desirable qualities. Last human-to-human
transmission of plague
600
First use of penicillin
Microbes as Research Tools
Rate
400 First polio vaccine introduced
Microorganisms are wonderful model organisms to study
because they have the same fundamental metabolic and genetic Passage of
First continuous Vaccination Assistance Act
properties as higher life-forms. All cells are composed of the 200 municipal use
of chlorine in water
same chemical elements, and they synthesize their cell struc- in United States
tures by similar mechanisms. They all duplicate their DNA, 0
and when they degrade foods to harvest energy, they do so via 1900 1920 1940 1960 1980 2015
the same metabolic pathways. To paraphrase a Nobel Prize– Year
winning microbiologist, Dr. Jacques Monod: What is true of
FIGURE 1.4 Trend in Death Rates Due to Infectious Diseases
elephants is also true of bacteria, and bacteria are much easier Crude death rate for infectious disease, United States, per 100,000
to study! In addition, bacteria can be used to obtain results population per year.
very quickly because they grow rapidly and form billions of
? Why would the creation of health departments lower the disease rate?
cells per milliliter on simple, inexpensive growth media. In
fact, most major advances made in the last century toward
Past Triumphs
understanding life have come through the study of microbes.
The Golden Age of Microbiology included an important
period when scientists learned a great deal about pathogens.
Microbes and Disease Between 1876 and 1918, most pathogenic bacteria were iden-
Although most microbes are beneficial or not harmful, some tified, and early work on viruses had begun. Once people real-
are pathogens, meaning they can cause disease (a noticeable ized that microbes could cause disease, they tried to prevent
impairment in body function). The disease symptoms can their spread. As illustrated in figure 1.4, the death rate due to
result from damage caused by the pathogen’s growth and prod- infectious diseases has decreased dramatically over the last
ucts or by the body’s defense mechanisms inadvertently dam- 100 years or so, due largely to preventing the spread of patho-
aging host tissues during the attempt to control the infection. gens, developing vaccines to provide immunity, and using
To appreciate the effect an infectious disease can have on antibiotics to treat bacterial diseases when they do occur. To
a population, consider that more Americans died of influenza maintain this success, we must continue to develop new medi-
in 1918–1919 than were killed in World Wars I and II and the cations, vaccines, and disease-prevention strategies.
Korean, Vietnam, and Iraq wars combined. The COVID-19 pan- Perhaps the most significant triumph with respect to dis-
demic has resulted in the death of more than 1,000,000 people ease control was the eradication (elimination) of smallpox. This
worldwide, including over 200,000 Americans. viral disease was one of the most devastating the world has ever
Epidemics are not limited to human populations. The great known, killing about one-third of those infected. Survivors were
famine in Ireland in the 1800s was due, in part, to a microbial sometimes blinded and often left with disfiguring scars. When
disease of potatoes. A bacterial disease that kills olive trees was Europeans carried the disease to the Americas, the effect on the
found in southern Italy in 2013, and it has since spread to Spain populations of native inhabitants who had not been exposed before
and France, contributing to a recent worldwide drop in olive oil was catastrophic. A worldwide vaccination program eliminated
production. A fungal disease called “wheat blast” that devastated the disease in nature, with no cases being reported since 1977.
wheat crops in South America spread to Bangladesh in 2016, Laboratory stocks of the smallpox virus remain, however, raising
resulting in the loss of over 35,000 acres of crops that year. In the possibility that the virus could be used in bioterrorist attacks.
2001, a catastrophic outbreak of foot-and-mouth disease of live- Polio, a disease that can cause paralysis and sometimes
stock occurred in parts of England. To contain this viral disease, death, was once relatively common, but it has been nearly elim-
one of the most contagious known, almost 4 million pigs, sheep, inated because of vaccination. In fact, the disease now occurs
and cattle were destroyed. More recently, over a million pigs in in only a few countries, and the goal is to eradicate it globally.
China either died from African swine fever or were killed to con- Plague is another major killer that has largely been
tain the disease, and officials in other countries are trying to limit brought under control. In the fourteenth century, one-third of
its spread. Meanwhile, frog populations around the world have the population of Europe, or approximately 25 million people,
been decimated by chytridiomycosis, a fungal disease. died of this bacterial disease in only 4 years (1347–1351).
We now know that rodents can carry the bacterium, and their the globe. COVID-19 is caused by a virus officially called SARS-
fleas can transmit the disease, so we take measures to control CoV-2 (for severe acute respiratory syndrome coronavirus 2) but
the rodent populations. We have also learned that the pneu- commonly referred to as the COVID-19 virus. Like COVID-19,
monic form of the disease (meaning that it is in the lungs) can many EIDs are new or newly recognized; examples include
spread from human to human through respiratory secretions, Ebola disease (EBOD), congenital Zika syndrome, Candida
so special precautions are taken when a patient has pneu- auris infection, hepatitis C, severe acute respiratory syndrome
monic plague. In addition, the discovery of antibiotics in the (SARS), Middle East respiratory syndrome (MERS), certain
twentieth century made treatment possible. As a result, fewer types of influenza, Lyme disease, acquired immunodeficiency
than 100 people worldwide die from plague in a typical year. syndrome (AIDS), mad cow disease (bovine spongiform enceph-
alopathy), and hantavirus pulmonary syndrome (figure 1.5). Oth-
Remaining Challenges ers such as malaria and tuberculosis have been present for years
Although progress has been impressive against infectious but have spread or become more common recently.
diseases, much more still needs to be done. On a worldwide Some diseases arise as infectious agents evolve to infect
basis, infectious diseases remain too common, particularly new hosts, cause different types of damage, or become more
in developing countries. Even in developed countries with difficult to treat because of antibiotic resistance. Genetic
sophisticated healthcare systems, infectious diseases remain a analysis indicates that the virus that causes COVID-19 arose
serious threat, costing lives and money. from a strain that infects bats. HIV-1 (human immunodefi-
ciency virus type 1), the most common type of HIV to cause
Emerging Infectious Diseases An emerging infectious disease AIDS, arose from a virus that infects chimpanzees. A bac-
(EID) is an infectious disease that has become more common terium called E. coli O104:H4, which caused a severe food-
in the last several decades. The EID that everyone is now likely borne diarrheal outbreak in Europe, appears to have gained
familiar with is COVID-19 (for coronavirus disease 2019), the the ability to make a specific toxin by acquiring genes from a
disease that emerged in late 2019 and then spread rapidly around related organism. Tuberculosis and malaria have increased in
2012 2019
Middle East respiratory COVID-19 2002
syndrome (MERS) China Severe acute
Saudi Arabia respiratory
syndrome (SARS)
China
1986
Bovine spongiform
1982 encephalopathy
United Kingdom 1977
E. coli O157:H7 Hantaan virus
United States 1980 Republic of Korea
1981 Hepatitis D
2009
AIDS Italy
1989 Candida auris
United States Hepatitis C Japan
1976 United States
Legionnaires’ disease
1997
United States
1992 Avian flu (H5N1)
2009 1991 China
Venezuelan Vibrio
Swine flu cholerae 0139 2013
Mexico hemorrhagic Avian flu (H7N9)
fever India
1976 China
Cryptosporidiosis Venezuela
United States 1999
1994 Malaysian
Brazilian encephalitis
hemorrhagic Malaysia
fever 1976
Brazil Ebola disease 1994
Democratic Republic Human and equine
of Congo morbilivirus
2015 Australia
Congenital Zika
syndrome
Brazil
FIGURE 1.5 New and Newly Recognized Infectious Diseases or Disease Agents in Humans and Animals Since 1976 Countries where
cases first appeared or were identified appear in a darker shade.
? Why might so many of the diseases first appear or be identified in the United States and Western European countries?
FOCUS ON A CASE 1 .1
A 24-year-old woman had suffered from returned when she stopped taking the medi- Discussion
recurrent severe episodes of an intestinal cation. She also tried oral supplements con- 1. Antibiotics kill or inhibit not just patho-
disorder called Clostridioides (formerly taining Lactobacillus GG, a bacterium that gens, but also beneficial members of the
Clostridium) difficile infection (CDI) for sometimes appears to be effective in pre- normal microbiota, a group that protects
the past 13 months. She routinely experi- venting antibiotic-associated diarrhea. against infection in at least two gen-
enced profuse watery diarrhea, abdominal Because the patient’s health was declin- eral ways. First, the normal microbiota
pain, and fever. In addition, she was feeling ing, doctors suggested a fecal transplant, quickly uses nutrients that would other-
tired and hopeless because she did not seem an experimental procedure that involves wise be available to C. difficile and other
to be getting well, despite long attempts at inserting feces from a healthy person into pathogens. Also, some members of the
multiple different treatments. the patient’s intestinal tract in order to normal microbiota make compounds that
As with most patients who develop repopulate that environment with appropri- are toxic or inhibitory to other organisms.
CDI, the woman had been taking an oral ate microbes. They chose to use her sister The environment of the intestinal tract is
antibiotic shortly before her symptoms as a fecal donor, screening both the donor quite complex, however, so other factors
began—in this case, to treat a tooth infec- and the patient to ensure that neither was might also be playing a role.
tion. The antibiotic had successfully killed infected with certain microbes, includ- 2. Physicians screen the fecal donor to
the bacteria that caused her tooth infection, ing various intestinal pathogens and HIV. decrease the likelihood that disease-
but it also killed some members of her nor- Approximately ¼ cup of fresh feces was causing microbes could be transferred to
mal intestinal microbiota. As a result, the mixed with 1 quart of water and delivered the patient by the procedure. The doc-
bacterium C. difficile—often referred to to her intestinal tract via a colonoscope. tors screen the patients to ensure that
simply as “C. diff”—thrived in her intesti- Within days after the transplant, the patient they are not already infected with the
nal tract, growing to much higher numbers began feeling better, and she soon recov- pathogens. For example, if this patient
than it could before. The strain that caused ered completely. developed symptoms of a Salmonella
her infection was able to make a toxin that 1. Why would certain oral antibiotics infection after the procedure, how would
damaged the lining of her intestinal tract. allow C. difficile to thrive in the the physicians know that she acquired
When the patient first started experienc- intestinal tract? the infection as a result of the procedure
ing CDI, her doctor told her to stop taking if they had not checked beforehand?
2. Why would the doctors screen both the
the antibiotic prescribed for her tooth infec-
patient and the fecal donor for certain 3. Feces contain many types of bacteria
tion, hoping that her CDI would resolve on
infectious agents? that cannot yet be grown in the labora-
its own. When that did not help, the doctor
3. Why would the doctors transplant feces tory. In addition, scientists do not yet
prescribed a different antibiotic that is often
rather than introducing isolated bacteria know which types of fecal bacteria pro-
effective in treating CDI. The patient started
from feces to repopulate the colon? tect against CDI.
feeling better, but the symptoms quickly
incidence in recent years, in part because the causative organ- with misinformation about vaccines, and some people develop
isms became resistant to many of the available medications. irrational fears, falsely believing that vaccines are more harm-
As the rapid spread of COVID-19 around the globe cer- ful than the diseases they prevent. When this happens, parents
tainly demonstrated, mobile populations can contribute to dis- often refuse to vaccinate their children appropriately, leading
ease emergence as people may inadvertently carry pathogens to situations where the diseases become more common again.
to different regions. Even diseases such as malaria, cholera, Measles had been declared eliminated in the United States in
plague, and yellow fever that have largely been eliminated from 2000, but outbreaks in 2019 resulted in the highest number of
developed countries can be carried to other places if travelers cases in 25 years. Outbreaks generally start with unvaccinated
to regions where they still exist become infected and then move travelers who bring the disease into the country, where it then
on before recovering. Meanwhile, as city suburbs expand into spreads among others who are not vaccinated.
rural areas, human populations come into closer contact with Chronic Diseases Some chronic illnesses once attributed to
animals as well as the mosquitoes and other arthropods that nor- other causes may be due to microorganisms. Perhaps the best-
mally feed on those animals. Consequently, people are exposed known example is stomach ulcers, once thought to be due to
to pathogens they might not have encountered previously. stress. We now know that stomach ulcers are often caused by
The preventive measures used to control certain infectious a bacterium (Helicobacter pylori) and are treatable with anti-
diseases can become victims of their own success, a situation biotics. Chronic indigestion may be caused by the same bacte-
that can also lead to disease emergence. Decades of vaccination rium. Another example is cervical cancer, which we now know
have nearly eliminated measles, mumps, and whooping cough is caused by human papillomavirus (HPV) infection; a vaccine
in developed countries, so most people no longer have first- against HPV prevents that cancer. Infectious microbes may play
hand knowledge of the dangers of these diseases. Couple this important roles in other chronic diseases as well.
Acellular Infectious
Organisms
agents
Protists
FIGURE 1.6 The Microbial World Although adult helminths (worms) can often be seen with the naked eye, some stages in the life cycle of
helminths are microscopic.
? Members of which two domains are prokaryotes?
MicroAssessment 1.2 should be impressed by the assortment of what you see. That
range of types, however, is dwarfed by the huge variety of
Microbial activities are essential to human life as well as being
commercially valuable. Microbes are important research tools. microbes! The extent of that diversity makes sense considering
Although most microbes are beneficial or not harmful, some cause that microbes have inhabited this planet for billions of years and
disease. Enormous progress has been made in preventing and curing have evolved to thrive in every conceivable environment—from
infectious diseases, but some diseases are becoming more common. the hydrothermal vents at the bottom of the ocean to the icy tops
4. Describe two microbial activities essential to life and three of the highest mountains. Many people associate microbes with
that make our lives more comfortable. disease, but their contributions to our world go far beyond that. In
5. Describe three factors that cause certain infectious diseases fact, as section 1.2 described, we could not survive without them.
to become more common. Living organisms are all composed of cells with one of
6. Why would it seem logical, even inevitable, that at least two basic structures—prokaryotic (pro means “prior to” and
some bacteria would attack the human body and cause karyote means “nucleus”) and eukaryotic (eu means “true”).
disease? Prokaryotic cells do not have a membrane-bound nucleus.
Instead, the genetic material is located in a region called the
nucleoid. In contrast, the genetic material in eukaryotic cells
1.3 ■ Members of the Microbial World is contained within a membrane-bound nucleus. Eukaryotic
cells often have a variety of other membrane-bound organelles
Learning Outcomes as well, and they are typically much larger and more complex
8. Compare and contrast characteristics of members of the than prokaryotic cells. Organisms that consist of one or more
Bacteria, Archaea, and Eukarya. eukaryotic cells are called eukaryotes, whereas those com-
9. Explain the features of an organism’s scientific name. posed of a prokaryotic cell are called prokaryotes. Prokaryotes
10. Compare and contrast the algae, fungi, and protozoa. fall into two very different groups—bacteria and archaea—as
11. Compare and contrast viruses, viroids, and prions. different from each other as they are from eukaryotes.
Because of the fundamental differences between bac-
Considering that small size is the only shared feature of all teria, archaea, and eukaryotes, all living organisms are now
microbes, the group is tremendously diverse. If you look at the classified into three different domains: Bacteria, Archaea,
macroscopic world around you—the plants and animals—you and Eukarya (sometimes spelled Eucarya) (figure 1.6).
10
Nucleus
Atoms Lipids Ribosomes Smallest Most Most eukaryotic cells Adult roundworm
bacteria bacteria
Human height
Electron microscope
Light microscope
The basic unit of length is the meter (m), and all These units of measurement correspond to units
other units are fractions of a meter. in an older but still widely used convention.
nanometer (nm) = 10-9 meter = 0.000000001 meter 1 angstrom (Å) = 10-10 meter
micrometer (µm) = 10-6 meter = 0.000001 meter 1 micron (µ) = 10-6 meter
millimeter (mm) = 10-3 meter = 0.001 meter
1 meter = 39.4 inches
FIGURE 1.7 Sizes of Molecules, Non-Living Agents, and Organisms Note that the scale here is logarithmic (rather than linear), and each
labeled increment increases by a factor of 10.
? Why is a logarithmic scale useful when comparing sizes of members of the microbial world?
F O C U S YO U R P E R S P E C T I V E 1 .1
Every Rule Has an Exception
We might assume that because microorgan- In 1999, an even larger prokaryote was intestinal bacterium Escherichia coli. The tiny
isms have been so intensively studied over isolated from the muck of the ocean floor organism was found attached to a much larger
the past hundred years, no major surprises off the coast of Namibia in Africa. It is a microbe, a member of the Archaea grow-
are left to be discovered. This, however, spherical organism 70 times greater in ing in an ocean vent where the temperature
is far from the truth. In the mid-1990s, a volume than E. fishelsoni. Since it grows was close to the boiling point of water. The
large, peculiar-looking organism was found on sulfur compounds and contains glis- larger organism is an Ignicoccus species (igni
in the intestinal tract of certain fish from tening globules of sulfur, it was named means “fire” and coccus means “sphere”).
both the Red Sea in the Middle East and Thiomargarita namibiensis, meaning “sul- The tiny one, also a member of the Archaea,
the Great Barrier Reef in Australia. This fur pearl of Namibia.” has been named Nanoarchaeum equitans
organism, named Epulopiscium fishelsoni, In contrast to the examples of large (meaning “tiny archaea” and “rider”). N.
cannot be cultured in the laboratory. Its bacteria, a unicellular alga found in the equitans cannot be grown in the laboratory by
large size (600 μm long and 80 μm wide) Mediterranean Sea is 1 μm in width. It is a itself, but Ignicoccus grows well without its
make it clearly visible without any magni- eukaryote even though it is about the size of Nanoarchaeum “rider.”
fication and suggested that this organism a typical bacterium. These exceptions to long-standing rules
was a eukaryote. However, it does not have How small can an organism be? A point out the need to keep an open mind
a membrane-bound nucleus. A chemical microbe discovered off the coast of Iceland and not jump to conclusions! They also
analysis of the cell confirmed that it is a is only about 400 nm (nanometers) in diam- serve as excellent reminders that in a sub-
prokaryote and a member of the domain eter and has one-tenth the amount of genetic ject as complex as microbiology, there will
Bacteria. information (DNA) compared to the common almost always be exceptions!
part indicating the genus is commonly abbreviated, with the Members of the same species may vary from one another in
first letter capitalized—as in E. coli. minor ways, but not enough to separate the organisms into differ-
The origin of one or both parts of the name often reflects ent species. A genetic variant within a species is called a strain.
a characteristic of the organism or honors a particular sci- In situations where genetic differences are important, such as in
entist (table 1.2). In the case of Escherichia coli, the name research, a particular microbe and its progeny may be indicated
of the genus honors Theodor Escherich, who discovered the with a strain designation—for example, E. coli B or E. coli K12.
bacterium; the species designation indicates the site where Groups of microbes are often referred to informally by
E. coli typically lives: the colon (large intestine). Within a names that resemble genus names but are not italicized. For
given genus, there may be a number of different species. For instance, members of the genus Staphylococcus are often
example, the genus Escherichia includes species other than E. called staphylococci.
coli, such as E. vulneris, which was first isolated from human
MicroByte
wounds (vulneris means “of a wound”). E. vulneris is geneti-
Bacterial species outnumber mammalian species by more than
cally related to E. coli, but not closely enough to consider it in 10,000-fold!
the same species.
Escherichia coli (bacterium) Honors Theodor Escherich, the scientist who discovered Derived from the word “colon,” the body site
the bacterium. inhabited by the bacterium.
Haemophilus influenzae (bacterium) Derived from haemo (blood) and phil (loving), reflecting Derived from the word “influenza,” the disease
that the bacterium requires certain components of blood mistakenly thought to be caused by the
for growth. bacterium; we now know that influenza is
caused by a virus.
Saccharomyces cereviseae (fungus) Derived from saccharo (sugar) and myces (fungus). Derived from cerevisia (beer), reflecting that the
fungus (a yeast) is used to make beer.
Shigella dysenteriae (bacterium) Honors Kiyoshi Shiga, the scientist who discovered the Derived from the word “dysentery,” the disease
bacterium. caused by the bacterium.
Staphylococcus aureus (bacterium) Derived from staphylo (bunch of grapes) and kokkus The term aureus (golden) indicates the common
(berry), reflecting the grouping and shape of the cells. color of visible masses of the cells.
XIII
Serge was now nineteen years of age. He occupied a small room on
the second floor, opposite the priest's, and there led an almost
cloistered life, spending much time in reading.
'I shall have to throw those old books of yours into the fire,' Mouret
said to him angrily. 'You'll end by making yourself ill and having to
take to your bed.'
The young man was, indeed, of such a nervous temperament, that
the slightest imprudence made him poorly, as though he were a
young girl, and thus he was frequently confined to his room for two
or three days together. At these times Rose inundated him with herb
tea, and whenever Mouret went upstairs to shake him up a little, as
he called it, the cook, if she happened to be there, would turn her
master out of the room, crying out at him:
'Leave the poor dear alone! Can't you see that you are killing him
with your rough ways? It isn't after you that he takes: he is the very
image of his mother; and you'll never be able to understand either
the one or the other of them.'
Serge smiled. After he had left college his father, seeing him so
delicate, had hesitated to send him to Paris to read for the bar there.
He would not hear, however, of a provincial faculty; Paris, he felt
sure, was necessary for a young man who wanted to climb to a high
position. He tried, indeed, to instil ambitious ideas into the lad,
telling him that many with much weaker wits than his own, his
cousins, the Rougons, for instance, had attained to great distinction.
Every time that the young man seemed to grow more robust, his
father settled that he should leave home early the following month;
but his trunk was never packed, for Serge was always catching a
fresh cold, and then his departure would be again postponed.
On each of these occasions Marthe contented herself with saying in
her gentle, indifferent way:
'He isn't twenty yet. It's really not prudent to send so young a lad to
Paris; and, besides, he isn't wasting his time here; you even think
that he studies too much.'
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