Chapter 4

CHAPTER 4
4 Prokaryotic Organisms
BENEFICIAL BACTERIA
Ch 04: Chapter Opening Video
The Staphylococcus aureus bacteria growing on the cilia of human nasal epithelial cells
make up part of our personal prokaryotic world
.
Prokaryotes, which include bacteria, are the smallest, simplest, single-celled organisms. Most
people equate bacteria with pathogens, but only a small fraction of these microorganisms cause
human disease. In fact, the extensive bacterial community living in and on us (see the photo)
composes our normal microbiota, which can protect us from infection. Because these well-
established microbes are better competitors for resources, they often prevent pathogens from
colonizing our bodies. Microbiota also improve our health by aiding digestion and producing
vitamins.
Other beneficial prokaryotic roles include bioremediation, or the use of unique bacterial enzymes
to degrade environmental contaminants. Bacteria increase soil fertility, enhancing agriculture
yields, and are used to manufacture foods such as cheese, yogurt, pickles, and chocolate.
Alterations of bacterial genomes create microbes that make environmentally friendly pesticides,
biofuels, and medications. Currently, investigators are experimenting with biocryptography, or
the use of bacteria to store hack-proof data, text, and videos. Microbiologists estimate that for
every one pathogenic bacterium, there are 30,000 beneficial ones. Because less than 1% of
prokaryotes can be grown in a laboratory (see the photo) and a much smaller fraction have been
characterized, a wealth of bacterial benefits await discovery.
This chapter will survey the structures that enable prokaryotes to adapt to a myriad of
different environments. It will also summarize prokaryotic evolution, classification, and
how prokaryotes have changed the chemical and physical makeup of the planet.
4.1 The Prokaryote's Place in the Living World

LEARNING OBJECTIVES
 1.Describe the roles of prokaryotes in sustaining life.
 2.Explain the symbiotic interactions prokaryotes can have with other living organisms.
Prokaryotes are small, unicellular organisms that lack a distinct nucleus and complex
membrane-bound organelles. Their cellular makeup is distinctly different from the cells
of eukaryotes—the plants, animals, and fungi of the macroscopic world and the microscopic
protozoans (Table 4.1). From a clinical perspective, both prokaryotes and microbial eukaryotes
can act as pathogens. However, because of their significantly different cellular compositions,
correspondingly different drug therapies must be used to cure these infections. Bacterial
infections are treated by administering drugs designed to target prokaryote-specific structures to
prevent host damage. To protect the host while curing an infection caused by a eukaryotic
microbe, medications typically must selectively damage features unique to the specific pathogen.
A comparison of prokaryotic and eukaryotic cellsTable4.1
Feature Prokaryotes Eukaryotes
Size 0.2-5 μm 10-100+ μm
Genetic Genetic information carried by DNA in one nucleoid or circular Genetic information carried
material chromosome; may contain plasmids carrying accessory genes chromosomes
Protein 70S ribosomes 80S ribosomes
synthesis
A comparison of prokaryotic and eukaryotic cellsTable4.1
Feature Prokaryotes Eukaryotes
Cell wall Structurally complex, composed of peptidoglycan Absent or structurally simpl
silica
DNA No membrane-bound nucleus True membrane-bound nucl
organization
Organelles No true membrane-bound organelles Many complex membrane-b
Cell division Replicate by binary fission, a simple cell division process that Replicate by mitosis/meiosi
separates copied nucleoids into daughter cells separate many copied chrom
Example
Euglena gracilus
Escherichia coli
Prokaryotic microbes are the dominant form of life on Earth (see What a Microbiologists Sees).
The two types of prokaryotic cells—archaea and bacteria—are widely distributed and live in
nearly every possible environment on the planet. The total prokaryotic biomass is 10,000 times
greater than the biomass of all humans currently living on Earth. Incredibly, there are about 10
million times more archaea in the oceans than there are stars in the visible universe, and there are
10 times more bacteria living in and on us than there are cells in our bodies.
WHAT A MICROBIOLOGIST
SEESProkaryotes—The Dominant Form of Life on Earth
Examine Figure a and identify the various forms of life in this community. The list of organisms
compiled by most people would include the plants, aquatic creatures in the pond, the boy, and
the cow. But at the top of a microbiologist's list would be the invisibly small prokaryotes. What a
microbiologist sees is the dominant form of life on Earth. As Figure b shows, prokaryotic
microbes are the most numerous of all life forms. They also demonstrate the greatest species
diversity. It is estimated that there are about 1000 times more bacterial species than animal
species on Earth. The number of archaean species is reported as TBD, or as yet to be determined,
because their diversity is so great that we do not yet have an accurate count. Finally, prokaryotic
microbes occupy every imaginable habitat. In addition to the number of prokaryotes living in the
locations indicated in Figure a, many survive deep underground or flourish in extreme
environments such as boiling hot geysers, crude oil-polluted water, and deep sea vents.
 a.
 b.
Organism group Estimated number of species
Plants 500,000
Fungi 1,500,000
Protists 1,000,000
Animalsa 6,000,000
Bacteria 1,000,000,000
Archaea TBD
 a
Insects and mites make up approximately 40% of animal species.
Think Critically
According to Figure a, there are approximately 10,000 times more bacteria living in your
intestines than in your mouth. Why does such a difference exist between two locations in the
same person?
The intestines are much larger, providing more room for bacteria to grow, and they contain an
enormous food resource to promote bacterial growth.
Sustaining Life
The activities of prokaryotes sustain all life on the planet in today's world. Photosynthetic
bacteria along with plants and marine algae produce glucose and O2, which are used by aerobic
organisms. Additionally, prokaryotes are the only organisms capable of nitrogen fixation, the
enzymatic process by which atmospheric N2, which cannot be used by other living organisms, is
converted into usable nitrogen-containing organic compounds. On land, bacteria living within
the roots of certain types of plants generate NH4+ that the plant can use to synthesize proteins
and nucleic acids. This process not only benefits the plant, but also the nitrogen-fixing bacteria
receive nutrients stored in the roots. Without microbial nitrogen fixation, plant life on the planet
would run out of usable nitrogen in about a week. If the plants died, then the herbivores, or plant-
consuming animals, would starve and the food web would unravel.
Prokaryotes also sustain life by helping cows and other grazing animals break down the cellulose
in the plants they eat. They have specialized chambers in their stomachs that culture prokaryotic
microbes that digest much of the plant material, making these nutrients available to the animal.
Prokaryotes are also critical components of biogeochemical cycles that allow chemicals such as
carbon, phosphorus, and sulfur to cycle between the living and nonliving worlds. Clearly,
prokaryotes are a foundational component of the food webs on which much of the life on the
planet depends.
Symbiotic Relationships
For about 2 billion years, until eukaryotes evolved, prokaryotes were the only living things on
Earth. As a result, eukaryotic microbes evolved in a soup of prokaryotic competitors. When
eukaryotic organisms developed into multicellular forms, it was done in the constant presence of
prokaryotes growing on and in them. This type of relationship in which one organism lives in or
on another is called symbiosis. In these relationships, one organism is generally supplied with
food, oxygen, a safe living environment, or some other necessary factor. Symbiotic relationships
benefit one of the species involved, but can be classified into types depending on how the other
species in the relationship is affected. In a symbiotic relationship between a prokaryotic
organism and a macroscopic organism, the prokaryotic organism always benefits, with different
effects on the other organism.
In mutualism, both species benefit; in commensalism, one species benefits and the other
species is neither harmed nor benefited; and in parasitism, one species benefits and the other
species is harmed (Figure 4.1). Bacteria can live on and in humans without causing significant
benefit or harm, a commensal relationship. Because normal microbiota are important in
inhibiting the growth of bacterial pathogens, this relationship is often considered mutualistic.
However, when tissues are damaged by illness or injury, these bacteria may also cause
infections, resulting in a parasitic relationship.
Symbiotic relationships• Figure 4.1Symbiotic relationships do not necessarily fit into discrete
classes, but form a continuum from relationships that are mutually essential to those that are so
harmful as to be fatal to the host.
Interactive Graphic: Figure 4.1
Think Critically
Fibrobacter succinogenes digests cellulose in the rumen of a cow. What type of symbiotic
relationship do you think they have and why?
They have a mutual relationship because the bacteria degrade the cellulose, providing
nourishment for the cow, and the cow provides protection and nutrients for the bacteria.
Bacteria that live on and in humans have all different types of symbiotic relationships.
Bacteria that live on our skin can at times have a commensal relationship with no significant
benefit to us. However, bacteria that live in our large intestine can at times have an important
role in inhibiting the growth of bacterial pathogens; hence, that relationship is often mutualistic.
Bacterial pathogens do cause harm when they damage tissues and cause illness. Such bacteria are
considered parasites.
Copyright © 2017 John Wiley & Sons, Inc. All rights reserved.
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4.2 Bacterial Cell Shapes and Arrangements

LEARNING OBJECTIVES
 1.Describe the common shapes of bacterial cells.
 2.Identify the different bacterial arrangements, how they form, and their clinical
significance.
Bacteria can have different arrangements depending on how they adhere to one another after cell
division. The size range for bacteria is usually 0.2 to 2.0 μm in diameter and 2 to 8 μm in length
(Figure 4.2a). However, Mycoplasma gallicepticum is known as an ultramicrobacterium
because, at 200 to 300 nm in diameter, it is substantially smaller than most bacteria. On the other
end of the size spectrum, Thiomargarita namibiensis, reported to be the largest bacterial species
at 750 μm, is visible to the naked eye.
Microbiology InSightBacterial morphology • Figure 4.2
Bacterial species are characterized by their size, shape, and arrangement. These features can
often be used to identify the bacterial genus presumptively, facilitating diagnosis and allowing
for rapid initiation of patient therapy.
 a.Bacterial size
The range of different sizes of bacteria is dramatic, with an approximately 4000X
difference in diameter between the smallest and largest bacterial species. Some species
such as Mycoplasma pneumoniae are as small as a large virus, whereas Thiomargarita
namibiensis can be seen without a microscope. Most bacteria of clinical significance,
such as Streptococcus pneumoniae, are approximately 1 μm in diameter.

 b.Bacterial shapes and arrangements
Most bacteria are coccus, bacillus, or spiral shaped, but tremendous variations on these
forms also exist. In addition to variety in shape, cell division in different planes results in
various bacterial arrangements when daughter cells remain attached. The names for
different morphologies describe their shapes and groupings, as shown in the smaller
table.
Basic shapes Groupings

Term Meaning Term Meaning
bacillus staff diplo- double
vibrio wave strepto- twisted chain
Basic shapes Groupings
Term Meaning Term Meaning
spirillum coil staphylo- cluster
coccus kemel tetrad group of four
spirochete coiled hair sarcinae bundle
 c.The clinical significance of bacterial morphology
Examination of a Gram-stained patient specimen for morphological determination is
often the first step in diagnosis. For example, a sputum specimen containing gram-
positive, club-shaped rods in a palisade arrangement strongly suggests the
pathogen Corynebacterium diphtheriae and a patient suffering from diphtheria, a serious
respiratory infection.
Ask Yourself
How would you describe rod-shaped bacteria linked together to form long chains?
 a.
staphylococci
 b.
staphylobacilli
 c.
streptococci
 d.
streptobacilli
 d.
streptobacilli
3D Animation: What do bacteria look like?
Bacterial Shapes
Because there are so many bacterial species, it is not surprising that they demonstrate variable
forms. The three common shapes of bacteria are coccus (plural cocci), or
spherical; bacillus (plural bacilli), or cylindrical; and spiral (Figure 4.2b). Each of these basic
shapes demonstrates some species-specific modifications. For example, the two cocci that
compose Streptococcus pneumoniae are slightly angular or diamond-shaped. Cocci can also be
modified to be chubby ovals or flattened circles. Bacilli range in width from slender to thick;
their ends may be angular or rounded; they can be cylindrical or flat; and, although they are
usually straight rods, vibrios curve, resembling elbow macaroni. Spiral bacteria are elongated
bacilli that coil into flexible spirochetes or rigid helical spirilla.
Some bacterial species deviate from the standard morphologies. Fusiform cells are exaggerated
bacilli with distinctly tapered tips. Other bacilli resemble a club, having one swollen end.
Filamentous bacterial forms may be straight chains or branched, looking similar to fungi.
Appendaged bacteria are characterized by the presence of bumplike projections.
Pleiomorphism describes slight shape variations of bacterial cells of the same species. These
morphological modifications may represent differences in individual genetic makeup, nutrition,
or environmental factors. Because the characteristic form of a bacterial cell is maintained by the
presence of its cell wall, anything that affects this supportive surrounding structure can influence
shape. Mycoplasma species are noted for their variable cell shapes because members of this
genus lack cell walls.
Bacterial Arrangements
Bacterial morphology is also characterized by the species-specific grouping or arrangement of
cells (Figure 4.2c). In addition to single cells, common arrangements include pairs (described by
the prefix diplo-), chains (strepto-), and clusters (staphylo-). Some bacterial species form
flattened sets of four (tetrads) or three-dimensional packets of eight (sarcinae). Short stacks of
bacilli can also group to resemble Chinese character writings and are known as palisades (Figure
4.2d). The various bacterial arrangements occur when the daughter cells resulting from cell
division in different planes remain attached to each other.
Although the terms listing the descriptions of the shapes and groupings of different types of
bacteria are relatively straightforward, they can sometimes be confusing because these same
names are used for certain genera designations. For example, streptococcus is the general term
for any group of bacteria that is coccus-shaped and grouped in chains. However, there is also a
genus named Streptococcus that contains several important pathogenic microorganisms. As you
read through future material, it will be important to notice whether terms such as staphylococcus,
bacillus, and vibrio are lowercase (simply descriptive) or uppercase and italicized (referring to a
particular genus of bacteria).
Because specific sizes, shapes, and arrangements characterize bacterial species, microscopic
analysis in the clinical laboratory to determine these features is often used as the first step in
pathogen identification (Figure 4.2d). When a patient specimen is received in the laboratory, it
is Gram stained (see Remember This!) for morphological analysis. These results, coupled with
the type of patient specimen, can be diagnostic or at least suggest follow-up testing for accurate
determination. For example, if gram-negative diplococci are found in a type of white blood cells
called neutrophils from a sample of penile discharge, then Neisseria gonorrhoeae is identified,
the patient diagnosed with gonorrhea, and the appropriate antibiotic therapy initiated. The ability
of clinical microbiologists to correlate bacterial morphology with particular pathogens is
essential for positive patient outcomes.
REMEMBER THIS!
Gram staining is a foundational skill in microbiology. This differential staining technique reveals
key cell wall variations between bacterial species and highlights cell morphology. Refer to
Figure 3.6 to review the steps and outcomes of the Gram staining procedure.
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4.3 The Bacterial Cell Wall

LEARNING OBJECTIVES
 1.Diagram the peptidoglycan structure of cell walls.
 2.Differentiate between the structures of gram-positive and gram-negative cell walls.
 3.Describe the cell walls of acid-fast bacteria and the structure of wall-less bacteria.
Nearly all bacterial species possess a cell wall, which provides support and protection for the cell
and determines cell shape. A cell wall prevents cell lysis, or bursting. Bacteria living in
freshwater are in a hypotonic environment, a medium with a solute concentration that is lower
than that found inside the cell. As a result, water spontaneously diffuses into the cell by the
process of osmosis (Figure 4.3). Cell walls prevent cell lysis by resisting the pressure from the
osmosis of water into a cell.
The role of the cell wall in preventing osmotic lysis• Figure 4.3As water flows into a
bacterium with a cell wall, the cell wall expands by about 25%, but does not rupture. Because the
plasma membrane is delicate and nonelastic, a bacterium without a cell wall will rupture in
response to increased internal pressure and the cell will die.
Think Critically
In which direction would water flow for a cell placed in a hypertonic solution (one with a higher
solute concentration than the cell contents), and what would happen to the cell contents?
Water would flow out of the cell, and the cell contents would shrink.
Cell Wall Structure
Peptidoglycan, the primary component of bacterial cell walls, consists of peptides linked to
sugars called glycans. Bacteria are classified into two main groups based on the chemical
composition of the peptoglycan in their cell walls: gram-positive bacteria and gram-
negative bacteria.
The basic peptidoglycan unit consists of a disaccharide composed of N-acetylglucosamine
(NAG) and N-acetylmuramic acid (NAM), modified glucose molecules. In the cytoplasm, a
tetrapeptide side chain attaches to NAM before the disaccharide subunit is transported across the
plasma membrane (Figure 4.4a). The sugar subunits are joined together enzymatically outside
the cell to form glycan chains typically 20 to 30 units long.
The structure of peptidoglycan• Figure 4.4Peptidoglycan is composed of disaccharides linked
to peptide chains. It provides the physical strength to maintain the structural integrity of the cell.
a. Peptidoglycan monomer
The peptidoglycan monomer is synthesized sequentially inside the cell beginning with the
disaccharide, which is then linked to the amino acid chains and transported across the membrane.
b. Crosslinking of glycan chains

After peptidoglycan monomers are joined together enzymatically to form glycan chains, the
strands are crosslinked to form a three-dimensional network. Gram-positive bacterial species
accomplish this by joining adjacent tetrapeptide side chains with pentaglycine cross bridges,
whereas gram-negative species bind the tetrapeptides directly together.
Ask Yourself
NAG is linked to ________.
 a.
NAM
 b.
the pentaglycine crossbridge
 c.
the tetrapeptide side chain
 d.
Both a and b are correct.
 a.
NAM
Animation: Peptidoglycan
Next, the glycan chains are linked together to form a sheet. Gram-positive bacteria join the
glycan chains together using pentaglycine crossbridges between the tetrapeptide side chains
(Figure 4.4b). The crossbridges are enzymatically attached to the number three position on one
tetrapeptide and the number four position on the other tetrapeptide. In gram-negative bacteria, a
covalent bond directly joins adjacent tetrapeptide side chains at the number three and four
positions. In this way, the associated peptidoglycan chains can form rings to encompass the cell.
Further crosslinking joins adjacent rings and chains to form a three-dimensional coating that
surrounds the cell.
There is free diffusion of molecules through the peptidoglycan layer. Small molecules and
globular proteins diffuse through spaces in the peptidoglycan. The pores result from incomplete
crosslinking between the glycan chains.
Pharmaceutical chemists use their knowledge of peptidoglycan cell wall structure to develop
effective antibiotics. For example, vancomycin inhibits peptidoglycan synthesis by preventing
the transport of disaccharide subunits outside the cell for assembly into the cell wall. Penicillin
binds to the enzyme responsible for joining the crossbridges between tetrapeptides. This
inactivates the enzyme, preventing cell wall synthesis and leading to bacterial death. Another
antibacterial compound that targets peptidoglycan is the lysozyme secreted in tears, mucus, and
sweat. This naturally occurring compound cleaves the peptidoglycan chain at specific intervals,
weakening the wall and allowing osmotic lysis.
Gram-Positive and Gram-Negative Cell Walls
In addition to the structural differences in the chemical composition of the peptidoglycan just
discussed, gram-positive bacteria have a thick layer of peptidoglycan, and gram-negative
bacteria have a thin layer of peptidoglycan. These differences in cell wall structure play an
important role in distinguishing bacterial species in the lab. Gram-positive bacteria stain dark
purple or violet after Gram staining because of the thick layers of peptoglycan in the cell wall. In
contrast, gram-negative bacteria stain pink or red because of the thin layer of peptidoglycan (see
Remember This!).
REMEMBER THIS!
The Gram stain is used to identify gram-positive bacteria and gram-negative bacteria. If
necessary, review the Gram staining procedure from Section 3.3 before continuing.
Gram-positive cell walls
In the cell walls of gram-positive bacteria, the peptidoglycan consists of 6 to 12 layers of glycan
sheets and measures 70- to 80-nm thick. There is a space between the plasma membrane and the
peptidoglycan known as the inner wall zone. This space serves as reservoir for metabolites for
cell wall synthesis.
Chains of teichoic acids weave through the peptidoglycan (Figure 4.5a). This polymer is
composed of sugar-phosphate subunits called wall teichoic acids and lipoteichoic acids. Wall
teichoic acids attach to cell wall NAMs, whereas lipoteichoic acids attach to phospholipids in the
plasma membrane. Teichoic acid strands often have additional sugars and alanines attached
along their sides. Extending beyond the surface of the cell wall, teichoic acid strands compose
approximately 60% of the carbohydrate component of a gram-positive cell wall.
Bacterial cell wall structure• Figure 4.5The key structural component of both gram-positive
and gram-negative bacteria is peptidoglycan. However, there are several significant differences
between the two cell types.
a. The gram-positive cell wall
Gram-positive bacteria have cell walls consisting of several layers of peptidoglycan interwoven
with chains of teichoic acids.
b. The gram-negative cell wall

Gram-negative bacteria have cell walls that consist of one or only a few layers of peptidoglycan
and an outer membrane with porin proteins and lipopolysaccharides in the outer layer.
Ask Yourself
In gram-negative bacteria, the outer membrane, unlike the plasma membrane,
contains lipopolysaccharide and porin proteins .
Teichoic acids perform many functions, including the regulation of cation flow through the thick
layer of peptidoglycan and the maintenance of cell shape. In some pathogenic streptococci,
teichoic acids enable the bacteria to attach to the host tissues. The negative charge from the
phosphate groups of teichoic acid repels negatively charged phagocytic cells (cells able to engulf
microbes), allowing some pathogens to avoid immune attack. Teichoic acids are also linked to
regulation of cell growth and division and enhanced survival when microbes are subjected to
elevated temperatures and solutes. Because teichoic acids are such an important cell wall
component, pharmaceutical chemists are developing antibiotics to target teichoic acid synthesis
to induce fatal bacterial cell wall damage.
Gram-negative cell walls
In gram-negative bacteria, the peptidoglycan layer is made up of one to three layers of glycan
sheets and is only 3- to 7-nm thick. There are no teichoic acids, and crosslinkage between glycan
strands occurs directly between tetrapeptide side chains (Figure 4.4b).
The gram-negative cell wall includes an outer membrane (Figure 4.5b), a unique lipid bilayer
anchored to the peptidoglycan by lipoproteins. The outer layer is mostly composed of
a lipopolysaccharide (LPS) and the inner layer is composed of phospholipids. The LPS has two
basic components—the external o-oligosaccharide and the membrane-bound lipid A. The o-
oligosaccharide is anchored to a core polysaccharide that attaches to the lipid A. It extends
beyond the outer membrane and its exact sugar composition can be used to distinguish different
gram-negative bacterial species. The o-oligosaccharide is functionally similar to teichoic acid.
The lipopolysaccharide is toxic and is called the endotoxin. It is released upon cell lysis and is an
important determinant of pathogenicity.
The outer membrane has embedded porin proteins that allow small metabolites, such as sugars,
amino acids, and nucleotides, to pass through the outer membrane (Figure 4.5b). Without porins,
these metabolites would have to be actively transported across both the plasma membrane and
the outer membrane, doubling the cell's energy costs to bring nutrients into the cell.
The outer membrane also pinches off small microvesicles that contain autolysin, an enzyme that
degrades the peptidoglycan of unrelated bacteria. The enzyme is also released when nutrients are
scarce. Competing bacteria are destroyed because they can't repair their cell walls rapidly when
nutrients are limited, and the gram-negative bacteria can use nutrients released from the killed
cells for survival. Autolysin also plays a role in regulating bacterial growth and consequently is a
current target of antibiotic research.
The thin peptidoglycan layer of gram-negative bacteria is located between the outer membrane
and plasma membrane in the periplasmic space (Figure 4.5b). This region is about 10-nm thick
and contains the periplasm, a gellike material composed of proteins that bind amino acids,
sugars, vitamins, and ions as well as components needed for cell wall synthesis and enzymes that
detoxify chemicals.
Differences in cell structure make gram-negative cells more susceptible to lysis. Gram-negative
bacteria can withstand about three atmospheres of pressure. The thicker layer of peptidoglycan
found in gram-positive cell walls can withstand about 25 atmospheres of pressure before
rupturing (Figure 4.3). Although their peptidoglycan layer is thinner, gram-negative bacteria are
well protected by the selective permeability of their outer membrane. Any molecules too large to
move through porins are excluded from entering the bacterium by the barrierlike outer
membrane. This is clinically significant because gram-negative bacteria are resistant to large
antibiotics such as vancomycin that cannot pass through the porins.
Atypical Cell Walls
Although most bacteria have either a gram-positive or gram-negative cell wall, several groups
have diverged during evolution and now differ significantly from their gram-positive ancestors.
Bacteria with atypical cell walls include the acid-fast bacteria and the wall-less bacteria.
Acid-fast bacteria
Acid-fast bacteria have a cell wall made of a thick layer of peptidoglycan similar to gram-
positive cells. However, the peptidoglycan is linked to disaccharides that are attached to mycolic
acids, which are long-chain fatty acids. The sugar/mycolic acid layer is then overlaid with a
hydrophobic layer high in lipids. Porins are required to transport small hydrophilic molecules
through the acid-fast cell wall. The combination of mycolic acids and lipids make the bacteria
impermeable to many chemicals; consequently, the cells hold fast to the primary dye used in the
acid-fast staining procedure (see Remember This!). The cell wall composition also makes these
bacteria highly resistant to many antibiotics and disinfectants.
REMEMBER THIS!
The acid-fast stain is used to identify acid-fast bacteria from non-acid-fast bacteria. If necessary,
review the acid-fast staining procedure from Section 3.3 before continuing.
Wall-less bacteria
Wall-less bacteria include the Ureaplasmas, which cause urogenital infections, and
the Mycoplasmas, which cause walking pneumonia (see the Case Study). Because these bacteria
lack a cell wall, they can only survive in isotonic environments (ones with solute concentrations
the same as that of their cytoplasm) or they risk osmotic lysis. By living inside the cells of the
organisms they infect, the wall-less bacteria receive the protection normally provided by a cell
wall. Because they are wall-less, these bacteria have undefined shapes. Most are shaped like
cocci, but there are elongated and irregular forms as well. With a diameter of approximately 300
nm, the wall-less bacteria are also some of the smallest bacteria. They also have very small
genomes, with some consisting of only 500 genes—too small to actually code for all the
enzymes needed for life. As a result, these organisms must acquire some of their enzymes and
nutrients from their host.
CASESTUDYA Walking Pneumonia Outbreak at a

University
Working as an epidemiologist with the Georgia Department of Public Health, Janet completed
her final assessment of the walking pneumonia outbreak that occurred during fall semester 2012
at the Georgia Institute of Technology (Figure a). She was relieved that the largest pneumonia
outbreak at a U.S. university in 35 years was finally over.
a. The number of Mycoplasma pneumoniae cases among students at Georgia Institute of
Technology in 2012
(Data from: Mycoplasma pneumoniae Outbreak at a University — Georgia, 2012. (2013, August
2). Retrieved July 10, 2015, from
http://www.cdc.gov/mmwr/preview/mmwrhtml/mm6230a2.htm#Fig.)
1.
How long did the outbreak last?
2.
How many total students were diagnosed?
When the physician at the university health center called Janet in October reporting a sudden
increase in students presenting with sore throat, fever, headache, fatigue, and a dry cough that
sometimes occurred as violent spasms, she immediately suspected the infectious culprit to
be Mycoplasma pneumoniae (Figure b). She knew this extremely small, wall-less bacterium was
easily transmitted via respiratory droplets from coughs and sneezes by people living and working
in crowded places, like a university. This mild form of pneumonia occurs most often in people
younger than age 35 and is sometimes called walking pneumonia because patients are usually
still able to function.
b. An electron micrograph of Mycoplasma pneumoniae cells
3.
Why do the Mycoplasma bacteria shown have such unusual cell shapes?
4.
Predict the effectiveness of treating a Mycoplasma infection with penicillin, an antibiotic that
inhibits cell wall synthesis.
Janet collaborated with the university health center staff to test respiratory samples from the
affected students and quickly confirmed M. pneumoniae as the causative agent. To curtail the
outbreak, Janet and school administrators implemented an outreach campaign to alert the
university community and educate them on healthy practices to minimize pathogen transmission
in the college setting.
INVESTIGATE:
5.
Should Janet's prevention plan include administering a vaccine against walking pneumonia?
Using a survey to assess the effectiveness of their outreach campaign, Janet was pleased that
79% of the students who were aware of the outbreak reported following her recommendations to
reduce infection spread. She was frustrated, however, to discover that 54% of the campus was
completely unaware of the outbreak, putting them at high risk both for acquiring the infection
and for serving as a pathogen reservoir that could extend the outbreak.
6.
What methods of communication do you think would be the most effective means of reaching
college students with this vital information?
Think critically and try the interactive case study related to pathogens discussed in this chapter.
Ch 04: Interactive Case Study
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4.4 External Structures of Bacterial Cells

LEARNING OBJECTIVES
 1.Describe the structure and functions of the glycocalyx and capsule.
 2.Describe the structure and functions of fimbriae and pili.
 3.Explain the structure and functions of flagella.
As bacteria evolved, natural selection produced variants that were better adapted for survival in
new and changing environments. Over time, specialized structures evolved that provided
selective advantages to new species of bacteria. Specialized structures located outside the plasma
membrane and cell wall interact directly with the environment. These structures have functions
that expand a bacterium's potential habitat or enable it to survive during unfavorable
environmental conditions. They include the glycocalyx, fimbriae and pili, and flagella (Figure
4.6).
External structures of a bacterial cell• Figure 4.6The external structures of bacterial cells
include the glycocalyx, pili and fimbriae, and flagella. The glycocalyx protects the cell, whereas
the pili and fimbriae help in adherence, and flagella contribute to motility.
Animation: Prokaryotic Cell Structure
Ask Yourself
Between the pilus and the flagellum, which structure is not anchored to the plasma membrane?
the pilus
The Glycocalyx
The bacterial glycocalyx generally consists of a network of simple polysaccharide chains layered
just outside the cell wall, making it the outermost coating for those bacteria that possess it. If the
glycocalyx is gellike and firmly attached to the cell, it is known as a capsule. If it is more fluid
and loosely attached, it is known as a slime layer. These structures can protect a cell from
desiccation or enable it to colonize a new niche. For example, Streptococcus mutans synthesizes
a sticky capsule that enables it to adhere to the enamel surface of a tooth. Because cells of S.
mutans and other bacterial species attach to the teeth, they form dental plaque, an oral biofilm, or
microbial community. Synthesis of the bacterial capsule requires the breakdown of the sugar
sucrose. Sucrose can also be fermented to acid that can damage the enamel and cause tooth
decay.
Capsules also a allow bacteria to avoid phagocytosis. Different variants of S. pneumoniae that
can synthesize a capsule are pathogenic, but those that are unable to make a capsule do not cause
disease.
Fimbriae and Pili
Many bacterial species have hairlike structures composed of the protein pilin that extend from
the surface, called fimbriae and pili. A fimbria (plural fimbriae) is a straight, stiff, short filament
that is 6 to 7 nm in diameter (Figure 4.7). Fimbriae are numerous, with approximately 100 to
400 per cell and function in bacterial attachment to surfaces. They can be found on both gram-
positive and gram-negative bacteria.
A comparison of fimbriae and pili• Figure 4.7Most bacteria possess numerous fimbriae that
encourage their adhesion to various surfaces, including host cells, which facilitates infection. The
longer conjugative pilus joins two Escherichia coli cells allowing the transfer of genetic material.
Ask Yourself
The multiple hairlike extensions of the bacterial cell used for attachment are fimbriae , and the
long extension used for the transfer of genetic material is a pilus .
A pilus (plural pili) is a longer, thicker appendage found on the surface of all gram-negative
bacteria and on a few gram-positive bacteria. Some pili are essential for attaching bacterial
pathogens to host cells and initiating infection. For example, N. gonorrhoeae attaches to the cells
of the mucosal lining of the reproductive tract with proteins found on the tips of the pili that bind
to receptors on a host cell. As a result, the pathogen is not washed away by vaginal secretions or
urination. A conjugative pilus is longer and cells have fewer of them (approximately one to six
per cell) than common pili. A conjugative pilus forms a fragile hollow tube that can connect to
another bacterium of the same or a closely related species (Figure 4.7). This allows the transfer
of genetic material from the pilus-forming cell to the recipient cell. Still other pili function in
motility. The tip of a pilus adheres to a surface then shortens, causing the cell to be dragged
forward. As the process is repeated, the cell advances in an irregular manner called twitching
motility.
Flagella
Motility is an important evolutionary adaptation for most bacteria. The ability to move enables a
bacterium to potentially find a new microenvironment with additional resources for growth. For
pathogenic bacteria, motility allows migration to sites where they can more easily avoid host
defenses.
Flagella are the most common bacterial structure for motility. A flagellum (plural flagella) is a
stiff, helical, protein filament that rotates to propel a bacterium through its liquid environment.
The numbers and arrangements of flagella differ with the species (Table 4.2).
Interactive Graphic: Table 4.2
A flagellum is made up of three parts: the filament, a hook, and a basal body (Figure 4.8a). The
long, hollow, helical filament made of the protein flagellin is 5 to 10 μm long and attached to a
hook, or flexible connector. The hook anchors the flagellum to the basal body, a structure
embedded in the cell wall and plasma membrane. It consists of a rod attached to four rings in a
gram-negative bacterium and two rings in a gram-positive bacterium. The basal body is the
molecular motor that rotates the filament at a rate of 300 revolutions/second using energy
derived from the diffusion of a hydrogen ion gradient.
The structure and function of bacterial flagella• Figure 4.8Most bacteria possess one or more
hairlike projections, or flagella, that permit locomotion.
a. Flagellar structure
The long flagellar filament attaches to the curved hook region and rotates rapidly when the rings
and rod of the basal body spin, using energy released from the diffusion of a hydrogen ion
gradient.
b. Negative chemotaxis
This peritrichous bacterium coordinates the rotation of its flagella to run (swim) away from
repellent chemicals. Despite periodic tumbles as the flagella temporarily rotate in the opposite
direction, the net result is movement away from a harmful substance.
Think Critically
1.
If the bacterial cell in part b had encountered an attractant chemical like galactose rather than a
toxin, how would its swim pattern differ?
The bacterium would run and tumble toward the galactose.

2.
How would its flagellar movement differ?
The flagella would spin in the opposite direction.

Bacterial movements in response to specific stimuli are called taxes (singular taxis). Some
bacteria exhibit chemotaxis, a response to a chemical gradient, or phototaxis, a response to
variable light intensity. In chemotaxis, receptors on the bacterial surface bind to the chemical,
which initiates a chain of chemical reactions that affect the direction of the flagellar rotation.
When the flagella rotate counterclockwise, bacteria with polar flagella and peritrichous flagella
run, or move forward. When the flagella rotate clockwise, bacteria with polar flagella move in
reverse, and those with peritrichous flagella tumble in random directions.
Binding attractant chemicals such as galactose or oxygen send internal signals favoring
counterclockwise rotation and chemotactic progress. As the bacterium gets closer to the
attractant chemical source, it has longer run times and fewer tumbles. The binding of repellant
chemicals, such as toxins, triggers clockwise rotation and runs away from the chemical. Run
times decrease and tumbles are more frequent as the bacterium successfully avoids the repellent
(Figure 4.8b). Because both chemotactic and phototactic responses are composed of a series of
runs and tumbles, there are small random movements, but the net movement is directional.
In one group of bacteria, the spirochetes, two flagella are assembled within the periplasmic space
between the outer membrane and the plasma membrane. These two internal flagella,
termed axial filaments, are anchored at the end of the cell by a basal body. As the axial
filaments rotate, they cause the entire cell to move forward with a corkscrewlike motion. This
enables spirochetes to move through viscous mucous that impairs the movement of externally
flagellated bacteria.
The presence of axial filaments or standard flagella correlates with increased bacterial virulence
because these motile pathogens are better able to invade host tissues. Because flagellated
microbes are often clinically significant, microbiologists can use their presence, number, and
arrangement for bacterial identification (see The Microbiologist's Toolbox), the first step in
determining antimicrobial therapy.
THE MICROBIOLOGIST'S TOOLBOXThe Flagella Stain
Although their flagella can be several times the length of a bacterial cell, they are too thin (15- to
20-nm thick) to be seen with a light microscope. Flagella must be coated with special stains to
increase their diameter or they must be viewed with an electron microscope.
Simple staining procedures use a mordant to form an insoluble compound that binds to the
flagella, increasing their diameter and giving them color. The thickened and stained flagella can
then be viewed with a light microscope (see the Figure).A light micrograph showing clearly
stained flagella of Salmonella spp.
This staining technique is a useful tool for the microbiologist because it allows analysis of
flagellar arrangement. This is clinically significant because flagellar arrangement can be used for
pathogen identification and correlates with the microbe's ability to spread within the host.
Put It Together
Review Table 4.2, and answer this question.
The special staining of this bacterial specimen demonstrates a peritrichous flagellar
arrangement.
Ch 04: Microbiologist Toolbox Video
4.5 Internal Structures of Bacterial Cells

LEARNING OBJECTIVES
 1.Describe the plasma membrane.
 2.Explain the functions of the nucleoid.
 3.Describe the ribosomes.
 4.Describe the plasmids, inclusion bodies, and other internal organelles of bacterial cells.
 5.Describe the endospores.
The interior of bacterial cells is filled by the fluid cytoplasm. The cytoplasm is the reservoir of
all the cell's water-soluble metabolites and the location of the cell's internal structures. It is a
mixture of soluble inorganic ions, metabolic precursors, and macromolecules. Metabolites
diffuse through the cytoplasm to carry out the chemical processes of life. Several filamentous
cytoskeletal proteins add to the viscosity of the cytoplasm. These proteins function in
determining cell shape and positioning internal structures. The chemical makeup of the
cytoplasm also determines cellular pH and the osmotic pressure a cell experiences in its
environment. The cytoplasm is viscous enough that the rate of diffusion of a protein is about 10
times slower in the cytoplasm than in water. This reduced rate of diffusion is probably a
significant factor in limiting the size of prokaryotic cells.
The most significant internal structures of bacterial cells include the plasma membrane, nucleoid,
and ribosomes. Although prokaryotic cells are very small, they also contain a variety of other
specialized structures, many of which adapt the microbe to a specific habitat.
The Plasma Membrane
All living cells have a plasma membrane, also called a cell membrane, that separates the cell's
internal components from its environment. This membrane is a fluid phospholipid bilayer with
embedded proteins. The plasma membranes of bacterial cells differ slightly in structure from
those of eukaryotic cells. They contain different steroids to promote fluidity and have a
significantly higher proportion of proteins that participate in enzymatic reactions, signaling, and
transport, shown in Figure 4.9). Overall membrane function, however, is the same. It functions
to regulate the passage of material into and out of the cell. The phospholipid bilayer serves as a
general permeability barrier and traps ions, polar metabolites, and macromolecules inside the
cell, but allows free diffusion of small hydrophobic molecules such as methane or ethylene.
O2 and CO2 are small and nonpolar and therefore diffuse freely across the membrane. Water is
small but polar. Aquaporins, a special class of proteins, form small pores in the membrane that
aid in the diffusion of water through the plasma membrane. Other small molecules can be
excluded from moving across the membrane. For example, because hydrogen ions are charged,
they cannot diffuse through the bilayer and a hydrogen ion gradient can exist across the
membrane. Because hydrogen ions carry a charge, this chemical gradient also produces an
electrical gradient. The electrical gradient across the membrane can store energy similar to a
battery.
PROCESS DIAGRAM
Transport across the bacterial plasma membrane• Figure 4.9The hydrophobic tails of the
fatty acids of the phospholipid bilayer serve as a barrier to the diffusion of small polar
metabolites across the plasma membrane. Steroids help to provide fluidity to the membrane.
Integral membrane proteins regulate the passage of some substances across the phospholipid
bilayer.
Think Critically
Why are the steroids oriented as they are pictured in the membrane?
The polar -OH groups interact with the water, and the nonpolar steroid rings interact with the
hydrocarbons.
Passive transport is diffusion of small, nonpolar molecules across the membrane down their
concentration gradients. To diffuse polar molecules across the membrane requires assistance
from transmembrane channels, or permeases, and is known as facilitated diffusion. This
spontaneous transport process allows many extracellular molecules that the cell could use as
energy sources or as building blocks to diffuse across the plasma membrane at rates fast enough
for cell survival and growth. When metabolites must move against their concentration gradients
to enter or leave the cell, active transport processes are used. Active transport is an energy-
requiring process that couples the movement of molecules or ions across the membrane with the
hydrolysis of adenosine triphosphate (ATP) or the flow of another molecule down its
concentration or electrical gradient.
In addition to transport proteins, the plasma membrane also contains important enzymes and
enzyme complexes. These are responsible for running the biochemical reactions essential for
bacterial survival. Other plasma membrane proteins serve as receptors, binding the pathogen to
the surface of a host cell or participating in cellular communication. When proteins extend
completely through the phospholipid bilayer, they are known as integral membrane proteins.
Peripheral membrane proteins are associated with only one side of the phospholipid bilayer.
The Nucleoid
In prokaryotes, the nucleoid is the region that contains the chromosome, which carries the cell's
genetic material. Nearly all bacteria have a single circular bacterial chromosome that contains
their genetic information. It ranges from about 0.5 to 10 million base pairs in length, making the
DNA 100X to 1000X longer than the cell itself. To fit, the thin DNA is in a highly folded and
twisted state. Because the DNA is so condensed, transcription of the DNA into RNA only
occurs at the outer surface of the nucleoid. Environmental factors and even the process of
transcription lead to constant twisting and supercoiling of the bacterial chromosome. This
exposes different genes at the surface of the nucleoid, where they have an opportunity to be
transcribed.
Chromosomes need about 800 genes to synthesize essential enzymes and produce the structural
components of the cell. However, bacterial chromosomes have thousands of genes, allowing
bacteria to adapt to changes in their environment. For a soil bacterium, there are significant daily
and seasonal changes in temperature, pH, osmolarity, and nutrient sources. All of these different
conditions require changes in metabolism and gene regulation to optimize growth and
competition with other soil microbes. Consequently, the cell needs many more genes than just
those essential for survival in a single environment.
Ribosomes
Ribosomes are the smallest functional structures in bacterial cells, measuring only 70 Svedberg
units (70S) in size. A Svedberg unit (S) is a measure of structure size based on its rate of travel in
a tube subjected to high centrifugal force, therefore considering both mass and shape so the units
are not additive. Consequently, the two parts of the ribosome, the small 30S subunit and the large
50S subunit, come together to make a full-sized, but different-shaped, 70S ribosome (Figure
4.10a).
Bacterial ribosomes• Figure 4.10Bacterial ribosomes are found in the cytoplasm, where they
use information in the genetic sequences in messenger RNA (mRNA) to code for the amino acid
sequences of proteins.
a. A bacterial ribosome is composed mostly of rRNA with many smaller r-proteins embedded in
or attached to the surface.

b. This simplified diagram of the actual structure of the ribosome shown in Figure a shows the
general shape of a 70S ribosome bound to mRNA and synthesizing a protein.
Ask Yourself
This figure does not show a necessary component of protein synthesis. This component is
the ________.
 a.
rRNA
 b.
riboproteins
 c.
tRNA
 d.
amino acids
 d.
amino acids
Bacterial ribosomes have about 50 different ribosomal proteins (r-proteins) in their two subunits,
which are assembled on a ribosomal RNA (rRNA) scaffolding. There are three rRNAs (16S
rRNA, 23S rRNA, and 5S rRNA), which have enzymatic activity and catalyze the assembly of
amino acids into proteins by the process of translation (Figure 4.10b). The r-proteins stabilize
the rRNA structure, unwind the messenger RNA (mRNA), and provide docking sites for
regulatory proteins.
In addition to the mRNA and ribosomes, transfer RNAs and other translational protein factors
are required. Protein synthesis is a major cell process and determines how fast it can grow. In
rapidly growing cultures of Escherichia coli, there are 200,000 ribosomes per cell and protein
synthesis consumes about 90% of the ATP produced by the cell.
Plasmids, Inclusion Bodies, and Membranous Structures
In addition to the chromosome, many bacteria carry plasmids, which are extrachromosomal
genetic elements. Plasmids range from one to hundreds of genes and from one to hundreds per
cell. The genes on plasmids generally give the bacteria a selective advantage in unusual
environments. For example, it is common to find plasmids in Pseudomonas bacteria that have
been isolated from soil polluted from an oil spill. The proteins produced by the plasmids help the
bacteria break down petroleum products. Other soil bacteria carry antibiotic resistance genes that
give them a survival advantage in their communities. Plasmids with such genes are common in
health care systems. Plasmids are also key tools in molecular biology and gene cloning.
Inclusion bodies
Inclusion bodies are generally small cytoplasmic structures whose composition and function
varies with bacterial species (Table 4.3). Often they act as a molecular stockpile, storing
substances needed by the cell (Figure 4.11a). Inorganic chemicals such as calcium, sulfur, or
phosphate can be stored as cystoplasmic granules in the cytoplasm. Organic polymers such as
glycogen serve as energy-storage compounds because they can be degraded for ATP production.
Bacterial inclusion bodiesTable4.3
General function Inclusion Specific use
Storage of carbon/energy polymers PHA (poly-ß- Serves as carbohydrate source for synthes
hydroxyalkanoate)
Glycogen Serves as carbohydrate source for synthes
Storage of molecules needed for Polyphosphate granules Provides phosphorus used in synthesis of
biosynthesis acids
Sulfur granules Required for the synthesis of some amino
Storage of enzymes Carboxysome Stores rubisco, which acts in CO2 fixation
Enterosome Uses enzymes to degrade toxic compound
Trapping gas Gas vesicle Confers buoyancy for aquatic cyanobacte
Magnetotaxis Magnetosome Directs swimming by interaction with the
attachment to the appropriate substrate
Photosynthesis Thylakoid Converts CO2 into glucose
Inclusion bodies• Figure 4.11Inclusion bodies are a diverse group of prokaryotic intracellular
structures.
a. Carboxysomes store the principle enzyme needed to initiate photosynthetic reactions.
b. Thylakoids are stacked, flattened membranous sacs with the appropriate proteins embedded to
run photosynthesis.
Put It Together
Review Table 4.3, and answer this question.
What type of inclusion body buoys cyanobacteria up to receive sufficient light for initiation of
photosynthesis in thylakoids and use of CO2-fixing enzymes in the carboxysomes?
gas vesicles
Inclusion bodies have many other functions. Magnetosomes allow bacteria to respond to Earth's
magnetic field and swim downward until encountering an appropriate substrate for
attachment. Gas vesicles provide buoyancy so aquatic photosynthetic bacteria float in the water
column at a depth where they receive optimal light intensity. Although bacteria typically lack
membrane-bound internal structures, some photosynthetic bacteria that have evolved simple
membranous compartments to localize chemical reactions and increase the surface area of
membrane-based reaction centers (Figure 4.11b). These structures, called thylakoids, organize
the enzymes and electron transport proteins needed for the light reactions of photosynthesis.
Similar internal membrane structures are found in bacteria that obtain their energy from the
oxidation of inorganic chemicals, such as hydrogen sulfide or reduced iron, rather than from
light.
Endospores
Several genera of bacteria can produce dormant cells known as endospores that are highly
resistant to harsh environmental conditions. Not only can endospores survive temperature
extremes, they are unaffected by exposure to ultraviolet radiation, toxins, antibiotics, and
desiccating conditions.
Bacillus and Clostridium are the most clinically significant genera of endospore forming
bacteria. Different species of these microbes are responsible for causing infections such as
anthrax, food poisoning, gas gangrene, botulism, and tetanus. Because their thick protective
walls are resistant to disinfectants and antibiotics, it is critical to prevent endospore
contamination in medical settings (see the Clinical Application). All surgical instruments and
indwelling devices must be sterilized by heat plus pressure in an autoclave or by treatment with
gamma radiation. Only drastic practices such as these are able to damage the endospore wall,
allowing destruction of the cell within. Alcohol-based gels are ineffective at decontaminating the
hands of health care providers, who are encouraged to wash with soap and water to physically
remove endospores.
Clinical ApplicationEndospore-forming Bacteria
Although not many pathogenic bacteria form endospores, those that do are very difficult to treat
because they are only susceptible to antibiotics when actively growing as vegetative cells. One
endospore-forming pathogen, Clostridium difficile (Figure a), is a common hospital-acquired
pathogen. It doesn't cause disease in healthy individuals, but if the normal bacteria in the large
intestine are killed by long-term, broad-spectrum antibiotic therapy, C. difficile is still able to
survive and grow. As a result, it can cause pseudomembranous colitis (Figure b), a type of
bloody diarrhea that is very difficult to treat. Although antibiotics can work, therapy is long,
expensive, and recurrence rates are high (Figure c). The most successful treatment strategies are
those that restore the normal intestinal microbiota.a. Clostridium difficile, a common hospital-
acquired pathogen, forms endospores. b.

Pseudomembranous colitis, or significant damage to the intestinal mucosa, occurs when C.
difficile endospores germinate and thrive in the gut.
c. Percent failure and/or recurrence of infection

after vancomycin treatment
(Data from: Aslam, S., Hamill, R., & Musher, D. (n.d.). Treatment of C. difficile-associated
disease: Old therapies and new strategies. The Lancet Infectious Diseases, 549-557.)
Interpret the Data
1.
What year was the lowest failure and/or recurrence rate for treating Clostridium difficile-
associated diarrhea?
1994
2.
What year was the highest rate?
1985
Under favorable conditions, bacterial cells grow and reproduce by binary fission (Figure 4.12a).
However, when normal vegetative bacterial cells encounter a harsh environment, metabolically
inactive endospores form. The process of sporulation, which takes about 6 to 8 hours, begins
with the invagination of the plasma membrane (Figure 4.12b). Known as a spore septum, this
membrane ingrowth encloses a small amount of cytoplasm, a newly replicated nucleoid,
ribosomes, RNA, and enzymes. Layers of peptidoglycan are deposited between the double layer
of the spore septum, forming a resistant cortex. Next, a thick, protective protein coat forms
outside of the peptidoglycan layers to produce the spore wall. Inside the developing endospore,
calcium dipicolinate dehydrates the cell. Because DNA and proteins are only vulnerable when in
solution, this practice protects these enclosed molecules when the endospore is exposed to
extreme heat. The mature endospore is released when the original cell lyses.
PROCESS DIAGRAM
Endospore formation• Figure 4.12In bacteria, vegetative cells reproduce by binary fission.
Some bacterial species can also undergo sporulation.
Ask Yourself
In the first step of bacterial sporulation, the ________.
 a.
endospore is released from the cell
 b.
DNA is replicated
 c.
chromosome is surrounded by a tough cortex and spore coat
 d.
plasma membrane encompasses the DNA
 b.
DNA is replicated
When the endospore encounters water and other favorable conditions, such as suitable
temperatures and appropriate nutrients, it germinates. The enzymes within the endospore degrade
the cortex and spore coat, allowing the uptake of water and nutrients. It takes about 1.5 hours to
revive the dormant cell, which then grows and reproduces by the vegetative cycle (Figure
4.12a).
Because one endospore develops into just one new vegetative cell, sporulation is not a
reproductive process, but a highly effective survival mechanism. In fact, endospores are so
resistant they can be freeze-dried and dispersed by the explosion of a bomb and still remain
infectious. Endospores of Bacillus anthracis, which causes a lethal form of anthrax when the
spores are inhaled, have been turned into a bioweapon. Endospores have been germinated into
growing cells after being frozen in ice for 10,000 years and after being fossilized in amber for 25
million years. Endospores are so heat resistant that they can be boiled without being killed. As a
result, food products must be canned under high temperatures and pressures to kill endospores
and prevent foodborne illness.
3D Animation: What structures are found inside of bacteria?
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4.6 Prokaryotic Evolution and Classification

LEARNING OBJECTIVES
 1.Explain the early evolution of the three domains of life.
 2.Describe how bacteria and archaea are classified.
No one knows whether, billions of years ago, life evolved on Earth once or multiple times, but
by analyzing the many characteristics shared by all living things, it appears that all life
descended from a common ancestral cell. This ancestor had a plasma membrane that regulated
the passage of materials into and out of the cell and nucleic acids for genetic information. It also
used ATP as an energy source and had evolved basic metabolic pathways probably similar to
those that we see in cells today. Evolution has given rise to millions of different prokaryotic
organisms. To understand how these living organisms relate to each other, it is necessary to
classify them into groups based on their structures and biochemical makeup. In this section, you
will discover theories on how life evolved and how the early tree of life split into the major
groups of prokaryotic organisms, along with current schemes used for classifying bacteria.
The Tree of Life
Over the past 3.8 billion years, countless diverse organisms have evolved, including millions of
microbial species. The sequence of nucleotides in DNA holds a wealth of information about an
organism's evolutionary history. A comparison of the nucleotide sequences of different
organisms can reveal the evolutionary connections between them.
The nucleotide sequence in the gene that codes for the 16S rRNA of the small ribosomal subunit
has been used to establish the evolutionary relationships between all prokaryotic organisms.
Some regions within this gene evolve very slowly and maintain sufficient similarity to make
comparisons between the most distantly related organisms. Other gene regions change relatively
rapidly so even closely related organisms show genetic differences. Comparison of 16S rRNA
gene sequences demonstrated that prokaryotes include two distinct lineages—the Bacteria and
the Archaea. The third primary lineage of living things, the Eukarya, diverged from the
Archaea and includes all eukaryotic organisms. The Bacteria, Archaea, and Eukarya are
the domains of life—the three main branches on the evolutionary tree of life proposed by Carl
Woese (Figure 4.13a). Today, microbial organisms, especially those that can't be grown in the
lab, are routinely classified by their 16S rRNA sequence.
Bacterial classification• Figure 4.13Because bacterial classification is especially challenging,
multiple organization schemes are used to highlight the evolutionary relationships between
organisms, providing valuable information to microbiologists.
a. The tree of life
Because all living things share certain fundamental features, there was most likely one common
ancestor of all modern living organisms. On the evolutionary tree, organisms that are distantly
related to one another are separated by long distances; closely related organisms are placed on
short branches with adjacent or common nodes. Here, related organisms are shown in the same
color.
b. Taxonomic classification of bacteria based on genetic comparisons of 16S rRNA gene

sequences
Phylum Genus name General properties
Actinobacteria Mycobacterium Acid-fast pathogen
Propionibacterium Anaerobic propionic acid producer
Streptomyces Antibiotic-producing filamentous bacteria
Aquificae Aquiflex Hyperthermophilic, chemoautotrophic bacteria
Bacteroidetes Bacteroides Obligate anaerobe; significant component of intestinal flora
Cytophaga Gliding locomotion
Chlamydiae Chlamydia Intracellular pathogen
Phylum Genus name General properties
Chlorobi Chlorobium Green sulfur bacteria perform anoxygenic photosynthesis
Chloroflexi Chloroflexus Green nonsulfur bacteria that perform anoxygenic photosynthesis
Cyanobacteria Synechococcus Widespread, abundant blue-green alga that performs oxygenic pho
Deinococcus- Deinococcus Highly resistant to radiation and/or heat
Thermus
Firmicutes Lactobacillus Lactic acid producer; component of normal vaginal microbiota
Staphylococcus Component of normal skin microbiota; may be pathogenic
Clostridium Anaerobic, endospore-forming pathogen
Planctomycetes Planctomyces Possess a holdfast to facilitate attachment following budding; lack
in cell walls
Proteobacteria Bdellovibrio Predatory bacteria
Pseudomonas Opportunistic pathogen
Escherichia Dominant component of intestinal flora
Myxococcus Soil microbe that forms swarming aggregations
Agrobacterium Cause of tumor-like growth in plants
Rickettsia Intracellular pathogen
Neisseria Pathogenic and commensal diplococci
Spirochetes Treponema Spirochetes motile by axial filaments
Leptonema Aquatic and soil microbe
Tenericutes Mycoplasma Gram positive, low guanine + cytosine
Thermotogae Thermatoga Hyperthermophilic, chemoautotrophic bacteria
Ask Yourself
Which of the following groups are most closely related?
 a.
flagellates and spirochetes
 b.
animals and methanogens
 c.
ciliates and slime molds
 d.
cyanobacteria and extreme halophiles
 c.
ciliates and slime molds
A traditional classification scheme based on phylum, class, order, family, genus, and species
(Figure 4.13b) was integrated with the tree of life strategy by nonclinical microbiologists. This
allows DNA sequence analysis to determine phylogenetic relationships among different bacterial
species and their organization using a phylogenetic tree. This branching diagram depicts the
evolutionary relationships among various groups based on their implied descent from a common
ancestor. Organisms whose gene sequence indicates they shared a common ancestor are located
on the same branch of the tree. The length of the lines between nodes is proportional to the
evolutionary distance.
The Clinical Classification of Prokaryotes
Early classification schemes for prokaryotes were based primarily on cell shape and Gram-stain
reaction. This type of system is still used by clinical microbiologists because it allows rapid
pathogen identification for diagnosis (Figure 4.14). The species is the fundamental unit of
classification in biology. Its meaning is clearly defined for macroscopic plants, fungi, and
animals that reproduce sexually. For these organisms, a species is a group of actually or
potentially interbreeding populations. However, this definition fails for prokaryotes, which
reproduce asexually through binary fission rather than sexually. A prokaryotic species is more
accurately described as a group of genetically similar individuals that are adapted for life in
similar environments. An issue is how genetically similar different prokaryotes need to be to be
considered members of the same species. At present, it is generally accepted that members of a
bacterial species show a 70% or greater DNA relatedness as measured by DNA hybridization
studies and a less than 5% variation in the amount of G and C in their DNA. They also show a
clear phenotype distinction from other species.
Practical classification of bacteria for clinical use• Figure 4.14Bacteria can be grouped for
rapid pathogen identification for medical diagnoses using cell shape and arrangement, followed
by analysis of biochemical properties of the specimens.
Ask Yourself
Which of these bacteria is gram-negative and facultatively anaerobic—Bordetella or Bacillus?
Bordetella
Approximately 10,000 prokaryotic species have been identified, but microbiologists estimate 10
million species as a full count of total prokaryotic diversity. Because of this broad diversity, a
simple genus and species classification probably couldn't describe important genetic variants
within a species. Therefore, it is sometimes necessary to characterize a bacterial species by
recognizing multiple subspecies, or groups of individuals of a particular species that demonstrate
consistent, identifiable variations. A subspecies can be further subdivided into
different strains to highlight smaller, but important genetic differences. The taxonomic rules for
designating strains are not closely regulated. Serotypes are strains that can be identified by the
response of specific antibodies to bacterial surface molecules. For example, most species of E.
coli are harmless bacteria that live in the large intestine of animals. However, there are many
different strains and several different serotypes that are pathogenic. One such serotype, E.
coli 0157:H7, causes bloody diarrhea and has a mortality rate of 50% for young children.
Clearly, from an evolutionary prospective, prokaryotes are very successful organisms. They
demonstrate amazing diversity that allows them to colonize almost every habitat on the planet in
large numbers. Because of their ability to affect human well-being and environmental health,
these microbes will be the focus of many upcoming chapters.
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Summary
 4.1 The Prokaryote's Place in the Living World
o •Prokaryotes are small, unicellular organisms that lack a distinct membrane-
bound nucleus and complex membranous organelles. They have a single
circular chromosome. The cells of eukaryotes are larger and more complex,
with a membrane-bound nucleus containing multiple linear chromosomes and
complex membrane-bound organelles.
o •Nitrogen fixation and photosynthesis performed by prokaryotes sustain life
on Earth. Their activities are also essential for the biogeochemical cycles.
o •As eukaryotes evolved, they developed relationships with the prokaryotes
that lived on or in them. Such relationships are called symbiosis. Many of the
symbiotic relationships between humans and the prokaryotes are examples
of mutualism, in which both partners thrive. An example of mutualism is
when bacteria fix nitrogen in plants (see the
diagram). Commensalism and parasitism are two other types of symbiotic
relationships.Symbiotic relationships: Mutualism• Figure 4.1a
 4.2 Bacterial Cell Shapes and Arrangements

o •The size range for bacteria is usually 0.2 to 2.0 μm in diameter and 2 to 8 μm
in length.
o •Bacterial shape is determined by the cell wall structure. Common bacterial
shapes are the coccus (see the diagram), which is spherical; bacillus, which is
cylindrical; and spiral. Other forms may be club-shaped, filamentous, or
appendaged. Pleiomorphism describes the slight shape variations among
bacterial cells of the same species.Bacterial morphology• Figure 4.2b
o •When bacteria don't separate after undergoing binary fission, they form
groupings including pairs (described by the prefix diplo-), chains (strepto-),
and clusters (staphylo-). Some bacterial species form sets of four (tetrads) or
packets of eight (sarcinae). Microscopic determination of bacterial shape and
arrangement is often the first step in pathogen identification.
 4.3 The Bacterial Cell Wall
o •The bacterial cell wall prevents cell lysis by resisting pressure
from osmosis when the microbe is in a hypotonic environment.
o •Peptidoglycan is the primary structural component of bacterial cells.
Peptidoglycan is composed NAG and NAM joined together to form a
disaccharide.
o •Gram-positive bacteria have a thick layer of peptidoglycan with molecules
of teichoic acids weaving through it. Teichoic acids enable pathogenic
bacteria to attach to host tissues, regulate cell division, and avoid engulfment
by phagocytic cells.
o •The cell wall of gram-negative bacteria (see the diagram) consists of a thin
layer of peptidoglycan and an outer membrane. Porins in the outer membrane
allow free diffusion of small metabolites. The outer membrane
has lipopolysaccharides (LPS) on the outer surface; the lipid A portion of the
LPS is the endotoxin, whereas the o-oligosaccharide portion functions much
like teichoic acids.Bacterial cell wall structure: The gram-negative cell
wall• Figure 4.5

o •Acid-fast bacteria have a thick layer of peptidoglycan surrounded by a
sugar/mycolic acid layer that is overlaid with lipids. Acid-fast bacteria are
highly impermeable to most antibiotics and disinfectants. Wall-less bacteria
do not have a regular cell shape. They are the smallest and simplest self-
replicating bacteria.
 4.4 External Structures of Bacterial Cells
o •The glycocalyx and capsules are polysaccharide coatings firmly attached to a
bacterial cell that can enable some pathogenic bacteria to attach to tissues to
avoid phagocytosis.
o •Fimbriae and pili (see the micrograph) are used to attach to surfaces, for
twitching motility, or during conjugation.A comparison of fimbriae and
pili• Figure 4.7

o •Flagella and axial filaments are long protein polymers driven by basal
body motors that function in motility. Movement in response to a chemical
gradient is called chemotaxis. Movement in response to varying light
intensity is called phototaxis.
 4.5 Internal Structures of Bacterial Cells
o •The plasma membrane, which is found in all cells, controls the passage of
materials into and out of the cell, which is filled with cytoplasm. Materials
move into and out of the cell by passive transport, facilitated diffusion,
and active transport.
o •The nucleoid is the region of the bacterial cell that contains the genetic
material, usually a circular chromosome of DNA. The bacterial
chromosome codes for the proteins needed to build the structural components
of the cell and the enzymes needed for cell metabolism. The information in
DNA is copied into RNA by the process of transcription.
o •The ribosome is the smallest organelle of the cell. It is made of
approximately 50 r-proteins and three different rRNAs (see the photo).
Ribosomes are the sites of protein synthesis. The process by which the
information in RNA is used to synthesize proteins is
called translation.Bacterial ribosomes• Figure 4.10
o •Plasmids are small, circular self-replicating DNA molecules that carry
extrachromosomal genetic information in bacteria. They are important tools in
molecular biology and gene cloning. Inclusion bodies often store material for
the bacterial cell. They include glycogen and polyhydroxyalkanoate granules,
carboxysomes, enterosomes, and gas vesicles. A few groups of bacteria have
membrane-bound organelles including thylakoids used in photosynthesis,
and magnetosomes, which direct cells into sediments.
o •Endospores, which form by the process of sporulation, are highly resistant,
metabolically inactive cells produced by several genera of bacteria. Under
favorable conditions, endospores can germinate into growing vegetative cells.
 4.6 Prokaryotic Evolution and Classification
o •The sequences of 16S rRNA genes have been used to construct a tree of life
(see the diagram) to depict early evolutionary of connections between living
organisms.Bacterial classification• Figure 4.13
o •Members of the kingdom Bacteria are traditionally classified based on their

shapes, Gram-stain reactions, and physical features. More recent genetic
analysis has made it possible to classify bacteria according to their
phylogenetic relationships, allowing construction of a phylogenetic tree.
Phylogenetic trees have also been developed for the
kingdoms Archaea and Eukarya.
acid-fast
A staining method that identifies organisms containing mycolic acid in their cell walls.
active transport
Processes by which energy is used to move metabolites against their concentration gradient to
enter or leave the cell.
Archaea
A domain of prokaryotic organisms that do not have a membrane-bound nucleus, are genetically
distinct from bacteria, and can exploit extreme environments or produce methane.
axial filament
In spirochetes, internal flagella assembled within the periplasmic space between the outer
membrane and the plasma membrane.
bacillus
The cylandrical shape of some bacteria.

Bacteria
A domain of prokaryotic organisms that do not have a membrane-bound nucleus and inhabit
virtually all environments.
bacterial chromosome
In prokaryotes, the circular DNA-containing structure that holds the organism's genetic
information.
basal body
The molecular motor that rotates a flagellum to contribute to bacterial motility.

biogeochemical cycle
The circulation of chemical elements between the biotic and abiotic components of the
biosphere.
chemotaxis
Movement in response to a chemical stimulus.

commensalism
A type of symbiosis in which one species benefits and the other species is neither harmed nor
benefited.
conjugative pilus
A fragile hollow tube that can connect to another bacterium of the same or a closely related
species for the transfer of genetic material.
cytoplasm
The gellike solution of water and inorganic and organic substances that fills the cell; contains all
the internal cell structures.
endospore
A resistant asexual spore that develops inside some bacteria.

Eukarya
A domain of organisms that have cells with a membrane-bound nucleus and complex organelles.
eukaryote
An organism consisting of one or more cells containing a nucleus and complex membrane-bound
organelles, including plants, animals, fungi, and protists.
facilitated diffusion
The diffusion of polar molecules across the membrane of a bacterial cell with assistance from
permeases.
fimbria
(plural fimbriae) A straight, stiff, short filament that functions in bacterial attachment to surfaces.
flagellum
(plural flagella) (eukaryotic) An external hairlike structure composed of microtubules in a 9+2

arrangement that permits locomotion through wave-like movements.
gas vesicle
An inclusion body that traps gas and provides buoyancy to cyanobacteria.

glycocalyx
The extracellular carbohydrate and glycoprotein coating produced by some bacteria.

gram-negative
Bacteria that stain pink or red after Gram staining because of the thin layer of peptidoglycan in
their cell wall.
gram-positive
Bacteria that stain dark purple or violet after Gram staining because of the thick layer of
peptidoglycan in their cell wall.
inclusion body
A cytoplasmic aggregate that often consists of stored materials.

lipid A
The toxic lipid component of the lipopolysaccharide embedded in the outer layer of gram-
negative bacterial cell walls.
lipopolysaccharide (LPS)
A large molecule with its carbohydrate portion extending outwards and its lipid component
embedded in the outer membrane of gram-negative bacterial cell walls; it functions as an
endotoxin, illiciting strong host immune responses.
magnetosome
An inclusion body that allows bacterial swimming to be directed by magnetic field.

mutualism
A type of symbiosis in which both species benefit.

nitrogen fixation
The enzymatic conversion of atmospheric N2 into organic compounds.

o-oligosaccharide
The long carbohydrate component of lipopolysaccharide that extends away from the outer layer
of gram-negative bacterial cell walls.
passive transport
The diffusion of small, nonpolar molecules across the plasma membrane down their
concentration gradient.
peptidoglycan
A network of polysaccharide and peptide chains that make up the primary component of
bacterial cell walls.
periplasmic space
The region in the gram-negative bacterial cell wall that contains the periplasm, a gel-like
material composed of proteins that bind amino acids, sugars, vitamins, and ions, as well as
components needed for cell wall synthesis and enzymes that detoxify chemicals.
phototaxis
Movement in response to variable light intensity

phylogenetic tree
A branching diagram depicting the evolutionary relationships among organisms descended from
a common ancestor.
pilus
(plural pili) An appendage found on the surface of some bacteria that may serve in attaching
bacterial pathogens to host cells and initiating infection.
plasmid
A small, circular, DNA molecule that replicates independently of the bacterial chromosome.
prokaryote
Bacteria and archaea; a small, unicellular organism that typically lacks a distinct nucleus and
complex membrane-bound organelles.
ribosome
A small particulate structure composed of ribosomal RNA (rRNA) and proteins that is the site of
protein synthesis.
serotype
A group of microbes that share common surface antigens.

sporulation
The formation of metabolically inactive, physically protected spores.

subspecies
A group of individuals of a particular species that demonstrate consistent, identifiable variations

associated ecological or geographic distribution.
teichoic acid
A polymer composed of sugar-phosphate subunits that is the major carbohydrate component of a

gram-positive bacterial cell wall.
thylakoid
A bacterial inclusion body that organizes the enzymes and electron transport proteins needed for
the light reactions of photosynthesis; also found in plant cell chloroplasts.
transcription
The process by which the genetic information in DNA is copied into RNA; the RNA molecule is
synthesized using a DNA template.
translation
The process by which proteins are synthesized by ribosomes from information contained in
mRNA.
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Critical and Creative Thinking Questions
1.
Streptococcus mutans can initiate tooth decay. What type of symbiotic relationship would S.
mutans have in a person who brushes and flosses their teeth regularly?
As long as no decay occurred, it would have a commensal relationship.

2.
From an evolutionary perspective, would you expect ribosomal proteins or ribosomal RNA to
catalyze amino acid polymerization?
Ribosomal RNA would catalyze amino acid polymerization because it could act as both an
enzyme and as genetic information.
3.
Of the thousands of genes on the chromosome of most bacteria, why are only approximately 800
considered essential?
Those genes are all that are required to carry out the basic metabolism and maintain cell structure
in a stable environment.
4.
Why is regulating the osmotic pressure of the cytoplasm a critical activity for the cell?
An environment that is too hypotonic or hypertonic can be lethal to a cell.

5.
Explain how this diagram shows why both membrane proteins and energy are required for active
transport.
A protein is needed to shield polar molecules as they move through the membrane. Energy is
needed to move the molecules against their concentration gradient.
What is happening in this picture?
The photo shows a green mat that is actually a thick layer of photosynthetic bacteria submerged
beneath the surface of the water. The bubble coming off the bacterial mat contains oxygen
produced by photosynthesis, the same process that occurred on Earth about 3.6 billion years ago.
In the early history of Earth, photosynthetic bacteria contributed oxygen to the atmosphere,
making the evolution of eukaryotic life possible; today, the same bacteria help sustain that life.
Think Critically
What causes the pointed projections on the surface of the cyanobacterial mat?
Air bubbles rising through the mat push up projections on the mat surface.
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Self-Test
1.
Review What a Microbiologist Sees, and answer this question.
Relative to eukaryotic organisms, prokaryotes have the ________.
 a.
greatest numbers of individuals
 b.
greatest species diversity
 c.
most diverse habitats
 d.
greatest biomass
 e.
All of these are correct.
 e.
All of these are correct.
2.
Nitrogen fixation is the process by which ________.
 a.
plants break down NO2 and release O2
 b.
bacteria incorporate nitrogen from the air into organic compounds
 c.
plants synthesize organic compounds from nitrogen in the air
 d.
bacteria break down amino acids
 e.
plants break down amino acids
 b.
bacteria incorporate nitrogen from the air into organic compounds
3.
________ is a symbiotic relationship in which the host is ________.
 a.
Mutualism; harmed
 b.
Commensalism; not harmed
 c.
Parasitism; not harmed
 d.
Commensalism; benefitted
 e.
Parasitism; benefitted
 b.
Commensalism; not harmed
4.
The processes by which elements such as carbon and sulfur are moved between the living and
nonliving worlds are known as ________.
 a.
biogeochemical cycles
 b.
mutualism
 c.
symbiosis
 d.
commensalism
 e.
parasitism
 a.
biogeochemical cycles
5.
Review the Microbiology InSight, Figure 4.2, and answer this question.
The cell morphology and arrangement in which the bacteria look like a bunch of grapes
is ________.
 a.
spirochete
 b.
streptobacillus
 c.
staphylococcus
 d.
diplobacillus
 e.
coccobacillus
 c.
staphylococcus
6.
What type of environment would cause the cell change shown in the diagram?
 a.
isotonic
 b.
high salt concentration
 c.
hypertonic
 d.
high sugar concentration
 e.
hypotonic
 e.
hypotonic
7.
In this diagram of peptidoglycan, the aqua structures are ________ and the purple structures
are ________.
 a.
glycan chains; crossbridges
 b.
tetrapeptide side chains; glycan chains
 c.
N-acetylmuramic acid; crossbridges
 d.
glycan chains; teichoic acids
 e.
lipid A; glycan chains
 a.
glycan chains; crossbridges
8.
Teichoic acids in the bacterial cell wall ________.
 a.
are also known as exotoxin
 b.
regulate cation flow
 c.
prevent cell lysis
 d.
regulate gene expression
 e.
anchor flagella
 b.
regulate cation flow
9.
Which of the following is true of bacterial cell walls?
 a.
Gram-positive cell walls include an outer membrane.
 b.
Gram-negative cell walls include an inner wall zone.
 c.
Gram-positive cell walls are made up of one to three layers of glycan sheets.
 d.
Gram-positive cell walls contain lipoproteins.
 e.
Gram-negative cell walls contain lipopolysaccharide
 e.
Gram-negative cell walls contain lipopolysaccharide
10.
________ have a thick layer of peptidoglycan covered with a sugar/mycolic acid layer that is
overlaid with a hydrophobic layer high in lipids.
 a.
Acid-fast bacteria
 b.
Gram-positive bacteria
 c.
Gram-negative bacteria
 d.
Archaea
 e.
Wall-less bacteria
 a.
Acid-fast bacteria
11.
Flagella found within the cells of some bacteria are called ________.
 a.
conjugative pili
 b.
fimbriae
 c.
internal flagella
 d.
axial filaments
 e.
inclusion bodies
 d.
axial filaments
12.
In the diagram, the transport process numbered 1 is ________ and 2 is ________.
 a.
active transport; facilitated diffusion
 b.
facilitated diffusion; active transport
 c.
simple diffusion; facilitated diffusion
 d.
active transport; passive transport
 e.
facilitated diffusion; passive transport
 c.
simple diffusion; facilitated diffusion
13.
Protein synthesis is carried out by ________ and ________ at the ________.
 a.
DNA; fatty acids; ribosomes
 b.
RNA; lipopolysaccharides; nucleoid
 c.
plasmids; RNA; nucleoid
 d.
RNA; proteins; ribosomes
 e.
DNA; RNA; ribosomes
 d.
RNA; proteins; ribosomes
14.
Structures within bacterial cells that store nutrients, enzymes, or other needed materials
include ________.
 a.
endospores and gas vesicles
 b.
ribosomes and endospores
 c.
magnetosomes and permeases
 d.
glycan sheets and ribosomes
 e.
carboxysomes and enterosomes
 e.
carboxysomes and enterosomes
15.
Which bacterial structures are correctly paired with their functions?
 a.
flagella: form a bridge during conjugation
 b.
endospores: store nutrients for the cell
 c.
plasmids: synthesize proteins
 d.
capsules: anchor flagella
 e.
fimbriae: aid in attachment of the cell to surfaces
 e.
fimbriae: aid in attachment of the cell to surfaces
16.
Review The Microbiologist's Toolbox, and answer this question.
Bacterial flagella are ________ to be seen so they must be ________ and ________ to be
visualized.
 a.
too short; lengthened; colored
 b.
too thin; lengthened; stained
 c.
too thin; coated; stained
 d.
too thick; shortened; colored
 e.
too short; lengthened; bleached
 c.
too thin; coated; stained
17.
The small circular DNA molecules found in some bacteria are ________.
 a.
inclusion bodies
 b.
plasmids
 c.
nucleoids
 d.
ribosomes
 e.
endospores
 b.
plasmids
18.
Review the Clinical Application, and answer this question.
Pseudomembranous colitis is caused by ________.
 a.
Clostridium difficile
 b.
Streptococcus mutans
 c.
Aquifex
 d.
Bacillus
 e.
Pseudomonas
 a.
Clostridium difficile
19.
In this diagram of bacterial cell differentiation, the indicated structure is a(n) ________.
 a.
flagellum
 b.
capsule
 c.
glycocalyx
 d.
inclusion body
 e.
endospore
 e.
endospore
20.
Bacteria and archaea differ in their ________.
 a.
ribosome size
 b.
need for a cell walls
 c.
type of cell reproduction
 d.
basic 16S rRNA sequence
 e.
chromosome structure
 d.
basic 16S rRNA sequence

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Ch 04: Chapter Opening Video

4.1 The Prokaryote's Place in the Living World

Interactive Graphic: Figure 4.1

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4.2 Bacterial Cell Shapes and Arrangements

such as Streptococcus pneumoniae, are approximately 1 μm in diameter.

Basic shapes Groupings

3D Animation: What do bacteria look like?

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4.3 The Bacterial Cell Wall

b. Crosslinking of glycan chains

b. The gram-negative cell wall

CASESTUDYA Walking Pneumonia Outbreak at a

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4.4 External Structures of Bacterial Cells

Interactive Graphic: Figure 4.6

Animation: Prokaryotic Cell Structure

Interactive Graphic: Table 4.2

The bacterium would run and tumble toward the galactose.

The flagella would spin in the opposite direction.

Ch 04: Microbiologist Toolbox Video

4.5 Internal Structures of Bacterial Cells

Interactive Graphic: Figure 4.9

or attached to the surface.

acquired pathogen, forms endospores. b.

c. Percent failure and/or recurrence of infection

3D Animation: What structures are found inside of bacteria?

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4.6 Prokaryotic Evolution and Classification

b. Taxonomic classification of bacteria based on genetic comparisons of 16S rRNA gene

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 4.2 Bacterial Cell Shapes and Arrangements

wall• Figure 4.5

pili• Figure 4.7

o •Members of the kingdom Bacteria are traditionally classified based on their

The cylandrical shape of some bacteria.

The molecular motor that rotates a flagellum to contribute to bacterial motility.

Movement in response to a chemical stimulus.

A resistant asexual spore that develops inside some bacteria.

(plural flagella) (eukaryotic) An external hairlike structure composed of microtubules in a 9+2

An inclusion body that traps gas and provides buoyancy to cyanobacteria.

The extracellular carbohydrate and glycoprotein coating produced by some bacteria.

A cytoplasmic aggregate that often consists of stored materials.

An inclusion body that allows bacterial swimming to be directed by magnetic field.

A type of symbiosis in which both species benefit.

The enzymatic conversion of atmospheric N2 into organic compounds.

Movement in response to variable light intensity

A group of microbes that share common surface antigens.

The formation of metabolically inactive, physically protected spores.

A group of individuals of a particular species that demonstrate consistent, identifiable variations

A polymer composed of sugar-phosphate subunits that is the major carbohydrate component of a

Print this page

As long as no decay occurred, it would have a commensal relationship.

An environment that is too hypotonic or hypertonic can be lethal to a cell.

Print this page

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